^t^^U Ci —MEMOIR 38

JUNE, 1920

CORNELL UNIVERSITYAGRICULTURAL EXPERIMENT STATIONTHE CRANE-FLIES OF NEW YORKPART II. BIOLOGY AND PHYLOGENYCHARLES PAUL ALEXANDER,,.^Gr^an

l^

SE P

in

£7 1921^

ITHACA, NEW YORK

PUBLISHED BY THE UNIVERSITY

MEMOIR 38

JUNE, 1920

CORNELL UNIVERSITYAGRICULTURAL EXPERIMENT STATIONTHE CRANE-FLIES OF NEW YORKPART II. BIOLOGY AND PHYLOGENYCHARLES PAUL ALEXANDER

ITHACA, NEW YORK

PUBLISHED BY THE UNIVERSITY

Memoir 38

Plate XI

THEODOR BE LING
(1816-1898)

KARL WILHELM THEODOR BELING
No sketch of the life of Theodor Beling, the great German student of theimmature stages of the Tipuloidea, has ever appeared in English. It waswith considerable difficulty that the writer was able to get into communication
with relatives and friends of Beling and obtain the data presented herewith.*
Theodor Beling was born at Steterburg, in the Duchy of Brunswick,

Germany, on March 26, 1816.

He was a son of the district forester. From1828 to 18:34 he attended school at Wolfenbuttel. At Easter, 1834, he began
his chosen career as a forester, having served his prescribed apprenticeshipof two years with his father at Danndorf.
He attended the Royal Saxon
Academy at Tharand from 1836 to 1837, and in the following year visited the
University of Gottingen to round out his technical training in various branches
relating to forestry.
to October

1861, he

1,

1888,

Beling's career as a forester extends from Easter, 1834,

when he was officially retired with a pension.

In April,

was made Master of the Forest, a position which he held until his

retirement.

After his retirement from active public service, Beling long continued his
vigorous researches afield and at home. When far advanced in years, hewent alone on long trips to seek new stations for plants and to observe animal
in wood and field.
He died on December 17,
much of his finest work was accomplished.
life

1898, at Seesen,

whereDuring his lifetime Beling published one hundred and seven articles. The
papers which rank him as a pioneer in the field of crane-flj'
biology were published in the years 1873, 1879, and 1886, respectively. In
series of threethese papers the histories of sixty-nine species of crane-flies are discussed inand brief notes on five additional species are included. The otherdetail,pubhshed

articlesof Beling cover a remarkable range of subjects relating

His most important researcheson the life histories of insects, in addition to those on the Tipuloidea, are on
the coleopterous families Parnidae and Elateridae.
His published articles
are dated between 1850 and 1888, but practically all of his entomologicalstudies were published in the seventies and eighties.
Beling's excellent collection of dried plants, mounted birds, and sectionsof various woods, and his cabinet of insects
the last-named including thetypes of several species described by himself and by othersarc preservedin the collection of the Natural History Museum at Brunswick.
It is understood that the insects in this collection, including the alcohoUc larvae andpupae of the Tipuloidea, are still in excellent condition.
directly or indirectly to the science of forestry.

——* Sincere thanks are here extended to the following persons, who have added materiallyto the value of this biosraphical notice:Landesforstmeister K. Block, son-in-law of Beling. who published in April. 1800 (inAllgemeinen Forst- unci Jagd-Zeitung). a complete obituary notice rclatinp; to his fatherin law. and to. whom the writer is indobtf^rl for several additional unpublished notes andfor the loan of the excellent portrait of Peling reproduced herewith.
.1.
Meerwartz. Director of the Natural History Mu'^eum at Brunswick, who has furnished data concerning the present condition of the collection of the immature stages ofth'' Tipuloidea described by Beling. now preserved in the above-mentioned institution.

William Prindle Alexander and Elsa MuUer Alexander,
service in translating the account of Beling's Ufe.

who have rendered valuable

CONTENTS
PAGE
General considerations
Representative crane-fly life histories
Eriocera longicornis (Walk.)
Cylindrotoma splenderis Doane

704
704
704
708
710
710

Life activities of crane-flies

The adult
The egg
The larva
The pupa'714

Enemies
Predatory natural enemies
Vertebrates
Invertebrates
Parasitic natural enemies
External parasites
Internal parasites
Historical summary of the immature stages

Economic importanceSpecies affecting herbaceous plants
Species affecting woody plants

Methods of experimental procedure
Collecting
Killing and preserving

Study
Rearing
Types of the immature stages
External morphology

The larva
General features
Body form

Organs of respiration

The spiracular disk
Anal gills
The head
The pupa
General features

The head
The thorax
The abdomenInternal morphology
The digestive system
Herbivorous larvae
Carnivorous larvae
Respiration and circulation._

The nervous system
The reproductive system
The muscular system
The fat bodies, or adipose tissue
The imaginal disks, or histoblasts
695_:]

715
719
721
721
721
727
730
730
733
735
736
737
739
740
740
740
741
741
743
744
744
744
745
746
746
748
748
752
752
753
754
756
758
758
758
759
760
762
762
763
763
763

Contents

696

PAGE
Phylogeny

The geological record
Comparative morphology
Phylogenetic considerations
Life histories of crane-flies, with keys and descriptions of the species

Family Tanyderidae
Family Ptychopteridae
Subfamily Ptychopterinae
Subfamily Bittacomorphinae
Family Rhyphidae
Subfamily Trichocerinae
Family Tipulidae
Subfamily Limnobiinae

763
764
766
766
767
769
772
773
779
787
789
701

Tribe Limnobiini
Subtribe Antocharia
Subtribe Ellipteraria
Subtribe Limnobaria
Subtribe Dicranoptycharia
Subtribe Rhamphidaria
Tribe Hexatomini
Subtribe Ularia
Subtribe Epiphragmaria
Subtribe Pseudolimnophilaria
Subtribe Dactylolabaria
Subtribe Limnophilaria
Subtribe Hexatomaria
Subtribe Polymeraria
Tribe Pediciini
Subtribe Adelphomyaria
Subtribe Pedicaria
Tribe Eriopterini
Subtribe Eriopteraria
Subtribe Elephantomyaria
Tribe Styringomyiini
Subfamily Cylindrotominae
Subfamily Tipulinae
Tribe Tipulini
Subtribe Dolichopezaria
Subtribe Ctenophoraria
Subtribe Tipularia
Bibliography and references cited
Index

793
795
799
806
808

828
830
835
838
843
848
852
856
876
893
894
895
899
908
911
952
957
959
974
981
981

.

.

.

".

986
990
1019
1129

THE CRANE-FLIES OF NEW YORKPART n. BIOLOGY AND PHYLOGENY

THE CRANE-FLIES OF NEW YORKPART II. BIOLOGY AND PHYLOGENYi
Charles Paul Alexander

A preliminary classification of the immature stages of the Tipiilidae andrelated families, suggested to the writer by Dr.
ispresented in this memoir.

J.

G.

Needham in 1911,

But few of the sixty-odd famihes of Diptera,and comparatively few species of the Tipulidae, have as yet been studied
from this viewpoint, and therefore the arrangement herein adopted must
be considered as tentative. The majority of the specimens used in the
study were reared in New York State, the crane-fly fauna of which is
In 1913,
typical of a great area thruout northeastern North America.
thru the kindness of Dr. Charles D. Woods and Dr. Edith M. Patch,
the writer was enabled to continue his investigations in Maine.
It is the writer's purpose to outline the morphological characters avail-able for the classificationofthe larvae and the pupae, and to give

preliminary keys for the separation of the various groups; such keys will,accession of new hfe-history material, but

complete remodeUng with theit is at least hoped that they

may furnish a basis for future investigation.

The most important work onof necessity, require constant revision or

the European fauna, that of Theodor Behng (1873 to 1887) ,2 is rendered
incomplete by the total lack of illustration, the insufficiency of descriptionof the details of the larval head, and the artificial nature of the keys.

That this difficulty in using Beling's figureless descriptions is not confinedto the writer is shown by the following criticism by one of his fellowcountrymen (Czizek, 1911:7):" Leider fehlen uns bis jetzt fast vollstandig gute Abbildungen der Larven und Puppen,
ein fiihlbarer Mangel auch in Beling's Abhandlungen, da die genaueste Beschreibimg das
Bild nie ganz zu ersetzen vermag."

It is intended to include in this1.

Descriptions ofallpaper the following material:

new life-history material available, with noteson the biology and occurrence of the species.
'Part I of The Criine-Flies of AVic York, dealing with distribution and taxonomy of the adultpublished in 1919 as Memoir 25 of the Cornell Univers'ty Agricultural Experiment Station.
2 Dates in parenthesis refer to Bibliography and licfircnccs Cited, page 1019.

699

flies,wasCharles Paul Alexander

700
2.

Brief summaries of published life histories of genera and important

species not availal^le for study as specimens and included here to complete

the data.
3. Summaries and tabulations of life-history records, larval habitats,economic importance, and related subjects.
4. Keys to the families, tribes, and lesser groups.
The adult flies are not here considered in any tletail, since they havebeen discussed by the writer in an earlier paper (Alexander, 1919 d).
The life histories remaining to be discovered in the Nearctic fauna
are still numerous in species, tho few in genera.
There are but four orfive genera whose life histories when made known may upset the present
Until more is known of these missing groups,
ideas on arrangement.
they must be classified according to the adult structure.
It will be noted that a number of important changes in nomenclature
have been adopted in this paper. The system hitherto in vogue, based
entirely on the structure of the imagines, was conceived by Osten Sacken
and represented the culmination of research on the structure and affinitiesof the adultflies.

A casual survey of the immature stages is sufficient

to show the impossibility of many of the groups hitherto generally accepted.

The principal modifications adopted in this paper are as follows:
1. Theerection of the family Tanyderidae to receive the genera
Tanyderus and Protoplasa. These had hitherto been placed with the
Ptychopteridae, a group to which they are not closely allied.
2. The removal of the genus Trichocera from the Tipulidae to the
Rhyphidae, and the inclusion of the latter family as one of the four
existing families of crane-flies.
3.

— Antochini, Limno— as being based on a con-

In the TipuUdae, the eUmination of four tribes

philini,

Dolichopezini,and Ctenophorini

glomeration of forms referable to other tribes orinsufficient basis.

else

separated on an

The former tribe Antochini included members which

writer now refers to the Limnobiini(Antocha, Rhamphidia,
Dicranoptycha, and other genera) and to the Eriopterini (Teucholabis,
Elephantomyia) the Limnophilini are too close to the Hexatomini;the;and the tipuline forms constitute a very compact group which cannot
well be subdivided into tribes.
4. The erection of nineteen subtribes, or divisions, to include lesser
groups of genera within the tribes. In the followmg pages these are

The Crane-Flies of New York— Part II

701

treated in what seems to be their phylogenetic sequence from the generalized
to the speciaUzed.

The arrangement of families,
summarized as follows:
Families

subfamilies,

andlessergroupsmay beCharles Paul Alexander

702

A permanent series of the immature stages of the species studied has
been carefully preserved in the writer's collection, wliile additional material
has been placed in the principal collections of the United States.

A seriesof two hundred and seventy microscope sUdes showing details of structureof the larvae and the pupae has been prepared, representing some seventy

speciesin forty genera.
The typical larvae and pupae
and neanotypes) have been designated and preserved inarranged(nepionotypes

alcohol for future reference.

For the most part, the hfe histories discussed
been described hitherto.

in thispaper have not

Besides the reared material, the writer has invery large number of imknown larvae and pupae, many
them representing interesting and undescribed types. In this paper
but three or four of these are considered, and these only because they
introduce new features of organization (Eriopterine No. 1, TipuUni No.
1 and No. 2).
A vast amount of work remains to be done on the immature stagesof crane-flies.
Exact, detailed hfe histories of individual genera andspecies are especially needed and the reward for effort will undoubtedly
be great. Nothing is known of the tropical and the antipodal faunas,
and their life histories will probably be imraveled but slowly, due to theinaccessibility of most of the regions.
The majority of specimens studied in the preparation of this paper
were reared by the writer during the past nine years. Many specimens
were received from various sources, however, including many of the mostdesirable hfe histories, and these are herewith gratefully acknowledged:
Dr. James G. Needham, Ithaca, New York. The extensive collections received from Dr.
his collection aofNeedham include the material described in the various Adirondack reports, as well as
numerous previously unrecorded specimens. In r.ddition, the writer has accompanied
Dr. Needham on many of his field trips in quest of the immature stages of aquatic insects,
and has received much encouragement and inspiration from this association.
Charles W. Johnson, Boston, Massachusetts.
Immature stages of Aeshnasoma, Elephantomyia, Teucholabis, and other species.
The late Frederick Knab, Washington, D. C. Immature stages of Rhipidia bryanti,
Elephantomyia, Epiphragma solatrix, and other species.
Longurio (larvae), Oropeza, Gnophomyia.
J. A. Hyslop, Hagerstown, Maryland.
Rhipidia bryanli.
J. R. Malloch, Urbana, Illinois.
Protoplasa (supposition).
C. T. Greene, Washington, D. C.
H. L. Viereck, Washington, D. C. Protoplasa (supposition").
W. L. McAtee, Washington, D. C. Protoplasa (supposition) numerous larvae from the;

Pribilof Islands.

William Lundbeck, Copenhagen, Denmark. Helobia and other species.
Dr. C. Weseaberg-Lund, Hillerod, Denmark. Phalacrocera, Ptychoptera paludosa, Tipula

maxima, and other species.

The Cra.ne-Flies of New York— PartII

703

William G. Dietz, Hazleton, Pennsylvania. Notes on larvae of Dicranomyia macateei Alex.
Oskar A. Johannsen, Ithaca, New York. Hmnobia fallax, and other species.
Notes on Rhipidia fidelis.
Carl Ilg, Philadelphia, Pennsylvania.
Notes on Libnotes and Styringomyia.
O. H. Swezey, Honolulu, Hawaii.
H. K. Munro, Pretoria, Transvaal. Notes on Trentepohlia and Styringomyia.
Charles Bruch, La Plata, Argentina.
Notes on Triraicra, Helobia, and Epiphragma.
D. B. Young, Albany, New York. Notes on Gnophomyia.
.'. T. Lloyd, Cincinnati, Ohio.
Eriocera.
The late C. Gordon Hewitt, Ottawa, Canada. The Canadian-Arctic Tipuiidae, including
Tipula arctica.
Cylindrotoma splendens.
A. E. Cameron, Ottawa, Canada.
R. C. Shannon, Ithaca, New York. Elephantomyia, Brachypremna, Tipula fuliginosa,

and other species.
Speed Rogers, Grinnell, Iowa. Notes on Phalacrocera tipulina.
Harold Morrison, Washington, D. C. Holorusia, and Tipula usitata.
C. Hamilton Kennedy, Columbus, Ohio. Prionocera and Rhamphidia.
J.

J. Chester Bradley, Ithaca,

New York.

Longurio.

H. S. Barber, Washington, D. C. Oropeza.
Adam Boving, Washington, D. C. Tanyptera; notes on Helobia.

In addition to the preceding, the writer acknowledges specimens ordata received from the following:
From Orono, Maine: Edith M. Patch, H. M. Parshley, Cordelia Stanwood.
From Cambridge, Massachusetts: Roland Thaxter.
From It-haca, New York: H. H. Knight, L. G. Brown, C. C. Hamilton, V. R. Haber, E.
A. Richmond, Chih Ping, Waro Nakahara, M. D. Leonard, W. A. Riley, J. D. Tothill,
Mrs. J. D. Tothill, W. P. Alexander, W. A. Clemens, W. C. Woods, P. A. Munz, P. W.
Claassen, Axel Olsson.

From Washington, D. C: C. H. Popenoe, E. R. Kalmbach, Alex. Wetmore, A. T. Speare.
From Lawrence, Kansas: H. B. Hungerford, P. B. Lawson, H. L. Fackler.
From Urbana, Illinois: T. H. Prison, W. P. Flint.

The writer would express his great indebtedness to Dr. Ewald Bergroth,of Jamsa, Finland, for help in determining the derivation of some of themore obscure generic namesinthe Tipuhdae; to Professor T.

D. A.

notes on the stratigraphy of fossil-bearmg beds in manyparts of the world; and to Miss Lela G. Gross, assistant editor of theCornell memoirs, whose critical work on this and the preceding part

Cockerellforunder this title has been of the greatest value in assuring uniformity
and accuracy.
Further acknowledgments of assistance in determining plant andanimal associates of crane-flies are made thruout the text. In conclusion,
the writer expresses his indebtedness to his wife, ]\Iabel ]\I. Alexander,
for her untiring care and assistance in the typing and final preparationof this thesis.

Charles Paul Alexander

704

GENERAL CONSIDERATIONSREPRESENTATIVE CR.\NE-FLY LIFE HISTORIESThe life histories of but few species of crane-flies have been studied
in detail.
The ^•erv nature of the habitat (mud or earth) required bymost species of the family renders it a most difficult operation to rear
the species from the egg to the adult and note the various stages, theirmolts, their halnts, and other features.
The subfamily Cylindrotominae,
the immature stages of which live on the leaves of various liigher plants
and curiously resemble the caterpillars of certain Lepidoptera, furnishes
species whose habits are more readily studied than most others, and as
a result the immature stages of tliis group are better known, perhaps,
than those of any other section of the family. Two widely different
species have been chosen, and their life histories as they are known atpresent are here outlined.

The first is a species of Eriocera, a powerful,semi-aquatic carnivore; the second
restrial herbivore.

isa species of CyUndrotoma, a ter-In the text which follows, notes on the hfe activities

of various other species are given, but the gaps in theknowledge of thisphase of the subject are verj- considerable and there still remain innmnerable interesting facts to be ascertained.

Eriocera loiigicornis (Walk.)

The common crane-fly Eriocera longicornis is widely distributed overthe northeastern United States and Canada.

Altho the species is local

in its distribution, the fhes may be found in

abmidance wherever it does

The situations that favor the presence of these fhes are large
streams or rivers with sand or gravel bottoms and banks. The following
notes were taken, partly in Fulton County, New York, along the Sacanoccur.

daga River, and partly in Tompkins Coimty, along Fall and Cascadilla
Creeks.
Some of these data have already been published by Alexander
and Lloyd (1914:12-18) and by the writer (Alexander, 1915 c: 14^^-152).
On ]May 27, 1914, the adult flies were exceedingly numerous near thevillage of Northampton, Fulton County.
They were present in untold
myriads, and at every step they arose in clouds from mider foot or fromthe leaves of chokecherr\' on which they rested.
They sat on the leaves
with the head directed away from the observer, ready to take instant
flight, and at the first approach of a possible enemy they darted up into

The Crane-Flies of New York— Part II

705

the air and far overhead.
A few were in copulation on the leaves, butfrom observations made later it seems probable that mating begins in

the air an.d the united pair seek a support later on.

Toward twilightthe flies may be found in great numbers in company with many kinds of

caddice
fifty

flies

and may flies.

The swarms vary in munbers from about
Some of the larger

individuals to those including many thousands.

of theseswarms cover a vertical height of at least fifty feet, the lower

individuals

being about fifteen feet above the water.

Whendangerswarm either mounts into the air overhead or retreats
before the breeze, never advancing nor moving sidewise.
Out over the land in the smaller swarms, copulation was observed
several times.
The males in the swarm dart swiftly at the females andseize them almost instantly.
They then usually leave the swarm and go
approaches, thesailing away, the male above doing the flying, the smaller female hanging

The antennae of the male at this time are directedahead and are slightly divergent, the legs hang downward, thewings vibrate rapidly. The female hangs downward with the legs trailing
limply beneath, the wings nearly horizontal and motionless. The deadweight of the female continually pulls the male toward the water, andoften both fall into the river.
As a rule, copulation ends before the maleis exhausted.
The male opens his forceps and the female drops straight
downward for a foot or so, quite like a parachute released from a balloon.
limply beneath.
straight

If

the pair are near the surface at the time of separation, the released

The male darts upward again and backswarm. The female slowly flies away, usually upstream but
sometimes downstream, presumably to lay her eggs. She does not stop
for swarms that she may encounter, and may even make a wide detourin order to avoid them.
In a very few cases the female is the active
partner and succeeds in pidling the male where she wills altho she is muchsmaller.
No matings were observed during the hours of sunlight, andit was only at twilight or just after sunset that mating took place.
Often
a second male will seize a female already in copula and the three will
come tumbling down into the water together. One such pairing of thi-ee
individuals came down, and when they were about a foot above a board
in the water the male in copulation dropped the other two.
These rested
for a moment, and then the remaining male attempted to engage thefemale in copulation. She resisted but finally he managed to seize her
female drops into the water.
into theCharles Paul Alexander706

with his forceps. Thou he attempted to fly away, but she seized holdof the board with all her feet and he was unable to disengage her.
This
seems to indicate that the normal place for copulation is in the air.
The motions of the insects in the swarm were very rapid, almost Hke
those of bees, and the sound produced was at a very low pitch, much
lower than thatplane,

made by Culex.

The movements are on a horizontal

each individual flying mostly in the path of a figure 8, sometimes

slowly and at other times much more rapidly.

Many specimens were seen dipping down into the water, as tho engaged
All of the few specimens captured proved to be males,
but why this sex should go thru these motions is not clear to the writer.
This action has been observed several times in various species of craneflies.
It is very probable that the female lays her eggs in the water inthis manner.
The eggs are pale white or brown, not heavily chitinized as are thoseof Hexatoma but with the chorion feebly sculptured.
They vary innumber from 892 to 1034, with an average of 952. They are small, about
the same size as those of Hexatoma. The ovaries almost completely
fill the abdominal cavity, and the eggs are arranged in the ovaries likebananas on a stalk, with numerous pale nurse-cells in between.
The larval life is passed in streams, usually under rocks. The winteris spent in the larval condition, but the larvae do not attain full size until
the following spring. At tliis time they come to the land and live inthe sand and gravel along the banks of the streams. By the alternate
extension and contraction of the body and the inflation of the penultimate
segment of the abdomen at the moment of extension, the larvae arecapable of inflating tliis segment into an enormous globular structure
which serves as an aid to progression thru the soil. The food of the larvae
consists largely of animal matter, and often large species, such as chironomid larvae, are swallowed whole. The almost total lack of chitinizationof the mental region allows for great distension of this part of the body.
The powerful mandibles and the retrorsely roughened esophagus serve
the fimction of both holding the prey and preventing its ejection whenonce swallowed. Consideral)le gravel and particles of vegetable tissue
are also found in the proven tricular region.
When ready to transform to the pupal condition, the larva becomes
After molting the last larval skin, the pupa is disclosed, palesluggish.
in laying eggs.

The Crane-Flies of New York— PartII

707

The pupa forms burrows in the loose
The diaineter of
The length varies, thethe burrow is a little greater than that of the pupa.
yellowish whiteand very callow.

gravel, these being vertical, or, more often, a little oblique.

burrow for a young pupa being a mere chamber inclosing the individual
As the pupa
and located from one-half to one inch below the surface.
becomes older, by a bobbing up-and-down motion it lengthens the burrow
upward until finally it penetrates the surface layer and forms a small
Here the pupa rests, often bobbing up and down
shot-hke opening.
with a rhythmic motion, but ducking down into the burrow when danger
threatens.
The length of the burrow is rarely more, but usually less,than twice the length of the pupa. A few of the pupae seem to be
inclosed in a very deHcate silken tul^e and the saUvary glands seem tobe well adapted for the purpose of forming silk; but the great majority
The indoor pupal period varies fromof the pupae are entirely naked.
one hundred and seventy to one hundred and seventy-three hours, or alittle more than seven days.
The outdoor pupal period, however, isundoubtedly longer, unless the weather is very warm.
The emergence of the adults usually takes place during the late hoursof the morning, the greatest number emerging between ten o'clock andnoon.
When ready to emerge, the pupa pushes part of its body out of theearth, the posterior two- thirds or half remaining attached to the soil.
If it projects farther than this, its transformation seems to be a very
difficult operation.
The pupa bends backward and forward constantly,
This motion appears to exhaust it,
flexing the body dorso-ventrally.

since

it

frequently rests.

The skin splits lengthwise up the mesonotum

and the adult emerges. The male has difficulty in extricating its very
long antennae from their sheaths.
The tips of the fore femora are placed
underneath the sharp spines of the flagellmn, and by raising the legs the
These spines are regularly
insect pulls the antennae slightly outward.
spaced, and, since both fore legs work in unison, the spines function as cogs
and the whole antenna is gradually forced from the pupal sheath. The bodyis ca-rried very straight and stiff during the operation, and the abdomen
is very long and pale.
The drawing out of the extreme tips of the antennae
is usually accomplished l)y the insect flexing its whole body backward.
When the antennae are freed, the insect walks a few steps from the cast skin,withdrawing its abdomen from the case. (The emerge-nce of E. spinosais shown in Plate XII, 1.)
A drop of nearly colorless Hquid is excreted

708Charles Paul Alexander

from the body at this time. The teucral
achilt then waits quietly until
itgams more strength and color. This condition of
the insect is the mostdangerous period of its existence, since it is defenselessagainst all enemies

The insects are associated in the gravel withground beetles
genera Omophron, Sclaizogenius, Dyscliirius,
Bembidion,of theTachistodeswith click beetles of the genus Cryptohypnus;
and with rove beetles ofwhich Paederus, Lathrobium, and Gastrolobium
are the commonest forms
In addition, numbers of larvae of Tabanidae,
Leptidae (Atherix), Eriocera
spinosa E. fultonensis, E. dnerea, andEriovtera armata were foundNatural enemies of the pupae and the teneralimagines are the mediumsized black lycosid spiders, which
preyed in numbers on the weakuncolored adults. Dozens of these spiders
were noticed with individualsof the crane-flies in their grasp.
When alarmed they would run rapidly
away, but only in exceptional cases would
they release their victims
A
few spiders of other famihes, notably the
Attidae, were found with Eriocerae.
Dragon-flies appear to be the most serious
enemies of the active
adults. Helocordulia uhleri (Selys)
has been observed capturing the craneflies by dartmg back andforth thru the swarms of individuals.

Cylindrotoma splendens Doane

The hfe liistory of the species Cylmdrotoma
splendens has been
worked out m considerable detail by Dr. A.
E. Cameron, from material
obtamed near Westhohne, Vancouver Island,
British Columbia, in latea^«o^"^<^ is abstracted fromf^^^«^"^gpaper nlr,o^(1918) on tins interesting crane-fly:

The adults first appear on the ^ving about

the middle of May.

Cameron's detailed
Without food thpv ^^wmMmmmmmr™u«^,„„* NutJ, (Ra„u„c„Vcter,tSt;e:'thto'Sdtpltj:"^ t'The Crane-Flies of New York— Part709

II

Palmatelyjust within the margin of the serrate,
are usually deposited in series along and^or more, all arranged
oneofgroups
found
are
They
fobed leaTCe^f.
e^sso, to the margm of the leaf. .The
and wth their long axes perpendicular, or nearly
being exposed dorsally, he margms of theepidermis,
thebeneath
Mdden
par y
a?e
posteriorly.
to a lesser degree, both anteriorly andsht overlapping the egg laterally and,leaf margin a somewhat beaded appearance
the
to
gives
series
in
eggs
the
oi
Ais arraSemeut
brown and then the Sgs are older, the leaf margin turns
under side of the leaf, %vith the tip ol the
In ovipositing, the female rests on thedetected
theis slightly flexed ventra ly, andabdomen
The
edge.
leaf
the
towardabdomen directedvalves of the ovipositor which is app bed
bifurcatedthebetweenheldcaf
Tsmarg^ of thewith their blades, are appbed
rthruoper surface of the leaf, while the paired cutting valves,and fro, cuttmg a sht in the.are then moved tobladesThese
surface.
Lhistthrunder
actually touch one another
The eggs are then deposited. In no case do the eggs
upper
their eggs indiscriminately on both thelayfemales
cages
breedingtheInearher incases, when a leaf has been eatensomeInlefves.
theofsurfaceand the lower
damageddeposit her eggs along the ragged edge of thisthe season by larvae, the female wiUm

Pf "^^^roXWh

YT^T/ '^7S SnlfZ

Sermi
onSeafto eighteen days, but in the field^"^The indoor duration of the egg stage is from fourteen
first deposited the egg is translucen
Whenrequired.
be
m'fy
weeks
t£ee
asTong a periofas
unornamented, and measures on anand spindle-shaped, with the chorion

gLyS whke

The chorion of the egg

the'ThTl'rvf r'e'qu'irefab'oit'S hours to emerge fromextendmg
almost half the length ofegg.

mid-dorsal line, the slit
splits longitudinally down thewhite If ^''-^^/ibzes the body ubercles
In emergLg, the almost transparent, grayish
as it is freed from the shell, the ne.lysoonAseggshell.
thefromitself
ill freemi
Isof the host plant.
hatched larva begins to feed on the leaf tissue
i iq v,,.
bycolor, are translucent, and measure 1.19
The first-stage larvae are grayish white
the lower surface of the leaf,
and
upper
the
both
on
feed
They
37 mnhmetef in sTze
and feeding on the parenchymatousburrow ni thru the epidermal layer with their mandibles
are not readily disturbed whenandsluggish
very
are
larvL
young
The
While engaged in feedingleaf tissue.
theinembeddedfirmlybeing
feeding the mandiblesof nme days the larvae ha e
end
the
At
surface.
leaf
the
on
positionsthey assume various
characteristic behavior of the fuUlength to 5.S4 millimeters, and show all theincreasedcoveredfirst -olt the larvae becomethebefore
and
slow
very
is
S^Tntrv'ai.
ihe first larval moit
the skin,toreadilyadhereswhich
excrement,with particles of their
twenty-one days; in some cases, however, it does
occurs after a period of from eighteen toelapsed.
have
weeks^not occur imtil five or six
racolor as they continue to feed
The sec^d-stage larvae gradually assume a leaf-green
their food plant, the larvae, whichofdying-offthewithcoincident
Toward the end of July,
millimeters, become quiescent and cease to
haTe now reached a length of from 8.32 to 9completely and the larvae remam clinging
cease
gradually
feed Teding and movement
some cases,
the larvae drop off, and,
wither,leavestheAsmotionleS to the leaves.
usually fall among
they
however,
conditions,naturalunder
Attach themselves to the stems;
condit on.
these they pass the winter in a dormant
the dead leaves on the ground and underleaves, but some of them retain
dead
of
the
color
brown
the
match

?he egg

Ws

m

t™Sii

r Xowthj.

mMany of these larvae

March of the ff o^ing sprmg'^Tll'fySlZifr'^g larvae first show signs of activity in
Growth then P^.^l' .^^^
when Trautvetteria grandis sends up its new shoots.
^/"^^^
be one molt before hibernation a n^^^pupation in the middle of May. There seems to
fullj gro^sn
Thepupation.
topreviousskin,larvaltheofcasting

the last being theinvariably found on the upper surface of he leaf,
farVae measure 17 milhmeters. They areleaf the larvae usually begm
anITnTheTpringaTe actively engaged in feeding. On a fresh
Later on, however, large holes may
bv skeletonizTng it, leaving the lower epidermis intact.
grown larvae, and to a somewhat lesser fgree
be eaten completely thru fhe leaf. The fully
motion, which may be aptly compared to thatthe yo^ger larvae, progress by a looping
afterCharles Paul Alexander

710

of the measuring worms (Geometridae) , progression being accomplished by the aid of themandibles and the ventral tubercles, or pseudopodia. In a quiescent condition the thoracic
region of tlio body has a noticeable humped appearance.
The fully grown larvae are verysluggish and inactive.
When disturbed they relax their hold on the leaf surface and fallto the ground.
Altho several hundred adults were reared, not a single parasite was discovered.
Before pupation the larva attaches itself firmly to the surface of the leaf or to the leaf
petiole, by means of its anal pseudopodia.
The skin splits transversely behind but is only
partially sloughed off.
The head, the thorax, and the first four abdominal segments of thepupa are exposed, but the apex of the abdomen remains encased in the larval skin, tl>eterminal part of which, collapsed and wrinkled, is attached to the leaf surface. Pupation
may take place on either the upper or the lower surface, but it occurs oftener on the former.
If pupation takes place on the petiole, it is generally at the axil.
In the breeding cages theduration of the pupal period was found to vary from six to ten days.
When the adult first emerges it is of a pale green color, which is gradually replaced by theblack and yellow of the fully colored insect. From one and one-half to two hours are required
for the adult to emerge.
After emergence is completed, the fly rests for a short period
until the cuticle hardens and the wings expand.
It seems that the adherence of the larvalskin to the pupa is necessary for the emergence of the adult, at least in many cases.
There
appears to be a large disproportion of females over males, this sometimes being as high asfive to one.
Since a single male may copulate with several females, this disproportion ia
not so serious as it appears at first sight.

LIFE ACTIVITIES OF CRANE-FLIESThe adult
Emergence.

— Emergence from the pupalhull

may require but a few

seconds (as described for GnophomyiabyHyslop in Zz7i., the whole operation
requiring but eight seconds), or it may take several hours.

The emergenceof the strictly aquatic genus Antocha has not been observed, but it must bepractically instantaneous as in Blepharocera and the lotic caddiceflies.

— In several widely different species, the females as they emerge

Mating.-from the pupal hulls are at once seized in copulation by the males altho
they are still callow and uncolored. Mik (1882 b: 40, and 1886 a) discusses
this curious condition in considerable detail.

known

to the writer— Dicranomyia

irinotata,

In

all

the cases that are

Discobola caesarea,

andCylindrotoma distinctissima (Mik, 1886 a, the last-named also cited by Mik,

1882 b), Liogma glabrata (Miiggenberg, 1901), Tipida rufina (Giard, 1895),
and Tipula ultima (Caudell, 1913)
the abdomen of the female iselongate, flabby, and nearly colorless.
In Dicranomyia trinotata thefemales scarcely have time to remove their legs from the pupal sheaths
before they are seized in copulation.
In other cases the males emerge
before the females and wait beside the pupae for the emergence of
their mates, when they at once seize them in copula.
In most speciesof Tipula the males, when seeking the females, progress by a fluttering—

The Crane-Flies of New York— Part II

711

motion, partly flying and partly walking, over the ground or up the trunks
Tliis habit is discussed under the account of Tipula taughannockof trees.
(p. 1013),

this

genus

and has been observed in other woodland-inhabiting species ofIn
as T. Jiiacrolabis, T. fuliginosa, T. fragilis, and others.

—

when a male comes upon a pair already in copula he passes
on without interrupting them. Somewhat similar mating habits are foundin some species of Dicranomyia {D. trinotata, D. badia, and D. simulans),
Discobola, Antocha, Chionea, Dadylolabis montana, some Pediciini as
T. fragilis,

Dicranota, and the Cylindrotominae.

Many crane-flies have developed swarming habits for the purpose of
these including representatives of most of the tribes of the
Limnobiinae and a few tipuline forms. Dicranomyia morioides wasobserved by Needham (1908 a: 204) swarming in vast nimibers near
LikeIthaca, New York, but here the swarms consisted only of males.
wise, Erioptera armata (Needham, 1908 a: 206) was found swarming nearmating,Lake Forest, Illinois; but, out of several hmidred individuals captured,
The writer has observed swarming in numerous species of Ormosia, Molophilus, Erioptera, Gonomyia, Rhabdomastix,
Limnophila, Ula, Epiphragma, Eriocera, Dicranota, Rhaphidolabis,
Trichocera, and other genera, and here, too, the males were always
predominant. The specific data may be consulted under these various
The males of Dicranota swarm in rather large numbers preheadings.
liminary to searching for the females, which rest quietly on the branches
all except three were males.

of neighboring shrubbery.

Limnophila ultima, as noted at Gloversville,

New York, on September 7, 1916, was swarming at half past six o'clock
The swarms here consisted of from fifty to sixty individuals
in the evening.
and took place from ten to eighteen feet above the earth. Mating took
place frequently in the air, and as soon as a pair were in copula they flew
away to some point to rest, many pairs being observed hanging on a
There were three distinct swarms, whichclothesline a few feet away.
showed little tendency to fuse altho their flight area was very close. The
vast swarms of Eriocera longicornis and of Trichocera are mentioned or
discussed elsewhere in this paper.

When pairs are in copula, they readily

imited, the female usually trailing the male after her;altho in a few groups, in which the male is the larger individual of the"Brachypremna, the famihar " weaver
two, the situation is the opposite.
of the Southern States, has a very remarkable vertical dance of several

takeflight,still

Charles Paul Alexander712

spots, and has licen aptly termed by Johnson (1907-12"
king of the tlancing tipuhds." Doubtless many interestthe[1909] 123)

feet inshady:

ing facts remain to be discovered concerning the dances of the tropical
species of Brachypremna,

Tanypremna, and Megistocera.

Dancing.'— The dances of Thrypticomyia

saltens(Dol.)

According to Doleschallshouldbe
Jacobsonmentioned at this(1857),(De Meijcre, 1911:22-23), and others, tliis species is common in JavaThe insects have the habit of clinging
in shady places thruout the year.
fore
of
the
feet, or, if a fore foot is lacking, one
to spider webs by means
Often
twenty or more of these flies are seenof the middle legs is used.
anotherona horizontally spun web, all seesawing
hanging close beside one
sametime swaying to and fro, sometimes
rapidly up and down and at theThis ludicrous tight-rope dance is conrhythmically, sometimes not.
tinued for a long time. Somewhat similar habits have been describedfor Trentepohlia pennipes, Rhamphidia venustissima Alex., and other species
point.

with white tarsi.
Bobbing.

— SpeciesofDicranomyia and Geranomyia, as well as a few

down while
The species of the latterother crane-flies, have the curious habit of bobbing up andresting, the long, slender legs acting as springs.

this same oscillating movement while resting on a head
and feeding. H. K. Munro has recorded tliis bobbiiig habit

genus practiceof flowers

in a species of Trentepohlia (p. 943).

— When at

rest crane-flies assume various positions
which are often fairly characteristic. Many species (Erioptera, Molophilus, and some Dicranomyia) resemble spiders when flattened against
a tree trunk or some other vertical support. Styringomyia resembles
a bit of cobwel), the fore and middle legs stretched out in front, the hindIn a position of rest the tipuhne forms generally
legs directed backward.

Resting

habits.

hold their wuigs outspread or divaricate, exceptions being in the genera
Longurio, Oropeza, and others, and in a few species of Tipula, as T. ardica

and the woodland-inhabiting species of the marmorata group {T. Jragilis
and T. ignobilis). The limnol)iine forms usually fold the wings incumbent
over the abdomen, but here again exceptions are found in Pedicia,
Limnophila toxoneura, and other species, which normally rest with thewings outspread. Oropeza hangs to the roofs of bridges, culverts, andsimilar places, with only the fore legs attached to the support, the middle
legs divergent, the posterior legs hanging loosely behind.

The apparently

The Crane-Flies of New York— Part713

IIclosely allied Dolichopeza, on the contrary, has the four anterior legson the support, the hind legs dangling, the wings divaricate. Many ofas Oropeza, Dolichopeza, Dicranomyia badia, and exotic
these species
habitually rest
species of the genera Thrypticomyia and Trentepohlia
on spider webs (page 982; also, Knab, 1912). During heavy rains, craneflies rest on the lower, side of the broad leaves of deciduous trees or hide——

beneath loose flakes of bark.

—

Feeding.
The acUilt flies feed but little, the majority of the species
whose habits are at all known merely lapping nectar from open flowers.
Knuth (1909:579) and Wahlgren (1917) record the plants frequented bya number of European Tipulinae as well as by Ptychoptera. These species
are found on Umbelliferae (Aegopodiimi, Anthriscus, Heracleum, Carum,
Anethum, and other genera), on Rosaceae (Spiraea, Rubus, and otherA few of the local Limnobiinae
genera), and on a few other plants.
have the rostrum very greatly elongatedan obvious adaptation forsucking the nectar from tubular flowers. The species of Geranomyia
feedon various composite, umbelliferous, and lauraceous flowers.
Toxorhina frequents composite, rhamnaceous, apocynaceous, and ericaceous plants. The exact plants frequented are discussed under the—descriptions of the respective genera.
Ovipusitioji.-thelarvae.

Eriocera,— The females lay their eggs in the habitat frequented by
In

the

case

and other genera

the water surface, one oraquatic forms — as Antocha, Hexatoma,— the eggs are deposited dipping down toofinmore eggs being deposited

at each descent.

Forms that live in mud or moist earth lay their eggs in these situations.

Many species with acute ovipositors insert the eggs carefully into the soil
or other substance.

The oviposition of Lininophila {Eutonia) alleni as

noted by the writer may be regarded as typical of this class of species:
Observations made at Gloversville, New York, June 28, 1916. A female was noted
ovipositing in low, wet spots along a small woodland stream.
She flew about slowly andsilently, just skimming the ground, until a place suitable for egg-laying was found.
She
finally chose a much-decayed log and the eggs were driven home securely by the aciculartergal valves of the ovipositor.
Much effort is expended to place the eggs firmly and therate of oviposition is not more than eight or ten a minute, the female often pausing to rest
for several seconds.
While thus engaged, the fly is entirely unconcerned with other agencies

and may be picked up by hand.

The females of most crane-flies usually live but a short time after egglayingiscrane-flies

completed.
isIndeed, the entire duration ofprobably but a few weeks at the most.

life

of

the adult

714Charles Paul Alexander

The specialized methods of oviposition in the Cylindrotominae havealready been discussed on page 709 under the account of Cylindroioma

The species of Tipula and other genera the females of whichhave blunt valves to the ovipositor (Styringomyia, Macromastix, andothers), all probably scatter their eggs promiscuously or else have a specialsplendens.

ized method of egg-laying.
Similarly, the species of Tipula of the arctica
group, in which the dorsal valves of the ovipositor are very large, placed

horizontally,

and with the margin

finely serrated,undoubtedly have apeculiar method of oviposition, but this has not yet been observed.

The egg

The number of eggs laid by crane-flies varies from about forty-five in
Styringomyia to some two thousand in the larger species of Eriocera.
The following table indicates the general range in the group:
Number of eggs laid
Species

Reference

Maximum Minimum
Ptychoptera albimana
Styringomyia didyma
Eriocera longicornis
spinosa

Hexntoma megacera

.

.

.

.

.

Phalacrocera replicata

Liogma glabrataCtenophora angustipennis
Tipula cunctans
bicornistephrocephala ....
angustipennis ....
collaris

dejectaoleracea

587
45
1,034
2,061
372
60

60
400
297
255
eo2
329
366
600

Average

520
872
1,824
316

200
282

251

554
45
952
1,942
347
60

60
300
300
289
255
602
329
309
600

Topsent, 1914-16
Terry ms.
Alexander ms.
Alexander ms.
Alexander, 1915 c
Miall and Shelford,
1897
Miiggenberg, 1901
Lovett, 1915
Hvslop, 1910
Webster, 1893 b
Webster, 1893 b

Hyslop, 1910
Alexander ms.
Alexander ms.
Del Guercio, 1914

The eggs are in most cases elongate with the ends narrowed and rounded,other words, spindle-shaped.
The chorion in the species of Tipula,Hexatoma, and other genera is blackened and in some cases more or lessor, insculptured.
In Ctenophora angustipennis (Lovett, 1915) the egg is ebony
black with deep purple reflection, elongate-oval, uniform, without pits orridges,and measuring from 1.26 to

1.4 millimetersby 0.375 millimeter.

The Crane-Flies of New York— Part II715

In Tipula bicornis (Webster, 1893 b) the egg is 0.8 millimeter long by 0.3
to 0.4 millimeter in diameter, elon.gate-ovoid, with one side deeply concave,the surface highly polished. The egg of Nephrotoma ferruginea is smaller,
with five distinct grooves. The egg of Tipula gladalis (Pokorny, 1887:53)
is described

as being 1.2 millimeters long and only 0.3 millimeter in diame-ter, cylindrical, thetwo ends equally rounded, the surface smooth, shiny

black, with faint steel-blue or purplish red reflections.

generaIn

many otherof Tipulidae, especially the smaller forms, the eggs are soft

whitish or nearly hyaline.

andIn a few species the eggs take on a decided
The egg of Cylindrotoma splendens is describedgreen or greenish tinge.
elsewhere in this paper (page 709). The egg of Ptychoptera albimana, as
described by Topsent (1914-16), measures 0.825 by 0.284 millimeter,
and is pale yellow, slightly arcuated, and with the surface curiously

ornamented.

The larva

As a rule, the larvae live in the hamits where the eggs are laid.

The

duration of the larval stage varies from about a month in Styringomyia
Terry ms.) to the greater(larva and pupa together, thirty-seven days—

part of a year in most crane-flies.

Many of the smaller species of Eri-

optera, Ormosia, Rhaphidolabis, and other genera are on the wing in thespring and again in the fall, and with little doubt are double-brooded.

This would make the larval existence but a few months, but still probably
This problem of double
longer than the other stages taken together.

broods should be worked out carefully.

It oftenappears that there are

two broods, when in reality there may be two developing generations,each passing the winter as larvae, but one attaining its growth much more
slowly in the spring and summer and not maturing until late summer.
Phalacrocera and Cylindrotoma splendens spend about eleven months in(Bengtsson 1897, and Cameron 1918), while Tipula
paludosa spends nine months in that stage (Rennie, 1917). It is probablethat nearly all crane-flies in the North Temperate Zone winter normally
The growth during summer, fall, and winter is very slight, but
as larvae.

the larval stage

in the springitisgreatly accelerated and in amonth the larva may

Larvae of Tipula ignohilis taken at Ithaca, New
York, on April 23, 1917, measured only 6.5 millimeters in length; on
May 19 they had attained their full growth of 18 millimeters and wereattainits full

growth.

ready to pupate.

Cylindrotoma splendens spends the winter as a larva,

Charles Paul Alexander

716

9 millimeters in length, but grows rapidly during the spring, attaining
its full size

(17 millimeters) in

two months of growth.

Liogma glabrataspends the winter as a very small larva, but in the spring its growth isgreatly accelerated.

The haunts in which the larvae of crane-flies occur are exceedingly
varied.

In the case of single large genera, such as Dicranomyia andTipula, the species range from those that are almost strictly aquatic toothers that are entirely terrestrial, living in decaying wood or even mining
in the leaves of plants.

The transition between strictly aquatic and terrestrial forms is very
gradual, as was pointed out by Miall (1895:11) some years ago when
he wrote:

How did insects ever come to seek the water, seeing that their mode of respiration is
primarily adapted to another element? We can see almost all the steps of the adaptationon the shores of our rivers, lakes and seas. We can see dipterous larvae which, like the" leather jacket " (the larva of the daddy-long-legs), burrow in the ground for their vegetable
food, and devour the roots of grasses.
Other larvae of the same family (Tipulidae) prefermoist earth in the neighborhood of streams. Others again live immersed in water, or mud
saturated with water, though they come to the surface at times and push their tails, whichcarry the spiracles, into the air. Some few have become so completely aquatic that they
seldom, if ever, come to the surface, and all their supply of oxygen is obtained from thewater.

The culmination of this latter condition is reached in forms such as
species.
Crampton (1919:100) has
made similar observations on the subject.
The haunts of the larvae of crane-flies are best shown by the following
Antocha and related genera andtable,

infrom thewhich the various species are arranged according to habitat,strictly aquatic to the various terrestrial

Species

Habitat
Strictly aquatic,in silken cases

forms:

AntochaIn very rapid water (lotic) on or in submerged mosses (hygropctric association)

Dicranomyia simulant, Pedicia,
Tipuline No. 1, and others

Aquatic, on submerged plants

Phalacrocera, Triogma

Semi-aquatic (part of life spent in water,
but pupation taking place on land)

Dicranomyia simulans, Eriocera, Hexatoma,
Aeshnasoma, Longurio, Tipula ahdomi-

On cliffs and wooden walls, usually in silken

Dicranomyia simulans, Geranomyia,
tera, Dactylolabis, and others

nalis,

cases covered by water

In cold springs

T. caloptera, T. bella,

Pedicia, Thaumastoptera

Triogma,and others
Ellip-

The Crane-Flies of New York— Part717

IISpecies

HabitatIn stagnant water in the axils of bromeliaceous and other plants

Trentepohlia, Gnophomyia rufa

Amphibious in decaying wood

Protoplasa (supposition), Epiphragma

In mud or sanda. In open swampsb.

Ptychoptera, Rhamphidia,
Erioplera septemtrionis, E. chlorophylla, E.
vespertina, Limnophila adusta, L. macrocera,
Pilaria recondila, P. tenuipes, P. quadrata,
Tricyphona inconstans, Prionocera, Tipula
dejecta, T. sayi, T. tricolor, and others.Bittacomorphella, Ormosia innocens, Eriop-

Bittacomorpha,

In shaded woods

tera

c.

Along the margins of streams andother bodies of water

megophthalma, Molophilus hirtipennis,

Ulomorpha, Dicranophragma, Penthoptera,
Tipula cayuga, and others
Erioptera(Hoplolabis)
Limnobia
fallax,armata, Trimicra, Gonomyia (Leiponeura)alexanderi, G. kansensis, Eriocera, Hexatoma, and others

In or beneath damp cushions of moss

Dicranomyia badia,

In or beneath dry cushions of moss

Liogma, Dolichopeza, Oropeza, and others

In dry soil

Dicranoptycha, Cladura, Nephrotoma ferruginea,
Tipula cunctans, T. angustipennis,

In fungi

Limnobia

D. stidta, Tipula oropezoides, T. collaris, T. nobilis, T. ignobilis,

and others

and others
triocellala,L.

cinctipes,

Ula,

and

othersIn decaying vegetables, plant stems, manure,and like situations

Trichocera,
Limnobia indigena, Rhipidia
domeslica, R. maculata, and others

In wood:a.

In decaying wood, usually just beneath the barkDicranomyia
fidelis,rara,
D. macaieei, Rhipidia
R. bryanti, Elephantomvia, Teucho-

Gnophomyia, Limnophila unica,
Brachypremna, Ctenophora, Dictenidia,
Tipula trivittata, T. usitaia, and others
Tanypteralabis,

b. In nearly solid wood

On leaves of terrestrial plants:
a. On flowering plantsb. On mosses

Cylindrotoma
Liogma, Triogma (in some instances)

Mining in the leaves of plants

Dicranomyia foliocuniculator

Feeding.
in their— The various larvae

habits of feeding.

of crane-flies show a considerable diversity

The majority of species, as known, are herbiv-

orous, but a large group are decidedly carnivorous in their habits.

The Tipulinae feed on
occurring in their habitat,

the living vegetable tissue or plant remainsor,when pressed for food, they will eat earth-

Charles Paul Alexander718

worms (Patterson,

1908) and other soft-bodied animals.

Dissections of

Trpula abdmninalis show the food of this species to consist principally ofsmall filamentous algae, diatoms (Diatoma, Navicula, and others), androotlets of small

The alimentary tract is often crammed with
The species of Tipula and Nephrotoma that areplants.

sand or soil particles.

considered later in this paper under the heading
Economic Importance, effect their damage by devouring the living tissuesof the roots.
The Cylindrotominae feed on the living tissues of the plants
on which they dwell; in the case of Phalacrocera, Triogma, and Liogma
these are mosses, in Cylindrotoma they are the parenchyma and epiinjurious to plants,

dermis of higher plants(spermatophytes).

With this habit of feedingTheon plant tissues the extreme of sluggishness of motion is attained.

wood-l^oring species of Tipulinae (such as the species of Tanyptera andCtenophora) feed on the ligneous tissue of the trees in which they occur.
Other species of crane-flies injure young seedUngs by destroying the barkand the bast tissues.
The hexatomine and pediciine forms represent the opposite extreme,
being for the most part carnivorous or even cannibalistic in their habits.
In order to capture their prey they are of necessity rapid of movement,
and in this group the most graceful and active of all tipulid larvae arefound. The motions of the species are, at times, exceedingly agile andsnakelike.
The food consists of a variety of animal forms. Dicranota
has been recorded as feeding on worms of the genus Tubifex, Pedicia

usually feeds on the larvae of small insects, especially Chironomidae,

but the large species of this genus and of Eriocera are capable of capturing
almost any insect of a size equal to their own. The larvae of Eriocera
spinosa are able to inflict painful bites on tender parts of the skin of a man.

Penthoptera, Eriocera, Hexatoma, and most of the limnophiline groups
likewise feed largely on

midge larvae.

From one small larva of Limno-

phila (Dicranophragma) fuscovaria the writer has dissected out the remainsof two large midge larvae, whose heads were nearly half the size of that ofThe chitinized mentum in these predacious forms is very
weak or is lacking, allowing for a tremendous distension of the gular region.
The mandibles are always developed into powerful curved hooks which
serve well their purpose of grasping and holding the victims.
The
the captor.

esophageal region

is often retrorsely roughened to prevent the egress ofanything that has once started down the throat. In addition to theThe Crane-Flies of New Yorkvarious chitinized jaws,

legs,— Part719

II

heads, and other insect remains, the dis-tended proventricular regions usually show

a considerable

amountofsand particles and much plant tissue.

The pupa

When ready to pupate, the larva ceases feeding and becomes much
The pupa is formed within the last larval skin,which is then shed completely except in certain Cyhndrotominae and a
few scattered genera m other tribes, in which cases the larval skin adheres
The pupal existence is spent in or
to the posterior end of the abdomen.
near the haunts of the larva.
In the strictly aquatic genus Antocha the pupa lives in water in a silken
case, respiration being accomplished by means of the many-branchedbreathing horns. The species of Elliptera and certain Dicranomyia(simulans, for example) approach this aquatic condition.
The other
species of crane-flies with aquatic larvae known to the writer go to thesoil in order to pupate, this category including Eriocera, Hexatoma, Tipula caloptera, T. ahdominalis, and many others.
The pupae of some, at least, of the Cyhndrotominae attach themselvescontracted and sluggish.

to plant stems for the purpose of pupation.

The leaf-mining DicranoThe majority

myia foliocuniculator pupates within the larval passages.

of the limnobiine forms spend the pupal existence in silken cases to which

pebbles and particles of debris or plant tissues adhere.

The pupae of the Ptychopteridae have one of the two breathing horns
enormously elongated, the tip of this being projected above the waterlevel into the air for respiration.

Certain tipuline crane-flies have a some-what similar development of the breathing horns, discussed later.
The duration of the pupal existence is remarkably uniform thruout thegroup, averaging from six to eight days. The following table illustrates
this for the more representative genera and species.
Records whichhave not been determined sufficiently close, and which as stated areprobably too long, are indicated by an asterisk.

Chahles Paul Alexander

720

Duration of pupal existence (days)
ReferenceSpecies

Maximum Minimum
Ptychoptera rufocincla

Average41^2

nibimana
Bittacomorphella
Dicranomyia badia

10

11

7
*8

stultaRhipidia bryanti

7

Limnobia cinctipes
triocellata

Rhamphidia mainensis ....
Dicranoptycha winnemana

9
6
'10

Ula elegans

Epiphragma

6

Pseudolimnophila
Limnophila macrocera
fuscovaria.

.

.

.

9
9

8^
8i

^4

10
7

Uloraorpha7*9Eriocera fultonensis

7

7

7.3
6i
'14

7.3
61
10|

S

7h

Pilaria

longicornis

Hexfitoma megacera
Pedicia rivosa
Erioptera macrophthalma.

Molophilus hirtipennis ....

Gnophomyia tristissima

.

.

.

Ormosia nigripila
Teucholabis complexa
Elephantomyia

'10

5
'11

9
8
10

G

Liogma glabrata

11

10

Phalacrocera repHcala ....
Dolichopeza albipes
Ctenophora angustipennis
Diclenidia bimaculatas6

7

Cylindrotoma splendens.

.

.

6

.

7

Tipula maxima

'14

'10

oleracea

8

7

cayuga7.

ignobilis

Nephrotoma lineata
lunulicornis
analis7\6
107Beling, 1886(ass2/Zt;icoZa)'12

Beling, 1886

7^-

7.6
7i

,

dejectacollaris

7
S
10|

8
7
7
'12

Alexander ms.
Topsent, 1914-10
Alexander ms.
Alexander ms.
Alexander ms.
Shannon ms.
Alexander ms.
Alexander ms.
Alexander ms.
Alexander ms.
Alexander, 1915a
Alexander ms.
Alexander ms.
Alexander ms.
Alexander ms.
Alexander ms.
Alexander ms.
Alexander ms.
Alexander ms.
Alexander ms.
Baling, 1879:46
Alexander ms.
Alexander ms.
Hyslop ms.
Alexander ms.
Johnson ms.
Shannon ms.
Cameron, 1918
Miiggenberg, 1901
Bengtsson, 1897

8
7h
7'
7
'10

Lovett, 1915
Beling, 1873 b

DelGuercio, 1914:315
Alexander ms.
Alexander ms.
Alexander ms.
Alexander ms.
Beling, 1879 (as histrio)
Beling, 1879
Beling, 1886

The Crane-Flies of New York— Part II

721

ENEMIES
Crane-flies have many enemies and but few means of combating them.

Every stage of the crane-fly's existence is fraught with danger. The
larvae of the majority of species are soft-bodied, herbivorous creatures,
which form a choice morsel of food for carnivorous forms of many classes
and orders. The period when the adult fly has just emerged from thepupal hull is imdoubtedly the one in which the greatest danger is found,the insects are teneral and incapable of rapid motion.

for thenThe

adult flies are sometimes drowned in the sea or other large bodies of wateranddead fragments cast up

their

in

windrows on the

shore.

A veryunusual instance of this kind is recorded for Tipula oleracea by Patterson

At their best, the adult flies are poor, awkward fliers and are
by a wide range of species, as indicated in the foUowing(1908).

easily captured

pages.

Predatory natural enemies
Vertebrates

Mammalia.

— Undoubtedly manyof the smaller mammals

prey on the

In another paper (Alexander, 1919 d:value of the larvae of an undeterdiscussed
thewriterhas
776-777) the
Pribilof
Islands as an article of food for the
Tipulathespecies
of
in
mined

larvae of various species of Tipula.

arctic fox.

These larvae occur in enormous numbers beneath the lichensof the tundra and the foxes can easily get them at a time when other foodisscarce or unobtainable.

Mice, shrews, and moles find an importantWhite (1914) states that in North

element of their food from this source.

Wales the European mole, Talpa europaea, eats, on an average, twenty
crane-fly larvae a day, these constituting one of the main foods of thismammal.

— Birds are well-known enemiesof crane-flies, both in their immaHyslop (1910:129-130) lists ninety-one
Baer (1913)
species of birds which are known to feed on crane-flies.
describes ravens and starlings as feeding on the larvae of a species ofTipula.
While working with the United States Biological Survey, theHis thanks
writer was given the opportunity to examine files for records.
for assistance in this work are due to Messrs. Kalmbach, McAtee, andWetmore. The records of the Svu'vey are based on an examination of
the contents of tlie stomachs of many thousands of individuals, andfurnish very valuable and fairly complete data on the North American

Aves.

ture stages and as adult9flies.

Charles Paul Alexander722

birds which prey .on various stages of the Tipulidae.
These records arehere arranged according to the latest check Ust of the American Ornithologists' Union:•''

Longipennes:
Laridae
Larus delawarensis Ord.

Ring-billed gullFranklin's gull
Sterna hirundo Linn. Common tern

L. franklinii Rich.

Anseres:/

Anatidae

Mergus americanus Cass.
Anas platyrhijnchos Linn.

Merganser

MallardA. nibripes Brewst. Black duck
Mareca americann (Gmel.). Baldpate
Nettion carolinense (Gmel.).
Green-winged teal
Dafila acuta (Linn.).
Aix sponsa (Linn.).

Pintail

Wood duck

Herodiones:
Ardeidae:
Egretta candidissima candidissima (Gmel.).
Snowy egret
Butorides virescens virescens (Linn.).
Green heron
Limicolae:

Phalaropodidae
Phalaropus fulicarius (Linn.). Red phalarope
Lobipes lobatus (Linn.). Northern phalarope
Steganopus tricolor Vieill. Wilson's phalarope
Recurvirostridae
Recurvirostra americana Gmel.

Himantopus mexicanus (Miill.).

AvocetBlack-necked stilt

Scolopacidac:
Philohela minor (Gmel.).
American woodcock
Gallinago delicata (Ord).
Wilson's snipe
Arquatella maritima maritima (Briinn.).
Purple sandpiper
Pisobia aurita (Lath.). Sharp-tailed sandpiper
P. maculata (Vieill.). Pectoral sandpiper
P. bairdii (Coues). Baird's sandpiper
Limosa fedoa (Linn.). Marbled godwit
Totanus fiavipes (Gmel.). Yellow-legs
Heteractilis incanus (Gmel.).
Wandering tatler
Bartramia longicauda (Bechst.). Bartramian sandpiper

Charadriidae
Charadrius dominicus dominicus (Miill.).
Oxyechtis vociferus (Linn.).

,

Golden plover

Killdeer

Aphrizidae:
Arenaria inierpres interpres iLinn.)
Turnstone
Gallinae:
Tetraonidae:
Bonasa umbellus umbellus (Linn.). Ruffed grouse
Raptores:
Buteonidae:.'.

Ictinia mississippiensis (Wils.).

^

Mississippi kite

'Check list of North American birds, 3d ed., p. 1-130.

American Ornithologists' Union.

I

„

J

•

I

i

1910.

The Crane-Flies of New York
Coccyges:
Cuculidae:
Coccyzus americanus atnericanus (Linn.).

— PartII

Yellow-bil.ed cuckoo

Black-billed cuckoo
C. erythrophthalmus (Wils.).
Cuculus canorus telephonus Heine. Kamchatka cuckoo
Pici:

Picidae:
Downy woodpecker
Dryobates pubcscens pubescens (Linn.).
D. nuttallii (Ganib.). Nuttall's woodpecker
Sphyrapicus varius vnrius (Linn.). Yellow-bellied sapsucker

Williamson's sapsucker
Melanerpes enjthrocephnlus (Linn.). Red-headed woodpecker

S. thyroideus (Cass.).

Asyndesmus lewisi Riley.

Lewis's woodpecker
Flicker

Colaptes auratus auratus (Linn.).

Macrochires:
Caprimulgidae:
Antrnstomus carolinensis (Gmel.).

Chuck-will's-widow

Nighthawk
Chordeiles virgininnus virginianus (Gmel.).
Texas nighthawk
C. acutipennis texensis Lawr.
Cypselidae:
Chaetura pelagica (Linn.). Chimney swift
Vaux's swift
C. vauxi (J. K.Towns.).
Aeronautes melanoleucus (Baird). White-throated swift
Trochilidae:
Calypte anna (Less.). Anna's humming bird
Selasphorus rufus (Gmel.). Rufous humming bird
Passeres:

TjTannidae:
Tyrannus tyrannus (Linn.). Kingbird
Arkansas kingbird
T. verticalis Say.
T. vociferans Swains.

Cassin's kingbird

Myiarchus crinitus (Lum.). Great-crested flycatcher
Sayornis phoebe (Lath.). Phoebe
Say's phoebe
S. sayus (Bonap.).
Black phoebe
S. nigricans (Swains.).

Myiochanes virens (Linn.).

Wood pewee

M. richardsonii richardsonii (Swains.). Western wood pewee
Empiionax fl'wiventris (W. M. & S. F. Baird). Yellow-bellied flycatcher
E.
E.
E.
E.
E.
E.

difficilis difficilis

traillii traillii

Western flycatcher
Acadian flycatcher

Baird.

virescens (Vieill.).

(Aud.).

Traill's flycatcher

minimus (W. M. & S. F.- Baird).
wrighlii Baird.
griseus Brewst.

Least flycatcher
Wright's flycatcher

Gray flycatcher

Corvidae
Pica pica hudsonia (Sab.).

American magpie

Blue jay
Cyanocitta cristata cristata (Linn.).
Steller's jay
C. stelleri steUeri (Gmel.).
Corvus brachyrhynchos brachyrhynchos Brehm.
Fish crow
C. ossifragus Wils.
Sturnidae:
Sturnus vulgaris Linn. Starling
Icteridae:

Dolichonyx oryzivorus (Linn.). Bobolink
Molothrus ater ater (Bodd.). Cowbird

American crow

723

724

'

Charles Paul Alexander

Passeres (continued):
Icteridae (continued)

Xanthocephalus xanlhocephalus (Bonap.)- Yellow-headed blackbird
Red-winged blackbird
Agelaius phueniceus phoeniceus (Linn.)A. gubernalor calijoniicus Nels. Bicolored redwing

Meadow lark
Sturnella magna magna (Linn.)Orchard oriole
Icterus spurius (Linn.).
Baltimore oriole/. galbula (Linn.).
Bullock's oriole/. bullockii (Swains.).
Euphagus cyanocephalus (Wagl.). Brewer's blackbird
Quiscalus quiscula quiscula (Linn.). Purple grackle
Fringillidae:

Aleutian rosy finch
Leucosticie griseonucha (Brandt).
Plecirophenax nivalis nivalis (Linn.). Snow bunting
P. hyperboreus Ridgw. McKay's .«now bunting
Lapland longspur
Calcarius lapponicus lapponicus' {hinn.)
Passerculus sandwichensis savanna (Wils.). Savanna sparrow
White-crowned sparrow
Zonotrichia leucophrys leucophrys (J. R. Forst.).
White-throated sparrow
Z. albicollis (Gmel.).
J unco hyemalis hyemalis (Linn.). Slate-colored junco
Passer domesticus (Linn.). English sparrow
Melospiza melodia m^lodia (Wils.). Song sparrow
M. lincolnii lincolnii (Aud.). Lincoln's sparrow
M. georgiana (Lath.). Swamp sparrow
Fox sparrow
Passerella iliaca iliaca (Merr.).
TowheePipilo erythrophthalmus erythrophthalmiis (Linn.).
Passerina amoena (Say). Lazuli bunting.

Tangaridae:
Piranga ludoviciana (Wils.). Western tanager
P. erythromelas (Vieill.). Scarlet tanager
Summer tanager
P. rubra rubra (Linn.).
Hirundinidae:
Prague subis subis (Linn.). Purple martin
Cliff swallow
Petrochelidon lunifrons lunifrons (Say).
Hirundo erythrogastra Bodd. Barn swallow
Tree swallow
Iridoprocne bicolor (Vieill.).
Northern \iolet-green swallow
Tachycineta thalassina lepida Mearns.
Riparia riparia (Linn.). Bank swallow
Rough-winged swallow
Stelgidopteryx serripennis (Aud.).
Bombycillidae:
Bombycilla cedrorum Vieill. Cedar waxwing
Vireonidae:
Red-eyed vireo
Vireosylva olivacea (Linn.).
Philadelphia vireo
V. philadelphica (Cass.).
Warbling vireo
V. gilva gilva (Vieill.).
Blue-headed vireo
Lanivireo solitarius solitarius (Wils.).
White-eyed vireo
Vireo griseus griseus (Bodd.).
Bell's vireo
V. bellii bellii Aud.
Least vireo
V. bellii pusillus Coues.
Mniotiltidae:
Mniotilla varia (Linn.). Black-and-white warbler
Worm-eating warbler
Helmitheros vermivorus (Gmel.).
Dendroica aesiiva aestiva (Gmel.). Yellow warbler
D. coronata (hum.). Myrtle warbler

D. auduboni avduboni (Towns.).

Audubon's warbler

The Crane-Flies of New York— PartII

725

Passeres (continued)
Mniotiltidae (continued):

D. dominica domhiica (Linn.)- Yellow-throated warbler
Oporornis tolmiei (J. K. Towns.). MacRillivray's warbler
Geothlypis Irichas trichas (Linn.).
Maryland yellowthroat
Icteria virens virens (Linn.).
Yellow-breasted chat
Wilsonia pusilla pusilla (Wils.). Wilson's warbler
Setophaga ruticilla (Linn.). American redstart
Motacillidae:
Anlhus rubescens (Tunst.). Pipit

Mimidae:Mimus polyglottos polyglottos (Linn.).

Mocking bird

CatbirdDumetella carolinensis (Linn.).

Toxostoma rufum (Linn.). Brown thrasher
T. redivivum (Gamb.).
Californian thrasher
Troglodytidae:
Heleodytes brunneicapillus couesi (Sharpe).
Cactus wren
Salpinctes obsoletus obsoletus (Say).
Rock wren
Catherpes mexicanus punciulatus Ridgw.
Dotted canon wren
Thryothorus ludovicianus ludovicianus (Lath.). Carolina wren
Thryomnnes bewickii bewickii (And.). Bev/ick's wren
Nanrms hiemalis hiemnlis (Yieill.). Winter wren
Telmatodytes palustris palustris (Wils.).
Long-billed marsh wren

Paridae:

Baeolophus inornatus inornatus (Gamb.).

Plain titmouse
Penthestes atricapillus atricapillus (Linn.).
ChickadeeP. carolinensis carolinensis (Aud.). Carolina chickadee

P. gambeli gamheli (Ridgw.). Mountain chickadee
P. hudsonicus hudsoniciis (Forst.). Hudsonian chickadee
PsaUriparus minimus minimus (J. K. Towns.). Bush-tit

Chamaeidae:
Chamaea fasciata fasciata (Gamb.).

Wren-tit

Sylviidae:-Regulus calendula calendula (Linn.). Ruby-crowned kinglet
Turdidae:
Myadestes townsendi (Aud.). Townsend's solitaire
Hylocichla mustelina (Gmel.).
Wood thrush
H. fuscescens fuscescens (Steph.). Veery
H. aliciae aliciae (Baird). Gray-cheeked thrush
H. ustulata ustulata (Nutt.). Russet-backed thrush
H. ushdata swainsoni (Tschudi). Olive-backed thrush
H. guttnta pallasii (Cab.). Hermit thrush
Planesticus migratorius migratorius (Linn.).

American robin

The principal families that feed on the adult flies are the Caprimulgidae,
the Cypselidae,

the Tyrannidae,

the Icteridae, the Hirundinidae,

theand the Turdidae.
Many species of birds feed on the larvae of crane-flies, the more notable

Mniotiltidae, the Troglodytidae, the Paridae,of these being the water-fowl, Anatidae, the shore

and the thrushes, Turdidae.
sided warbler, DendroicaSim(1907)'pensijlvanica

birds, Scolopacidae,

has recorded the chestnut-(Linn.) as feedingon Tipula sp.

Charles Paul Alexander

72G

In Europe the raven and the starhng are important, as ab-eady stated.
Patterson (1908) records the starhng as eating great numbers of Tipuln
oleracea

and as boring into the

soil

in

searchofthelarvaeofthisspecies.

be noted that in the foregoing list no species of doves or pigeons
The following note

It willare recorded in this country as feeding on Tipulidae.

on an Australian pigeon shows the importance of the larvae as a food forthese birds: ^Mr. North exhibited the head, crop, and gizzard of a wonga-wonga pigeon (Leucosarcia
picata Lath.) shot by Mr. H. J. McCooey in a myrtle scrub at Upper Burragorang on the
21st inst.
The crop is absolutely crammed with dipterous larvae {Hahromastix cinerascensSk.) and undigested portions of them mixed with seeds, berries, and earth appear also in the
gizzard.
As the larvae are known to be destructive to grass, the wonga-wonga would appear
to be deserving of consideration.

Amphibia.

— Crane-flies, both larvae and adults, form a considerable

food of many Amphibia. The studies by Needhamshow this to be true in the case of the bullfrog, Rana catesMimz (1920) studied the food habits of eight species ofheiana Shaw.
Anura and found that five fed on crane-fly larvae or adults. These
species were Rana clamitans Latr., R. sylvatica Lee, R. palustris Lee,The following additional
R. pipiens Schreb., and Hyla crucifer Wied.

element of the(1905:13)

records are given:
Desmognathus fusca Raf. Dusky salamander. Wings of a Limnophila found in a specimen from Ithaca, New York. (A. A. Noyes.)
Rana clamitans Latr. Green frog. Two larvae of a Tipula, near dejecta Walker, foimd(S. W. Frost.)
in stomach.

Bufo lentiginosus woodhousei Girard. Centerville, Utah, April 27, 1912. Twenty-sLx(E. R. Kalmbach.)
per cent of the food of this species consisted of a large tipulid.
Bufo sp. Washington, D. C, May 17, 1890. One per cent tipulid larvae. (W. L.
McAtee.)Pisces.

— The larvae of

crane-flies furnish favorite morsels formany

carnivorous species of fish, and as a consequence they are in considerable

demand with fishermen

as bait for bass

and other game

fish.

These

include the larvae of the larger species of Eriocera and many large semi-aquatic species of Tipula, especially Tipula caloptera and T. abdominalis.

Fragments of the adult flies are often found in the stomach contents offish, notably species of trout, most of these pieces being of individuals that*

Linn. Sor. New South Wales.

Proc, ser. 2:9:585.

1894.

The Crane-Flies of New York— Part II

727

fallen into the water or were captured while newly transformed.
Forbes (1888) records Coregonus and Hyodon as feeding on the eggs,
and Notropis as feeding on the larvae, of crane-flies. Levander (1909:1)

hadrecords Perca fiudatilis Linn, as feeding on larvae of Tipulidae.

Invertebrates

Arachnida.

— Some spiders are notable enemiesofcrane-flies,

whicheasy victims while in a helpless, teneral condition. These spiders
represent many families, such "as the Thomisidae, the Lycosidae, the
fall

the

Attidae,also,

Epeiridae,

and others (Alexander and Lloyd, 1914:15;
An accomit of their preying on the larger

Alexander, 1915 c: 144).

crane-flies

ofthe genus Tipulataughannock (page 1013).
four species of crane-flies

given

is

under the discussionofT.

In a recent paper Bilsing (1920) has recorded
eaten by spiders.

thirteen species arranged in fourfamilies

the Epeiridae, and the Agelenidae.

J.

angustipennis which was being eaten

These spiders represented— the Lycosidae, the Attidae,

R. Malloch found a female Tipula

by a crab spider, determined by
The spider did not release its hold

Mr. Banks as Xysticus ferox Htz.
it was placed in a jar of cyanide.

until after

Hexajjoda.

— Odonata: The following records

feeding on adult crane-flies are available:
G. ventricosus Walsh, found feeding on

Indiana,ofdragon-flies

found

Gomphus vastus Walsh andTipula bicornis at South Bend,

May 30, 1914 (E. B. Williamson); Helocordulia uhleri (Selys),

foimd feeding on Eriocera longicornis (Alexander, 1915 c: 152); Agrion
puella Linn., found feeding on Erioptera flavescens (Campion, 1914:498).
C. H. Kennedy has sent the writer a male and a female specunen of a small
crane-fly, Teucholabis pabulatoria Alex., which he found in the mouth ofa damsel fly, Hetaerina tricolor Burm., collected in Guatemala by Professor
Needham and Hart (1901-03 [1901]: 47) record the nymphs of
Hine.

Anax Junius
Diptera:(Dru.) feeding on the larvae of Tipulidae.

The adult flies of at least three families of the order Diptera—-the Asihdae, the Empididae, and the Scatophagidae — and the larvae

The
Anthomyiidae, are notable enemies of crane-flies.
Kirby (1892), Poulton (1906-O7), Bromley (1914), Alexander(1915 c), and McAtee and Banks (1920) on the first three of thesefamihes may be summarized as follows:ofa fourth,

records ofCharles Paul Alexander

728

Lasiopogon ductus (Fabr.)

Prey
Nephrotoma lineata (Scop.) (Poulton, 1906-07,

Neoitamus cyanunis {L\\'.)(mcopu\a.t\onvfith.

Tipula scripta Meig. (Poulton, 1906-07)

female, the latter with the crane-fly)
Proctacanthus philadelphicus Macq.
Asilus sp.

Nephrntomn sp. (2 records, Bromley, 1914)
Nephrotoma ferniginea (Fabr.) (3 records,

Asilidae (robber flies)

as histrio [Fabr.])Alexander, 1915 c)
Tipula sp. (McAtee and Banks, 1920:30)

Asilus flavofemoratus Jiuxe
Asilus notatus Wied.

Nephrotoma ferniginea (Fabr.) (McAtee andAsilus paropus Walk.

Epiphragma

Asilinae

Banks, 1920:31)
Tipula lateralis Meig. (Poulton, 1906-07)

Banks, 1920:31)
solatrix(0.

S.)(McAtee and

The Empididae, or dance flies, are small, predacious flies which are closely related to therobber flies and largely replace them in moist shaded situations. They unquestionably
play a most important part in the economy of the Tipulidae, since both groups are practicallyconfined to the same general situations and, in the temperate regions at least, representatives
Macquart (Kirby, 1892:229), discussingof the two groups are invariably found together.
Empis livida Linn., writes as follows: " Among the thousands of pairs which I have noticed
resting on bushes or hedges, nearly all the females were engaged in sucking some insect,sometimes small Phryganidae or Ephemeridae, but more often Tipulidae. They busy
themselves with feeding and perpetuating their species at the same time." Howlett (1907)
The following records are fromrecords Empis bnrealis Linn, as feeding on Tipulidae.
Poulton (1906-07:380-382): Empis tesselata Fabr. prej^s on Tipula lunata Linn., T.
paludosa Meig., and other species; Empis livida Linn, preys on Dicranomijia sp. (It is
probable that the long-legged flies, Dolichopodidae, likewise play an important r61e inthe lives of the Tipulidae, but no records are available to the writer to conflrm this belief.)

The Scatophagidae (Cordyluridae)

or dung flies, unquestionably play an important partthe smaller crane-flies (Limnobimae). The following records indicate thisrelationship, the last being supplied by Malloch:,in the lives of

Enemy
Scatophaga suilla (Fabr.)
Scntophaga sp. {stercoraria?)
Scatophaga sp.
Scatophaga stercoraria (h.) and squalidaMeig.

Prey
Dicranomyia lulea (Meig.) (Poulton, 1906-07)
Brioptera sp., (Poulton, 1906-07)

Hexatomamegacera{0. S.) (Alexander, 1915 c^
Trichocera (Malloch, 1911)

Comparatively recently the larvae of Anthomyiidae have been foundto play a highly important role in the economy of other insects living inthe same haunts.

(1917) may beThis student found the follow-

The very important paper by Keilm

consulted in connection with this point.

ing Anthomyiidae associated with TipuUdae:

Graphomyia maculata Scop.,feeding on larvae of Ptychoptera contaminata (L.) (Keilin, 1917:354-360);

Allognota agromyzina Fall., associated with and possibly feeding on theof Ula macroptera Macq. (page 360-362 of reference cited);
Phaonia cinda Zett., feeding on larvae and pupae of Mycetobia pallipes(page 362-375 of reference); Phaonia goherti Mik, associated with andpossibly feeding on the larvae of Gnophomijia tripudians Bergr. (pagelarvae— Part II

The Crane-Flies of New York729

375-377 of reference); Mydaea pertusa Meig., possibly feeding on larvaeof Tipulidae (page393-396 of reference).

A species of Phaonia with presumably parasitic habits is discussed on
page 732 of this memoir.
Coleoptera:

The adults and the larvae of the following species of groundbeetles (Carabidac)
flies(Hyslop,

have been recorded as important enemies of cranePoedlus lucuhlandus Say; Micromaseus femoralis

1910):(Kirby); Platynus sp.; Harpalus pennsylvanicus Dej.; H. caliginosus Fabr.
Most of these were recorded by Webster (1893 a: 241) as feeding on the
injurious crane-fly Tipula bicornis.

The larvae of Elateridae (wireworms)

are enemies of crane-flies.

The

writer has seen several feeding on large larvae of Tipula trivittata.

Ants (Formicidae) midoubtedly capture and devour
many crane-flies, these generally being pupae or teneral adults. Hyslop(1910) cites the record of an Aphaenogaster fulva Roger which was observed

Hymenoptera:

Chapman (1918:191)

dragging a hving adult tipuhd over the groimd.

records feeding species of Myrmica on adult TipuUdae which were readily

eaten by the ants in preference to most other insect food.

On August 2,

1917, at Larned, Pawnee County, Kansas, along the bank of the Arkansas

was dragging a still living
pupa of Gonomyia kansensis from its bmrow in the sand at the bank of
the river. The ant had the pupa about the head and carried it off despite
its struggles.
Both the Gonomyia and the fonnicid were conunon andthe tragedy as described was not accidental.
The Pemphredonidae (Mimesinae) and the Crabronidae, fossorial wasps
in the Hawaiian Islands, bear an important relationship to the smaller
Tipulidae dwellmg in the same haunts. Writing of the Mimesinae, aRiver, the writer noted a small ant which

subfamily of the Pemphredonidae, Perkins (1913:bcxxv) says:
All the Hawaiian species of both genera are true forest insects and most of them may beseen in large numbers, where thej^ occur, flying around ferns and bushes in sunny places.
Tlie males are often much more numerous apparently than the females, but this is due tothe more retiring habits of the latter, which, when they have begun to provision their nests,
This consists of the endemic
frequent dark, shady and damp places in search of their prej-.
Although I have often
Limnobiidac or dadd.v longkgs, which live in such places.
watched females of these wasps returning with prey to their burrows, it was always Tipulidaethat they carrif^d to the nest. Their burrows are usually made in the ground and are often
drilled down from beneath a stone, this no doubt serving to keep the burrow sufficiently
.

dry.

.

.

730

Charles Paul Alexander

ReferringtoLimnobiinae,

Perkins

says

(page

clxxxiiofthe samereference)

The specios of Dicmnomyia are endemic, and to the five described,
others and perhangmany more will, no doubt, be added. They are common insects,

sometimes attracted bylight and sometmies observed at rest in thedaytime, or on the wing in the dark cavity ofsome hollow tree. In the latter case a number are usuallyseen flying together, rising andfallmg m their flight in the narrow space of a few
feet.
We have bred one or more spedesfrom decayedwood, overgrownwith damp moss. These fragile flies are the favourite
prey ofthe endemic predaceous wasps of the family
Mimesidae, and some of the Crabronidae likewise gather them.
Consequently one may find the females of these wasps
investigating
damp, dark places, where the Dicranomyia are likely
to be found, but which ordkiarilv
orcunaruy
would have but bttle attraction for such sun-loving insects.

Writing of the Crabronidae, Perkins says (page kxxviof the reference
cited)
Xenoa-abro hawaiiensis on one occasion was caught
carrying off Lispe [Anthomyiidae]-^numjiiaaej
and on another occasion a species of the Ihnnobud Dicranomyia!

De Meijere (1920:59) records Crabro (Rhopalum) tibiale as provisioning
its nest principally with Molophilus

armatus Meij.

Parasitic natural enemies

External parasites

Besides the predatory forms just considered, thevarious stages of themany crane-flies are threatened with parasites which areequally effective altho more insidious in their
method of attack. There
are unquestionably many parasites of crane-flies
concerning which nothingat all is known at this thne.
It is inconceivable that such speciesas
Holorusia grandis, Tipiila abdominalis, and other large
and common forms
do not serve as hosts for parasites as yet unknown.
It is this field ofmvestigation that now promises some of the most valuable
results.
Hexapoda.— i:\ie only parasitic insects of which the writerknows
are the dipterous Tachinidae.
Glover (1874) states that in Europe certain
Tipulae are destroyed by a proctotrupid parasiteof the genus Diapria
existence of

Latreille.

Kieffer, inliis

monograph of the Diapriidae

mention this and the writer has not seen it referred(1911), does notto elsewhere.

Itisprobable that the reference pertains to some ofthe smaller Tipula-Hke
forms of another family. However, it is very
strange that one or moreof themany families of parasitic Hymenoptera do not infest somestage
and it may be confidently predicted that suchparasitesof the Tipuhdae,
will one day

be discovered.

The Crane-Flies of New York
Most

Tachinidae:of— Part II

731

the species of the interesting genus Siphona(Bucentes) are parasitic on the caterpillars of various Lepidoptera.
have been recorded as parasites of the larvae of species of Tipula.

Two
The

Siphona cHstata (Fabr.), a constant parasite of
the larvae of Tipula maxima (Behng 1886, Czizek 1913, Riedel 1913,
Thompson 1915). The method of attack by the parasite has been
better-knownof theseis

The large, massive larvae of this species
amphibious or semi-aquatic, living near the margins ofstreams and other bodies of water. They live in the mud or in the water
beneath the surface, at times coming up to breathe the air thru the two
described by Roubaud (1906).
of crane-fly arelarge spiracles at the posterior end of the body.

It is at this time,when

the spiracular disk is projected thru the surface film and the spiracles areopen, that the tachinid is supposed to dart down and deposit its eggs in
the open stigmata.

The young parasite is somewhat elongated, yellow-

orange in color, and about a millimeter in length, and is completely
It is connected with one of the
inclosed in a firm, membranous cyst.
ish

two principal tracheal trunks of the host by a sort of chitinized calyx
which opens into the trachea. At the time of their pupation the parasites detach their organs of fixation, and with their heads perforate theIn nature this departureskin of the host and enter the humic earth.
coincides almost exactly with the time of leaving the water by the host

The pupal duration of the paraabout three weeks. Rennie (1912) gives Siphona geniculata (DeGeer) as a parasite of an undetermined species of Tipula.
Species of the genus Admontia are important parasites of the larvae of
crane-flies.
In Europe, Admontia arnica (Meig.) is parasitic on species offor the purpose of pupation in the earth.
site isTipula, while in America A. pergandei Coq. is an important parasite ofthesmoky crane-fly, Tipula cunctans (Hyslop, 1910:128).

In thecollec-Natural History Survey are several specimens of
A. pergandei bred from larvae of the smoky crane-fly (larvae taken at
Averin (1913)Ur])ana, Illinois, parasites emerged October 9 to 25, 1900).
and Lutchnik (1916), in Russia, record the larvae of a Tipula, in one casetionof theIlUnois— as being very heavily

at least— that of the economic species T.

infestedby an undetermined tachinid which may have been the Admontia

oleracea

species mentioned above.

A third genus of Tachinidae, Sturmia, has recently been reported to
the writer by J. D. Tothill as being parasitic on the larva of a large crane-

Charles Paul Alexander

732

undoubtedly a tipuline form. The larva was found at Chelsea, near
Ottawa, on May 27, 1906, by Dr. James Fletcher, the parasite issuing on

fly,

June 27. The tachinid is to be described as Sturmia tipulensis Tothill.
The genus Trichoparia is parasitic on Ctenophora and its allies.
Altho the Tachinidae are the only dipterous parasites that are definitely
known from the Tipulidae, the writer has a record of a species of Phaonia,
of the family Anthomyiidae, which possibly is a parasite.
Among some
Diptera sent to Malloch for naming was one specimen of a Phaonia bearing
the label " Bred from tipulid pupa, ace. no. 14022."
The writer isindebted to R. H. VanZwaluwenburg for the following facts concerning
The material was taken by Harry L. Parker on June 6,
this specimen:
Under litter
1916, on South Mountain, near Hagerstown, Maryland.
and growth of Virginia creeper on rock a tipulid pupa was found. The
pupa had lieen eaten out inside and there was a hole in the side of its body.
About a half inch away from this pupa was found a newly formed pupa
The adult emerged on June 23, 1916, and wasof an anthomyiid fly.
pinned. It is possible, of course, that the Phaonia was merely feedingon the tipulid, as recorded for this genus elsewhere in this paper (page728), and that the species is a predatory enemy rather than a parasite.
Arachnida.
Young mites of species of Trombidium, Rhyncholophus,and other genera of Arachnida, are frequently found attached to the bodyand the wings of adult crane-flies. This attachment is by anal filaments,
and it is doubtful whether any real injury results to the host therefrom.
These cases are probably instances of phoresy to provide a means of dispersal of the mite.
At various times the writer has found in northeastern
America a large number of adult crane-flies so infested, and practicallyevery genus and many species are included among the number.
Fungi:
Among the enemies of crane-flies, both to the immature andto the adult stages, probably none are more lethal in their action than——certain parasitic fungi.

The largest groups

are the Entomophthoraceae(Phycomycetes)of insect-fungous parasites

and the Laboulbeniaceae

Ascomycetes)
Dr. Roland Thaxter informs the writer that, altho thelatter group infest other Diptera, he has never seen specimens on Tipuhdae,(.

and none have been recorded in the literature. The writer is greatly
indebted to Dr. Thaxter and to A. T. Speare, Government MycoentomoloDr. Thaxter sent to the writer for detergist, for data on this subject.
mination a considerable number of specimens of crane-flies with their para-

The Crane-Flies of New Yorksites,— Part II

733

these including the types of many of his Entomophthoraceae andsome new species not yet described.
the following notes
Class,'

From these data and the htera^ture

may be given:

Phycomycetes

Family, Entomophthoraceae
1. Enlomophthora (Empusa) pachyrrhinae Arthur.
a manuscript name.

On the larvae of Nephrotoma ferruginea;

E. (E.) sepulchralis (Thaxt.). On an adult Tipiila of the subunicolores group.
E. (E.) caroliniana (Thaxt.) [= E. arrenoctona Giard]. On an adult male of Tipula
enlomophthorae.
On male adults of Tipula paludosa (Giard 1888, Villeneuve
4. E. (E.) arrenoctona Giard.
1910, Picard 1913).
5. E. (E.) iipulae Fres.
On Tipula maxima (Lohde, 1872).
A species which is probably this one is recorded by
6. E. (E.) conglomerata (Sorokin).
Thaxter on larvae and adults of Tipulidae.
Recorded by Thaxter from adult Tipulidae.
7. E. (E.) sphaerosperma Fres.
Recorded on at least two occasions from the adxilt flies of Tipula
8. E. (E.) grylli Fres.
ultima Alex. [= T. flavicans Fabr.].
On adult females of Pseudolimnophila noveboracensis.
9. E. (E.) punctata Thaxt., ms.
10. E. (E.) dipterigena (Thaxt.).
On the adult flies of several genera of Limnobiinae, as
2.

3.

follows:

Molophilus

hirtipennis,

Limnophila aprilina,

Pseudolimnophila nigripleura,

Uhmorpha pilosella,Penthoptera albitarsis.
11. Isaria sp.
On adults of Dicranomyia pubipennis.
of this same genus on various tropical Tipulidae.
Class,

Additional undetermined species

Hyphomycetes.

Family, Alucediniaceae.
12. Sporoirichum densum Link.
1910:130).

On

Tipula cunctans Say[= T.

infuscaia Lw.] (Hyslop,

Internal parasites

Crane-fly larvae and adults are infested by numerous protozoan
and bacterial parasites, the more conspicuous and constant being the
Sporozoa (Gregarinidae) and the Bacteria. General references to thissubject may be found in Leger (1892), Labbe (1899), and Minchin(1903).

More especial references are made to the parasites of Ptychoptera

by Leger and Dul^oscq(1909), to those of Dicranota by Miall (1893:
Ctenophora by Anthon (1908:542), and to those of
Tipula l)y Mackinnon (1913). In the case of Ctenophora the parasites
were frequent between the cells of the alunentary canal, especially in
The writer has fomid a large gregarine very
the proventricular caeca.
abundant in the alimentary canal of the larva of Pedicia albivitta,
many of the individuals being very heavily infested. Gamkrelidze(1913 b) records gregarine and nematode parasites in Gnophomyia237), to those oflarvae.

Charles Paul Alexander734

The more important recorded protozoan parasites are as follows:
Class, Sporozoa.
Subclass, Telosporidia.

Order, Cocoidiomorpha.
Family, Adcleidae.
1. Adelea tipulae Leger.
In the intestine of species of Tipula.
Order, CrcRarinida.
Family, (iri'gariuidae.
In the intestine of the larva of Limnohia sp.
2. Hirmoq/sti.s polijmorpha (L6ger, 1892:118).
In the alimentary canal of Tipula oleracea, Nephrotoma
3. H. ventricosa (Leger, 1892 :111).
pratensis, and other species.
4. Gregarina longa (Leger, 1892:117).
In the alimentary canal of a species of Tipula.

Family, Actinocephalidae.
Adinocephalus tipulae Leger (1892:141). In the alimentary canal of Tipula larvae.
Probably the same species has been recorded from the larvae of a species of Ctenophora.
6. Pileocephalus striatus LSger & Duhoscq (1909:887-89^).
In the mid-intestinal epithelium
The Pileocephalus live in the epithelium of theof the larva of Ptychoptera contaminata.
mid-intestine, attaching themselves to the epithelial cells and hypertrophying the adjoinThey obtain their nutriment from the food that penetrates into the cells.
ing tissues.
5.

Family, Stylorhynchidae.
Near Stylorhynchus (Miall, 1893:237).

7.

In the stomach of larvae of Dicranota bimaculata.

Subclass, Neosporidia.
Order, Cnidosporidia.
Suborder, Microsporidia.

Family, Nosematidae.
Nosema stridum Monz. (Moniez, 1887).

8.

of

In muscles, conjunctive tissue, and other parts

Nephrotoma pratensis.

Gurleya francottei L^ger & Duboscq (1909:894).
of the larva of Ptychoptera contaminata.

9.

In the epithelium of the mid-intestine

Class, Flagellata.

Family, Trypanosomidae (Herpetomonadidae).
Crithidia campanulata Leger.
At the juncture of the mid- and hind-intestines in thelarva of Ptychoptera contaminata (Leger & Duboscq, 1909:898-900).

10.

The writer is indebted to Dr. R. Kudo for assistance in determining theterminology used above.
Bacteria are frequently found in crane-fly larvae.

Leger and Dul)oscq(1909:900-901) record undetermined spirochaetes in the epithelialofthe posterior part of the mid-intestine of Ptychoptera

cells

contaminata.

Dr. Hugh Glasgow, of the Geneva Experiment Station, informs the writerthat in Illinois a large tipuhd larva, probably that of Tipula abdominalis,
living in the leaf-drift of prairie streams,

isheavily infested with bacteria.

Most of the specimens observed had an abundance of small coccus andspirochaete forms, with occasional specimens of a gigantic bacillus measuring from forty to eighty microns in length
bacilli infest

the hind-gut of the larva.

and disporous.

These large

The Crane-Flies of New York— PartII

735

HISTORICAL SUMMARY OF THE IMMATURE STAGESThe discovery and description of the immature stages of crane-flies
has by no means kept pace with the taxonomic work that has been doneon the adult flies. There are a very large number of common species inevery countiy concerning whose early stages nothing at all is known,
and the knowledge that exists concerning a comparatively small number
Among these may be mentioned
results from the labors of a few students.
Beling, Mik, Brauer, Hart, Needham, and Malloch.

Some other workers

have given excellent descriptions of single species, while still others havecontributed important treatises on the anatomy, morphology, histology,
and related subjects of the different species. The descriptions of theearlier

workersare,for themostpart, of historical interest only,

butin a few cases they are exceptional and are still the main source of informa-tion concerning certain species.

Many of the foremost workers on thetaxonomy of the adult flies have summarized the earlier writings on theimmature stages but have themselves contributed little to the subject,among these bemg Schiner, Loew, Osten Sacken, Skuse, and Brunetti.
The following chronological summary gives an indication of the more
important work that has been done on this phase of the subject:
Tipula oleracea
Ptychoptera, Tipula
Phalacrocera, Ctenophora, Tipula

Frisch1722
De Reaumur
1740
De Gear1776
Schellenberg
1803
Stannius
1829
1832 (Posthumous). Lyonet
Bouche1833
Von Roser
1834
Boie
1838
Dufour
1840
Zeller1842
Bremi-Wolf
1846

Cylindrotoma
Limnobia xanthoptera [=bifasciata]
Ptychoptera
Dicienidia sp., Tipula sp.

Hexatoma nigra
Cylindrotoma
Trichocera

Cylindrotoma
Trichocera, Limnobia

xanthoptera [-bifasciata],

Nephrotoma
Trichocera, Limnophila dispar, Ula, Tanyptera

1849
1854

Perris1867

Nowicki1872'.

Weyenbergh

Dactylolabis wodzicHi
Ctenophora, Dictenidia: anatomy and histology

1873-86

Beling

The immature stages of (.9 species of European

1875-76
1876

Hammond

187(3-1902

De Rossi

1880
1882-1900

Hermann
Mik

Brauer,
Ep:f!;er,Frauenfeld

GrobbenandChionea

Tipulidae, including 20 species of Tipula
Tipula oleracea anatomy
Ptychoptera: anatomy
Liogma glabrata, Tanyptera atrata
Tanyptera atrata
Discobola, Elliptera, Dactylolabis, Tipula rufina,:etc.

Charles Paul Alexander

736
1883

1884
1890
1S93-97
1895 [1898]
1897
1900-09
1901
1901
1901-08

1907-08
1908
1908
1908-09
1908-09
1900
1909
1909
1910
1910

BrauerImportant work on
immature stages

Gerckelihamphidia longiroslris, Tamjptera atrata
I'tychoptera: histology of alimentary canal
Dicranota, Phalacrocera
Pioneer American worker; Bittacomorpha, etc.
Phalacrocera: anatomj'Teucholabis, Aeshnasoma, etc.
Holorusia grandis (as rubiginosa) anatomy

Van Gehuchten
Miall

Hart
Bengtsson
Jolmson
Kellogg
Miiggenberg

Needham
Steinmann
Holmgrentheclassification

of

the

:

Liogma glabrata
Dicranomyia simulans, Pedicia albivitta, Ehaphidolabis, Tipula ultima (as flavicans)
Triogma trisulcata

AnthonPhalacrocera: mouth parts
Ctenophora angustipennis anatomy

Pastejrik
Miiller

Limnobia xatilhoptera, Ctenophora
Triogma trisulcata

BrocherGreeneTipula lunata
Tipula triviitata
Dicranomyia trinotata
Tipula maxima: anatomy
Tipida cunctans (as infuscata)
Ctenophora, Tipula oleracea mouth parts
Trentepohlia pennipes, Tipulodina pedata, and:Thienemann
Brown
Hyslop1911

Vimmer1911-16

De Meijere:

several European species

Trichocera: morphology
Tipula ultima (as flavicans)Anatomy of the respiratory system ofEuropean species of Tipulidae
Trentepohlia bromeliadicola

1912
1913
1913

CaudellGerbig

1913
1914
1914-16
1914-19

PicadoDel Guercio
Topsent
Alexander

1915
1915
1915
1915-17

SwezeyDicranomyia foliocuniculator

LovettCtenophora angustipennis
Dicranota, Phalacrocera, Triogma
Limnobia immalura, Gnophomyia, and, in his
uncertain material, Antocha, Elephantomyia
Tipula paludosa
Cylindroloma splendens

1916-17
1918
1920
1920

KeilinWesenberg-Lund
MallochRennieCameronHudsonmany

Tipida oleracea
Ptychoptera albimana
Dicranoptycha, Ila, Eriocera, Hexatoma, Penthoptera, Liogma, Prionocera, Tipula arctica

Gnophomyia rufa, Limnophila sinistra
Thaumastoptera, Cylindrotominae

Lenz

ECONOMIC IMPORTANCE

The larvae of some species of crane-flies, almost all belonging to thegenera Tipula and Nephrotoma, often do considerable damage to various
crop species, the injury being largely caused by the larvae devouring theroots

and thus Idlling the plants.

Certain of these crane-fly species, as

Tipula oleracea, T. paludosa, and Nephrotoma maculata in Europe, andTipula hicornis and Nephrotoma jerruginea in America, have long been

known, but several others have come into prominence only within theThe Crane-Flies of New York
past ten years, notably— PartTipula parva Lw. (supposition)

T. cunctans and T. simplex

m North America.

II

in

737
Japan, and

The more important out-

which have been recorded are as follows.

l:>reaks of this natureSpecies affecting herbaceous plants

Nephrotoma fcrruginea has been reported by Webster (1891, and 1893 a:
243-245) as injuring young wheat in Indiana, the injury in some cases
being estimated at fifty per cent.

Tipula oleracca is the best-known European species in this group, infesting a wide range of plants.
as

The more important outbreaks of this species
Ewert (1899) reports from Germany anrecorded are as follows:

unusually severe attack, in which the larvae of this species and of Tipula
nigra did a considerable amount of damage to the roots of grasses.
Theywere so abundant that from ten to twenty could be found in the area of

one square foot.

Grasses in the infested area were completely destroyed.

Ormerod (1885, 1886, 1900) records serious injury by T.oleracea to pasturesand meadows in England, and recommends apphcations of guano either
This species ha.s beenalone or mixed with salt, kainit, or superphosphate.
recorded also as being a serious pest in Ireland (Anonj^mous reference,and meadows, and careful plowing and the use of
Schoyen (1903), in Norway, reports
serious injmy to meadows and pastures, as well as to cereals and young
cabbage plants. Del Guercio (1914) records serious injury in the Italian1904 b)in grainfields

fertihzers for its control are advised.

rice fields.

A few additional outbreaks of Tipula oleracea may be mentioned.
Barthou (1913) records injury to canes (raspberry, strawberry, andothers) in France.

Desoil (1914) reports injury tomeadows in France,

and Ritzema Bos (1915) to meadows in Holland.
Tipula paludosa is reported by Lind, Rostrup, and Kolpin Ravn (1914
and 1915) as causing serious injury to oats and barley in Denmark, andRennie (1916 and 1917) reports the same species as injuring corn andpastures in England.

Tipula flavolineata is reported by Sopotzko (1916) as injuring clover
The specific identity, however, is probably erroneous, as thisin Russia.

species lives almost entirely in decaying

wood (Beling, 1873 b: 581-582).

Onuki (1905) records Tipula parva Lw. (supposition) as one of the serious rice pests in Japan, In some localities from sixty to nmety perCharles Paul Alexander

738

cent of the seedlings have })oon destroyedby this species.

The larvae

apparently cannot exist in water for any long period of time, and so may

be killed by flooding the

Guercio (1914)

offersfieldsforfromto thirty-six hours.

six

Del

the same recommendation for the control of T.

oleracea in the Italian rice fields.

Tipida bicornis has been found in grass and clover lands in Illinois byForbes (1890), and on clover in Indiana by Webster (1892 a).
Tipula cundans, as recorded by Hyslop (1910:126, as T. infuscata),
works largely on Japan clover and other leguminous plants. The larvae

many as two hundred having beenfound in an area covering but little more than one square foot of soil.
They destroy the plants by devouring the roots and sucking the juices.
It is recommended that the infested fields be plowed under before theadult flies emerge (in the autumn), and the following season be planted
are often exceedingly numerous, as

to corn or potatoes or else left fallow.

Tipula simplex has been foimd on pasture land and alfalfa in Cali-

by Doane (1908) and by Carnes and Newcomer (1912). Doaneby Hyslop, 1910:120-121) also reports an outbreak of this species
on wheat and grass lands and in clover fields in central California in
1907, and states that thousands of acres of these crops were stripped offornia(citedtheir verdure.

The following summary gives the various species of plants that aredamaged or destroyed by crane-flies in general. Very many of the
injuries reported for
FamilyGramiaeae

Crop

Wheat
Wheat
Wheat

Tipulaoleracea,

however, are omitted.

BarleyTipulidae
Tipula paludosa
Tipula sp
Tipula paludosa

Barlev

Tipula sp

BarleyTipula oleracea
Tipula paludosa

Corn
Corn

Authority

Species

Nephrotoma ferruginea Webster (Indiana), 1891, 1893a
Tipula simplex
Hyslop (California), 1910

Stedman (Missouri), 1902
Rennie (England), 1917
Kirk (New Zealand), 1895
Lind, Rostrup, and Kolpin Ravn
(Denmark), 1915
Wahl and JMiiller (Germany),
1914

Oats
Rice
Rice
Cereals
Cereals

Goriatchkovsky (Russia), 1915
Lind, Rostrup, and Kolpin Ravn
(Denmark), 1914, 1915
Del Guercio (Italy), 1914

Tipula oleracea
Tipula parva
Tipula oleracea

Nephrotoma pratensis.

•

Onuki (Japan), 1905

Anonymous.

reference (Ireland),1904 bHollrung (Germany), 1898

I

The Crane-Flies of New York
Crop

FamilyGramineae

Meadows

•

739

II

Authority

Speciesandpastures

Meadows— PartTipula bicornis

Forbes (Illinois), 1890

Tipula simplex

CarnesandpasturesandNewcomer(Cali-

fornia), 1912

Meadows

andMeadows

andpastures

MeadowsD^soil (France), 1914

Tipula paludosa

Rennie (England), 1917

Tipula sp

MacDougall (Scotland), 1915

Tipula sp

Ritzema Bos (Holland), 1915

Tipula sp

Marchal and Prillieux (France),andpastures

Meadows

Tipula oleracea

andpastures

Meadows(Ireland),1904 bandpastures

Meadows

Anonymous reference

Tipula oleracea

pasturesandpastures

1916

Meadows

andpasturesTipula oleracea

Ormerod (England), 1885, 1883,

Nephrotoma macule ta

Collinge (England), 1911

1900
Tulips (bulbs).

Liliaceae.

Onions and garlic

Cruciferae

Polygonaceae.

.

.

.

Rosaceae

CabbageCabbageCabbageCabbageCabbage
Buckwheat.
Roses
Raspberry

.

.

.

.

.

Nephrotoma lineata.

Alfalfa

Tipula oleracea
Tipula bicornis
Tipula bicornis
Tipula flavolineata
Tipula oleracea
Tipula cunctans
Tipula simplex

Tmbellif erae

Peas

Tipula paludosa

Solanaceae

Compositae

.

.

.

.

.

CloverCloverCloverCloverJapan clover.

I

Vassiliev (Russia), 1915.

.

Schoyen (Norway), 1903
Averin (Russia), 1913
Goriatchkovsky (Russia), 1915
Lutchnik (Russia), 1916
Lutchnik (Russia), 1916
Goriatchkovsky (Russia), 1915
Schaufuss (Germany), 1901

andstrawberry..

Leguminosae.

.

Tipula oleracea
Tipula oleracea
Tipula oleracea
Tipula oleracea
Tipula sp
Nephrotoma sp
Tipula oleracea.

.

Carrots
Potatoes
Potatoes

Tipulidae
Tipula oleracea
Tipula lateralis
Potatoes, stored Trichocera
Potatoes, stored Trichocera
CardoonTvpida sp

Barthou (France), 1913
Webster (Indiana), 1892a
.

.

.

.

Forbes (Illinois), 1890
Sopotzko (Russia), 1916
Del Guercio (Italy), 1914
Hyslop (Tennessee), 1910
Carnes and Newcomer (California), 1912
Lind, Rostrup, ai^d Kolpin Ra\Ti(Denmark), 1914
Lesne (France), 1905
Beling (Germany) 1887
Cameron (Scotland), 1917
Johannsen (Maine), 1910
Carpenter (Ireland), 1912
Lesne (France), 1908,

Species affecting woody plants

Species that injure or destroy living trees or shrubs are not common,and the damage that they doisahnost entirely of a minor nature.

Charles Paul Alexander

740

Ctenophora angusti'pennis is recorded as doing secondary damage to prune

trees
Oregon' (Lovett, 1915). Fuchs (1900) records four species —TipulaT. marginata, Nephrotoma cornicinn
andN. quadrifaria — as injuring young plants
both deciduous and conifin

scripta,[as iridicolor],oferous species. Taschenberg (1880:54) describes Tipula flavolineata andNephrotoma crocata as destroying year-old seedlings of fir and larch.
Matsuniura (1916:466) records the larva of Nephrotoma makiella as
injurious to the mulberry (Morus) in Formosa.

METHODS OF EXPERIMENTAL PROCEDURE
Collecting

The larvae of the larger species of crane-flies, such as those of the genera

may be readily discovered in the field,
but the immature stages of the smaller Tipulidae are not so easily found.
Tipula, Pedicia, and Eriocera,

It is necessary to bring into the laboratory large quantities of the material

which the immature stages are supposed to be living, and there to
examine it with considerable care. In the case of mud or earth, it isinbetter to wash away the finer silt particles and examine the residue.
The writer has found it most convenient to use a Simplex water-net,

placing in it a handful of earth and holding it underneath a running faucet

The mesh of this net is of sufficient diameter to retain any crane-fly larvae
When the fine particles have been removed

except the very young stages.

the residue can be examined in water in white enameled or porcelain
dishes,and the larvae and pupae may be easily detected in the water.

Numerous associated forms of life will be found, and these should be preserved or recorded.

The immature stages of wood-inhabiting species are most easily foundand painstaking examination of the removed
bark and the exposed parts of the tree stump or trunk.
Moss-inhabiting species, such as those of Liogma and many tipuline forms, may be
shaken from their haunts onto a piece of white oilcloth, where they arein the field Joy a patient

easily observed.

Killing and preserving

The larvae and pupae to be studied should be dropped into boiling water
for a brief instant. The larva, on contact with the water, at once expands
to its maximum size, the spiracular disk is spread wide open, and the anal\

The Crane-Flies of New Yorkgills

are completely everted.

cent formalin— Part741

II

The specimens should be placed in 4-per-or, preferably, 70- to 75-per-cent alcohol.

Large, fleshy

larvae, such as those of the Tipulinae, should be slit at the third or theabdominal segment to allow easy penetration of the preservative.
Other notes on preserving material are given by Banks (1909) and by

foiu'thWilhamson(1916).

Study
In the study of the gross material, both the binocular and the com-pound microscope may be used. A special word on the preparationof the head capsule of the larva may be helpful.
In removing the headcapsule from the body it is generally easiest to slit longitudinally the thoracic segments back of the head and pull the capsule thru this incision.
In forms with compact and massive capsules, the thorax and the head may
be snipped off and the head everted after the manner of turning thefinger of a glove.
of caustic

The capsule should bepotash until

should be dissected

allthe fleshy

inaparts have

5-per-centsolution

been removed.

Itthe mouth parts
Then follow the usual processes of washingout so that

rendered distinct for study.

left

all

are isolated

andand dehydration, and the final mounting in Canada balsam.
Various means are available for examining the spiracular disk. The
specimen may be placed in a watch crystal under water and held firmly
in place by a piece of glass, as, for example, a broken microscope slide.
By holding a heavy piece of glass over the anterior end of the body, theposterior end may be bent at a right angle and bolstered in place by twoor three other glass fragments so that the disk is directed straight upwardand its details are easily examined. Other methods, such as embedding
the anterior end of the larva in paraffin in a deep dish and submerging
the body, may be followed.
It must be emphasized that in alcoholicgross material the study should be conducted under alcohol or water.
The measurements of the pupa
abbreviated in the text as d.-s.
(dextro-sinistral) for the width and d.-v. (dorso-ventral) for the depth
are taken opposite the wing pad.

——

Rearing

In order to rear the immature stages of crane-flies successfully thenaturalhaunts should be imitated as closely as possible.

In the caseof wood-inhabiting species, large pieces of the material in which the larvae

CiiARLES Paul Alexander742

arc working may be l^rought into the lal)oratory and placed in the breed-ing cages.

Tliis

method may be followed withspecies of Tanyptera,Ctenophora, and other genera.
Fungicolous species, as a rule, also are easily reared. The entire fungns
in which the specimen is found should be brought into the la])oratoiy

and placed in a jar on a bed of clean sand. The sand takes up the liquidd;
pi-oduced by the disintegration of the fungus and provides a place forpupation. Species of Limnobia, Ula, and other forms are reared in thismanner.
The chances for error in rearing are many.

One must be certain that

there are no other larvae in the breeding jar with the one that is being
reared; else one of these other larvae might transformand emerge first,
and the results would be altogether misleading. The writer has hadtliis happen in his breeding cages, even after the utmost care had beenused to guard against it. Beling, the great German student of theinunature stages of crane-flies, made a few mistakes in the same way;as, forexample, in the case of his Trimicra, the larvae that he describes

being pediciine and probably a species of Dicranota or the yoimg larvas
What happened, presmnably, was that Beling foundof a Tricyphona.
these pediciine larvae and placed them in rearing; in the same cage, bub

unknown to the breeder, was a larva of Trimicra which emerged, andit came from one of the larvae that he hadplaced in rearing. It is usually easy to check up such errors. Thus,
naturally Beling thought

the writer has placed in rearing the larvae of Penthoptera and, to his
surprise,

had adultsof

Limnophila adusta emerge.

Obviously larvaeof L. adusta got into the cage in spite of precautions, and emerged

first.

When

hardclosely related species are concerned, however, it becomes amatter to straighten the tangle. Hence a species cannot be reared too
many times, since each rearing checks up the previous results.
The precaution to be taken in the case of mud-inhabiting or sand-mud or sand is baked or thoroly
Then it may be remoistened,
and the larva or larvae chosen to rear may be put into the earth without
the chance that some unknown larva may be lurking in the medium andmay emerge first, and so bring about confusion.
inhabiting species is to see that thedesiccated in order to destroy all life in it.

The writer has foimdthat the most satisfactory

way to rear small

tipulid larvae fomid in earth or sand is to place a small amoimt of bake I

The Crane-Flies of New Yorkearth from their haunts— Part II743

m a 4- or 6-dram shell vial, moisten the earth,and place a thin layer of moss over it. The vial should not be corked,
but should be covered with a piece of cheesecloth held in place by rubberbands.
In the case of carnivorous species, as the Hexatomini and the
Pediciini, individuals should be isolated in vials, but the herbivorous
The predatory
species may be bred in large numbers in single containers.
forms are usually distinguished without difficulty by their extremely
active, snakelike motions, the other species being

more sluggish in theiractions.
Strictly aquatic forms, such as Antocha, are reared only with great diffi-

These species can survive only in rapidly flowing, well-aerated
water, and it is usually a difficult matter to imitate this condition successfully.
The best plan is to place rearuig cages in the natural haunts ofculty.

the larvae.

hand andThis can be done if a suitable location is sufficiently close at
freefrom disturbance by inquisitive passers-by.

The

chief

source of danger to breeding cages in natural streams is that a sudden
rise of water may suffocate the larvae or wash the cages downstream,
or else may destroy the contents of the cage by a thick deposition of silt.

Lotic species that are discovered in streams far removed from the laboratory

can be transported alive for long distances in folds of wet cheesecloth.

The writer has found this to be the most satisfactory way to keep advanced
pupae alive until they are ready to emerge as adults.
In general, the writer has foimd species of the tribe Pediciini the mostdifficult to rear, and the Limnobiini and the Eriopterini perhaps theeasiest.
Mud-inhabiting species are easily reared, l)ut species from rushingtorrents are at the opposite extreme and it is almost an impossibility tobring some of these species -thru to the adult condition.
TYPES OF THE IMMATURE STAGESThe material on which this study is based was almost entirely reared.
seems desirable that these authentic specimens of the larva and thepupa should be so designated that they will have a value greater thanNo terms for the designation ofthat possessed by ordinary specimens.
types of the immatiu'e stages are known to the writer, and the two thatIt

are needed in this paper are here defined as follows:

Nepionotype, The type of the larva.

Neanotype, The type of the pupa.

Charles Paul Alexander744

The type specimen should, of course, be selected only when there isabsolute certainty of the identification, and in most cases this determination
can be made only hy rearing the species.
(this

After the species has been reared

many times, if possible, in order to check up thea good representative specimen may be chosen as the type of theshould be done

identity),In the cases in which the species is known only from a single
specimen, the nepionotype may be the larval skin, the neanotype thepupal skin. The remaining specimens of the original series become parastage.

types.
The types of the immature stages possess fully the value of thetype of the adult and should be as carefully preserved. The types herein

designated are in the collection of the writer.

They are preserved inalcohol, but the larval heads of most species have been removed, treatedwith caustic potash, and mounted in balsam.

EXTERNAL MORPHOLOGY

The larvae and the pupae of crane-flies show considerable diversity
The fundamental plan of structure remains muchin their general form.

the same thruout the group, but the details are widely different and furnish
the characters in use for the separation of the various tribes and lesser
divisions.

The immature stages of crane-flies have evolved more rapidly thanhave the adult flies, and in many features they show a greater specialization.
The head capsule of the larva seems to be the most constant feature,
the same fimdamental type of structure recurring in the generalized

members of all the various groups, indicating a close phylogenetic relationship.

On the other hand, the respiratory organs of both the larvae andthe pupae vary greatly in the different species and are obviously moldedby habitat. The often-repeated statement that the inside of an organismshows what it is, wliile the outside shows where it has been, is well illustrated here.

The larva
General features

The form of the larval body is, as a rule, moderately elongated autl
The head is eucephalous and non-retractile in, the threeusually terete.

families Tanyderidae, Ptychopteridae, and Rhyphidae.
It is incomplete
and more or less retractile in all the species of Tipulidae. The bodyis shortest in the more generalized forms, becoming greatly elongated— PartThe Crane-Flies of New York

many Eriopterini and some Limnobiini (Dicranoptycha).

in

up

745

II

It is

mads

the composite head capsule, three thoracic segmeiits, and nineofevident alxlominal segments.

In some species all the abdominal segments
narrow basal and a usually broader

are subdivided, respectively, into a

posterior ring, or annulus; in other species only the basal segments are

The integument is usually covered with a dense appressedso subdivided.

jiubescence and often bears setae, or pencils of hairs, or, in some CyUn-drotominae, spinous projections.
Respiration

is

often

in the Rhyphidae it is
The typical metapneustic forms

characteristically metapneustic

amphipneustic, in Antocha apneustic.

;

show vestigial lateral spiracles, but these are not functional in any
known to the writer and the peripneustic type of larva is still

speciesunknov\'n in this group of Diptera.

The spiracles are placed at the endsof the long breathing tubes in the

Tanyderidae and the Ptychopteridae.

In the Tanyderidae, the Tipulidae, and the Rhyphidae the disk is sur-rounded liy a varying number of lobes which are rarely indistinct, theseranging in number from two to eight. Anal gills are found in representatives of almost all the major groups of crane-flies, and their loss is
a result of habitat and non-usage. In wood-inhabiting species the gills
are often modified into blunt lobes, having the evident function of pro-pulsion by shoving.

Body form
As already stated, in the majority of crane-fly larvae the body is terete
or approximately so, but in some species it is decidedly depressed withthe ventral surface flattened.
Such forms are Dactylolabis, some Cjdindrotominae, and some Tipulinae. The integument is produced into
elongate spines and blades in almost all species of Cylindrotominae, similar
conditions being suggested in a few tipulines.

A definite arrangement ofThe basal abdominal ring is provided withcreeping-welt in the Limnobiini and in some Hexatomini

setae (chaetotaxy) obtains.

a transverse

and Pediciini, as well as in a few other forms. In some genera,
as Epiphragma, this welt is practically naked; in others it is covered witha microscopic scurfiness; while instillothers (Dicranota)

it isseparatedwhich are armed with circlets of cliitinizedhooks that lessen in size from the tips basally. The welts are both dorsal
and ventral in position in, many Limnobiini and in some Pediciini (Rhaphi-

into distinct paired prolegs,

Charles Paul Alexander

746

I

dolabina), or are ventral only (Rhamphidaria, Ularia, Epiphragmaria
and most Pediciini). In numljer they range from four (on abdominal
segments 4 to 7) in most Pediciini, to five or six (on abdominal segments 2to 7) in Ularia, Epipliragmaria, and other forms.

Organs of respiration—

The spiracular disk.
The posterior end of the body is usually truncated,bearing the two spiracles and surroimded by a number of fleshy lobes.
These lobes vary much in their shape and armature, and range in number
from two in the Pedicaria and the Antocharia to six or eight in the Tipuhnae.
The inner faces of the lobes are often fined with brown or black horny
plates, which serve as points of attachment for the longitudinal muscles.
Their arrangement and distribution are of great service in specificclassification.

The Limnobaria

usually have the lobes surrounding the spiracular

disk indistinct; the Antocharia have

two long ventral lobes; the Rham-

pliidaria five lobes; the EUipteraria and the

Dicranoptycharia four lobes.
In the Eriopterini, the Elephantomyaria and some Eriopteraria havefour lobes; the majority of species of the latter division havefive.

these last-named species the disk is almost squarely truncated,

and theInfive lobes are subequal and stellate in appearance.

The Hexatomini usually have four lobes, with the ventral pair longer
than the lateral pair. In a few cases only (Ularia, Dicranophragma,
Poecilostola) a reduced mid-dorsal lol)e is present.
In the more specialized
forms the lateral lobes also tend to reduce, simultaneously with the,

In this tribe the disk is fringed with longwhich are sometimes excessively elongate, especially on the ventral
lobes where they appear as long fringes of delicate hairs that spread out
on the surface film of the water as broad fans. Such fringes are foundin a large nimiber of Hexatomini and also in the Adelphomyaria (supposition).
Near the tips of the ventral lol^es, but inside this fringe ofhairs, are often inserted one or more stiff sensory bristles.
In the Pediciini, the Pedicaria have two lobes and the Adelphomyaria(supposition) have four.
In the former division the lobes are ventral
in position and are almost naked.
The Cylindrotominae have six rather indistinct lobes in Cylindrotoma,
and four in the other genera.
elongation of the ventral lobes.
hairs,

The Crane-Flies of New York — Part II

747

The Tipulinae show six lo])es in practically all genera, the only exceptions
being that there are eight in a few rare cases of Tipula and five in Dolichopeza, and that lobes are indistinct or lacking in Tanyptera.

In the Rhyphidae (Trichocerinae) the spiraciilar disk is very similar
In theto that in many Tipulidae, and is surrounded by four lobes.

Ptychopteridae the very reduced disk is borne at the tip of a slender,
In the Tanyderidae the condition is somewhat
retractile breathing tube.
similar, but here the disk is larger and is surrounded by five lobes at the
tip of a long, stout, non-retractile breathing tube.

The spiracles vary greatly in size from very large to small and vestigial,
They consist
or they may even be lacking in some species of Antocha.
foldedradiallybysurroundedapiece
middleof an apparently imiform
margin, or ring, of various widths, called the stigmal ring.

Many authors(De Meijere, Mik, Miiggenberg, Brown, Keilin, and others) hold thatthe middle piece is an imperforate chitinized plate and that respiration
Gerbig (1913), however, shows thattakes pl?.ce thru the stigmal ring.
the middle pieceissplit acrossoverlapping membranes.

the disk, the cleft being closed

Directlybehindthespiracles

theby twotracheae

enlarge into the felt chamber, whose walls are provided with long, branched,
Surrounding
treelike structures, the branches apparently anastomosing.

chamber in many larvae are dense masses of air tubes, whichmake up the tracheal lungs. These tubes are arranged in bundles, which
the

felt

arise in special cavities of the felt

there are about

fifty

chamber; thus, in Tipula paludosa,

bimdles, each of about twenty tubules, maldng

a total of one thousand of these air canals (Gerbig).
The early stages of the larva are quite different from the later developmental stages, as Gerbig (1913:137-140), working on Tipula paludosa,

has well shown.

The prominent six-lobed spiracular disk of the morematured larva

represented in the

is

first

developmental stage by fourwhich bear but few bristles on their outernot evident, but are replaced by eight
lobes
are
The
dorsalmargin.
from one another. The spiracles
equidistantabout
branched bristles,
larva, and project a little beyond
grown
the
circularasinare oval, notheavily chitinized projections,the level of the disk.

Thewriter has noted several first-stage larvae

with an appearance almost as described l)ut showing several points of
dilTerence.
The immature larvae of Phalacrocera are described elsewherein thiswork (page 963).

Charles Paul Alexander

748—

Anal gills.
In the Ptychopteridae and in the Tanyderidae there isbut a single pair of anal tracheal gills. In the former these are very small
and are cylindrical, in the latter they are large, branched, fanlike strucIn the Tipulidae the anal gills number from four in most Lim-tures.

The increase in number is brought
about by a division of the original gill of each side. In generalized forms,
as Antocha, Pedicia, and others, the anal gills are constricted into segments,
nobiinae to six or eight in the Tipulinae.

the apical ones being more or less telescopic into the preceding ones.

Asa rule the anal gills are entirely or almost entirely retractile within thebody.

In the Tipulinae the gills vary with the genera, being blunt and constructed for propulsion in the wood-inhabiting species, such as those of

Ctenophora, Tanyptera, Tipula, and other genera.

In the semi-aquatic

species of Tipula the number of gills varies from four to eight.

In thelatter case each of the four principal gills is deeply bifid and the gills arearranged transversely, as in T. ignohilis', in species with six gills the posterior
branches of the posterior gills are atropliied as a rule; in other species,

which have but four gills, the four anterior branches are preserved, theIn Longurio and "Aeshnasoma

posterior pair being usually atrophied.

the four anal
branches.

gills

are pinnately branched, each with aljout six lateral

A similar condition is found in the Tanyderidae.

The headThe

primitive

crane-fly

head, was undoubtedly of the eucephalous,

non-retractile type, as found in the Ptychopteridae, the Rhyphidae,

other families.

and

The retracted head capsule of the Tipulidae is a derived

condition.

The massive, compact capsule is fomid in all of the lower groups of thethree subfamilies of the Tipulidae.

Such a head is easily derivable fromthe condition in the Rhyphidae or in the Tanyderidae, for instance.
dorsalinThe

median sclerite, the jprefrons, is almost as large and conspicuous

Ula as in the eucephalous families.

The lateral plates that constitutethe capsule are shaped like a mussel in the generalized groups, with theposteriorincisionsshallow.

In the rAore specialized forms,

with thecapsule greatly dissected, the two plates of either side are entirely sepa-— the innermost, next to the prefrons, being theinternal-lateralof De Meijere, and the outermost being his external-lateral.

The prefrons

ratedThe Crane-Flies of New York— Part II

749

found in all forms with the capsule compact and massive, and is evenmany species with the dissected capsule. In many cases,
however, it is firmly fused with the internal-lateral plates; or, in someispreserved incases, all the plates of the capsule are firmly miited and their individual

are ill-defined.
In the generalized forms the clypeus and thelabrum are entirely distinct from each other, while in other forms the twolose their individuality by fusion.
The sclerites of the generalized primitive head bear bristles, or setae,
of various types and arrangement.
In the Tanyderidae the punctures
are multisetose; in most Ptychopteridae the setae are branched or somewhat plumose.
The literature on the head capsule and the mouth parts is as follows:
limits

Ptychoptera. De Meijere, 1910:188-191, figs. 14-20.
Bittacomorpha. Hart, 1S98 [1895]: 192, pi. 5.
Trichocera.
De Meijere, 1916: 191-194, figs. 21-23.
Limnobia. De Meijere, 1916: 198-201.
Dicranomyia. De Meijere, 1916: 197-198, figs. 32-35.
Dieranoptycha. Alexander, 1919b; 71, figs. 2, 3.
Elliptera.
Mik, 1886 b: 339, pi. 6.
Alexander, 1915 a: 5-6, pi. 1.
Ula.
Dactvlolabis.
Mik, 1894:261-266, pi. 2; Nowicki, 1867:341 (as Rhicnoptila).
Limnophila. De Meijere, 1916:204-206, figs. 49-51.
Eriocera. Alexander and Lloyd, 1914:21-22, pi. 1.
Hexatoma. Alexander, 1915c: 146, pi. 1.
Penthoptcra. Alexander, 1915c: 155, pi. 1.
Tricyphona. De Meijere, 1916: 195-196, figs. 29-31.
Pedicia.
Needham, 1903:286, fig. 19.
Dicranota. Miall, 1893:237-238, pi. 10.
Ormosia. De Meijere, 1916:201-204, figs. 37-47.
Gnophomyia. Malloch, 1915-17 b: 230-231, pi. 34, fig. 10.
Chionea. Brauer, Egger, and Frauenfeld, 1854:614, pi. 11.
Phalacrocera. Miall and Shelf ord, 1897:344-345; Bengtsson, 1897.
Cylindrotoma. Cameron, 1918.
Liogma. Miiggenberg, 1901; Alexander, 1914:111, pi. 1.
Ctenophora. Anthon, 1908:544.
Prionocera. Malloch, 1915-17 b: 199, pi. 32, figs. 1-3.
Holorusia.
Comstock and Kellogg, 1904:55, 61-62.
Tipula. Vimmer, 1906, and 1911:1-6.
In addition to the preceding, the genera Helobia (Beling, Hart, Malloch), Rhaphidolabis(Needham), and Tanyptera have been discussed less fully.

All of the above-named genera are considered in this paper, and thehead capsule and mouth parts of the following genera are describedProtoplasa (supposition), Bittacomorphella, Rhipidia, Rhamphidia,
Antocha, Epiphragma, Pseudohnmophila, Pilaria, Ulomorpha, Elephant-also:oniyia,Teucholabis,

Gonomyia,

Erioptera,

JVIolophilus,

Adelphom^^ia

750Charles Paul Alexander

feuppo >t.on)

RhapMolabma,

Oropeza, Longurio, Aeshna^oma
Thee«-np„o„. for the go.icra are either
entirely new or else thecharactershave heretofore been n.sufficiently
described a„<l figured.

:lw

The

detai

'" '"'' "''"'" "' "^"'^ «*>' ^--'^ of crane!fl£ :;':Wn.

The labrumand the epipharym;.The labrum is preserved in all cranefly brvae
It ,s usually broadly transverseor ovaThe <lomsXe.

clothed w.th short hairs, thesebeing longer on.sor papillae

Thelateral regions

ZeZ
Z Th "V'"""^""'""-

thea.>tror marrin

on the ventral, or epipharyngeal,ide''' '''''*"^"'' P™P^^ i/vari^sTy arnm the different groups of crane-flies.
T-Ae

Za6f»„.-The labial region of the capsuleisdof vast importance inr""P-'^ shouldr"
The confusion in terminology of thedipterous larval labium
has been partly outlined by
De Meijere (1910:253). The
appieciated.
Sed"''•''"^•"™"'"'

princh alpiincipai

synonyms are as follows:
Submentum.

(BSTniT luafpS'tT' '"""'• "' '"'"'" "" <M«-rt); s„h„.e„tu^ (Miall); ectol.bium

BoSST "'"'""'"" *'""• <"='°"''"™

<K»"i"); -<iol«biu„, (Holm^en,Vi„,„,er«!,P«pfer,„x-Labi™ (Meinert); n,e„,um (Miall. i„

The submentum is represented by
eacephalous groups of crane-flies,

mentum IS the usually chitinized

"

''

Tf

DicranoU.); endolahium (Keilin).

a narrowtransvei-se strip inthebeing well shown in Ptychoptera
anterior-ventral plate of^"*"™''*-"'*'''"'°"'

The

he capsule'-"ty ™™>"*eteeth 'tBi::
teeth.
In Bittacomorpha it is moreor less distinctly bilobed but
untoo hed. In the generalized
Tipulidae it is indistiifctly div
ded
an outer plate and, imme<liately

isLo

dorsad of this and closely applie

to or

tooth of theside.

mentum and m some cases an additional
tooth on either
The inner plate adds a varying numberof teeth to the mental
p ate, from two mEpiphragma to as many as ten or twelve in .some
speciesof Limnobia.
The chitinized plate is completely
divided me<lial y insome Enopterini (Molophilus an,l some
Erioptera' in them some Hexatomini (Pseudoliinnophila).
In

PeS'and

this

case,

each

half

is

The Crane-Flies of New York— Partdirectly continuous with the ventral bars of thementumrepresented

head capsule.

by a narrow,

Lininophi'laria

thechitinizcd

l)ar,

which

mentum islacking in chitinized parts, or apparently so, in

teriniisdelicately

is

751

II

grooved with parallel

In thetransverse,The

lines.

many Eriop-

and in the higher Hexatomini.

Directly behind the mentum is the prementum, rarely prominent andbearing the two small palpi when they are preserved, as in the Ptychopter-

The hypopharynx lies immediately dorsad of the prementum.

In
an enlarged, two-horned, fleshy lobe which
is difficidt to study.
In the Tipulidae the generalized condition, suchoccursinLimnobiini
and the lower Hexatomini, consists of twoasthecollar-like chitinized plates whose anterior margins bear from ten to
fifteen or more teeth.
In the Tipulini this plate is narrow, with theanterior margin few-toothed, there usually being five or in some cases
idae.

the Ptychopteridae thissixteeth.

Eriopterini

In theit is

The antennae.

is

Pediciinithehypopharynx

islabriform;

in

many

cushion-shaped and covered with numerous blunt setae.

— The antennae are borne on the anterior

lateral margin

They are one- or two-segmented,of the capsule, laterad of the labrum.

and in most cases have one or more apical papillae.

De Meijere andothers hold that the principal apical papilla constitutes an additional

In some cases, however, as in Ula for example, twomore papillae are found which are very similar to one anotherexcept for shght degrees of size. It is difficult in such cases to hold any
one of these papillae as a true segment. The writer prefers to consider
them as apical sensory papillae, and they are so treated in this paper.
The primitive antennae are not greatly elongated, and are usuallyantennal segment.
or evenshort-cylindricalor

subglobular (Ula).

An auditory plate, circular inouthne and lying on the face of the basal segment, is almost always present.
The apical papilla in. the Linmophilaria and the Eriopterini is egg-shaped,or oval, and is delicately sculptured by a network of api:>arently impressedlines.

In many species, as for example those of the higher Hexatomini,

In many Limnomany species of Tipula and related

the apical papillae are very long, tapering to the tips.
biini,

as for example Limnobia, and informs, the apical papilla is button-shaped, or diskhke, and is often very
reduced.

—The 7nandihle.
In the primitive type, such as occurs in the eucephalousgroups and in the generalized Tipulidae, the mandible is rather com-

Charles Paul Alexander

752

plicated, usually having a considerable number of teeth on the cuttingedge and a distinct sub-l)asal arm (prostheca) or prosthecal tuft of hairs.

At the same time there are usually one or more setae on the outside of
the mandible, near the " heel," or base, of the scrobal region. In thegcncrahzed tipulid type there are two apical teeth and two rows of cutting
teeth.

In other cases, as in the Limnobiini, there are but a single dorsal

tooth, a single apical tooth,and a varying number of cutting teeth on

the ventral face; in some species of Limnol)ia the number of teeth in theventral cutting row is six or seven.

In the Tipulini the mandible is often
reduced in size (as in Tanyptera), there being but a single dorsal anda single ventral tooth in addition to the apical point; the prostheca,
however,isusually well developed.

In the Eriopterini but one row ofteeth, the ventral, is strongly developed.

In the Pediciini and the higher
Hexatomini, the mandible is elongate and sickle-shaped, with the few
teeth on the cutting edge at about midlength.
In Ulomorpha the mandibleis hinged at about midlength, the basal part being hollowed out on the
inner face to receive the blade in a position of rest.

—

The maxillae.
The maxillae are usually of simple form. In thegenerahzed types, such as the eucephalous families and the lower groupsof the subfamilies of Tipulidae, they consist of a large and distinct trans-versely triangular cardo, a conspicuous stipes,

outer apical lobes.

anddistinct innerandIn the PecHciini the lobes are separate in the supposed

Adelphomyia larva Ijut are fused together in the highest types (Dicranot'a)
In the Hexatomini the lobes are reduced to a single long, flattened blade.
The outer and inner lobes are usually densely hairy, especially at the tip
and on the inner face of the latter. The palpus is uni-articulate andantenniform in the generalized forms, is short-cylindrical with a varying

number of tiny sense pegs at the apex, and bears on its face a circular
auditory plate similar to that of the antenna. The armature of the inner
lobes is varied, in some species being provided with stiif, comblike setae,

or projections.

The pupa
General features

The pupa of the crane-fly is of the so-called "free," or " munmiy,"type.

In only a few cases does the larval skin adhere to the end of theabdomen (Dactylolabis, Cyhndrotoma, and some other forms.) The headisusually small,

and

is

ventral in position.

Iimnediately behind theThe Crane-Flies of New York— Part II

753

head on the pronotiim are the two breathing horns, which are variously
developed in the different tribes and genera and furnish invaluable bases
The leg sheaths usually far exceed the short or but
for classification.
moderately elongate wing sheaths. The abdominal segments are often
provided with a suliterminal armature of stiff setae, or spines. At theposterior end of the body, the last two segments (cauda) are variously
modified to inclose the sexual organs of the adult flies.

The head
The head is usually small and flattened, occupying the anterior ventral
part of the body. The eyes differ in size in the various groups, in some
(Erioptera, Elephantomyia) being larger in the male than in thefemale; in the male sex they are approximated on the niedian line above
or beneath.

The front between the eyes is usually narrowed behind, dehmited bycompound eye, narrowed at the posterior end,

the iimer margin of theand bluntly rounded or pointed at the apex.
herein as the labrum, or labral sheath.

and the labrum of the imago.

This partisdescribed

It contains the fronto-clypeus

At its tip it bears two more or less diver-gent lobes, these being in some cases closely approximated so as to appear
as a single lobe; these are herein termed the labial lobes or sheaths, and they

contain the so-called paraglossae of the adult fly.

On either side of the labral sheath, and usually divergent and lying
along the posterior margin of its face, are the sheaths of the maxillary

which in most cases extend beyond the knee joint of the fore legs.
all the Limnobiinae these are short and stout and almost
straight, but in the majority of the tipuline forms they are curved attheir tips, which in most species are actually recurved and offer an easy
palpi,In almostof distinguishing members of this subfamily.
In many of the
Limnobiini the margins of the cheeks project as flattened ledges overlyingmeans

the joint of the fore legs.

The antennae arise from above or between the eyes and bend laterad
and thence caudad around the eyes, in some forms, such as Elephantomyia and the males of other species, lying across the face of the eyes.
The antennae usually end just beyond the roots of the wings, but in themales of some species (Eriocera) they extend beyond the tips of the wings.
The basal segments (scape) of the antennal sheath are often armed with3

Charles Paul Alexander754

spines or tubercles, and very often the organ is angulated at the end ofeach segment of the atUilt antenna inside.
At its vertex, between or just dorsad of the antennal bases, the headmay bear a crest which is usually bilobed and setiferous. In some species
this cephalic crestisquadrituberculate, there being a smaller secondary

crest behind or before the primary one.

In the Tipulini the crest is very
In most of the Limnobiini

inconspicuous and but weakly setiferous.
it islacking or nearly so.

The head may be variously armed with spines, tubercles,or setae; in

Eriocera spinosa, for example, there are spines or strong tubercles on theantennal scape, on the clypeal region, and even on the face of the eye.
In some cases there are setae on the front between the eyes, on theclypeus,and on the cheek.

The thorax
The pronotum of the thorax is small. The ventral part is closely
applied to the head and often has small setiferous tubercles close to thebreathing horns.

The pronotal breathing horns are variously developedin the different tribes and genera,

and are discussed here in general terms

only.

Many species are propneustic, the pronotal horns alone being functional.
Other species (in Hexatomini and Eriopterini) are peripneustic, the second
to the seventh abdominal segments being provided with functional lateralspiracles in addition to the breathing horns; other pupae have lateralabdominal spiracles, but in most cases they are merely vestigial. Somepupae are amphipneustic, there being in addition to the breathing horns
a conspicuous pair of spiracles on the dorsum of the eighth abdominal
segment (Rhamphidia, Ula, Epiphragma; in the typical species of Limnobia these are present but they are small and are probably nonfunctional).
In the Ptychoptei'idae the breathing horns are very unequally developed,
one being enormously elongated and filiform while the other is abortive.
In some Tipulini (Longiirio, Prionocera, Tipulodina) the horns are likewise greatly elongated, but in these cases they are shorter than the

body and are approximately subequal

in

size,

or at least are not so

disproportionately unequal.

In the Limnobiini the breathing horns are usually stout and broad,
Limnobaria (Limnobia, Dicranomyia) being subquadrate,in the typicalThe Crane-Flies of New York
subcircular, or even broader than long.

— Part II

755

In Elliptera they are earhke andIn Antocha a unique condition is found, in that the apparatus

very large.

is divided into eight stout filaments.

In the Eriopterini and the Hexatomini

the horns are usually long and cylindrical or sUghtly flattened, and straight

There are numerous deviations from type, how(alexanderi andkansensis, for example), the horns are trumpet-shaped; in Gonomyia sulphurella they are compressed and flattened into fanlike structures.
Inseveral widely separated paludicolous genera, such as Ulomorpha, Pseudolimnophila, and Prionocera, the breathing horns are split at their tips
into two conspicuous flaps with fimbriate margins, an obvious adaptation
Often the breathing horns are very small (as in
to an existence in mud.
Elephantomyia, Teucholabis, and Trichocera) or even microscopic(Dicranoptycha). Limnophila hyalipennis (Zett.) is described by Beling(1886:198-199) as lacking the breathing horns, but this is presumably anerror of observation which may be due to a defect in the material studied,or possibly the horns are sessile or reduced as in Dicranoptycha.
Insome species (Erioptera, Eriocera) the horns are stout at the base but
or but slightly curved.

ever.

In

Gnophomyia and some species of Gonomyia

taper rapidly to the acute points.
In many PecUciini the tips of thehorns are expanded and usually obliquely trimcated, with a row ofbreathing pores around the margin. The tipuline breathing horns are
remarkably uniform in general structure, being usually elongated cylindricalin shape, of moderate length, and with the tips slightly expanded.
The
most conspicuous deviations from type are the short, flattened, and somewhat clavate horns of Tanyptera.

The thoracic mesonotum is very convex, or gibbous in many species(Limnobiini and some Hexatomini).

In other species it is declivitous,
with a high transverse crest (Eriopterini). This crest may be armed
with numerous small spines (Helobia); from two to four powerful hooks(Molophilus, some species of Ormosia, some species of Erioptera, Teucho-

from six to eight tubercles set with stiff bristles (Gonomyia); or
abundant small setiferous tubercles on either side of the median line,
labis);

these

being

less

conspicuous

Ormosia and of Erioptera).

along

the

shoulder(most

In Eriocera longicornis thereisspeciesofa medianspine or tubercle on the scutellum.
In some Cylindrotominae themetanotiun bears spines. The extreme lateral or ventral margin is
produced into a blunt or somewhat pointed angle just above the wingCharles Paul Alexander756
root; this is usuallyarmed with from one to three setae.

Various othersetae usually occur on the mesonotum.

The leg sheaths lie along the ventral side of the pupa.
various groups, a considerable difference inand thetheThere is, in thelength ofthe sheaths,

arrangement of the ends of the tarsal
segments. In the Bittacomorphinae, the fore legs overlie the middle
In the Rhyphidae the fore legs overlie the middle pair, which,legs.
In the Ptychopterinae and in the Tipulidae,
in turn, overlie the hind legs.
all three pairs of sheaths lie parallel along the midventral area, thoseof the hind legs being outermost along the wing sheaths, and those of
the fore legs being on the inside and contiguous. In Gnophomyia theleg sheaths are very short, extending but little beyond the wing tips andreaching only the end of the second abdominal segment. In other craneflies they are longer, in some species reaching the end of the fifth abdominal
segment. Very often the tips of the tarsi are on a level. In the Limnobiini the hind tarsi are usually a little shorter, and the fore tarsi are a little
In the Eriopterini
longer, so that the ends of the sheaths form a bhmt V.
the middle legs are usually (except in Gnophomyia) much shorter thanthe other legs; in Helobia, moreover, the hind legs are conspicuously
longer than the fore legs, so that all three sets end at different levels.
A somewhat similar and parallel development is foimd in the tipuline
their position,

relativeforms.

The wing sheaths are moderately broad, usually ending oppositeora little beyond the apex of the second abdominal segment. The venationof the various species often shows clearly and with little distortion on
the wing pad, and in such cases it is of very great help in locating certain
Species with a heavy pattern in the adult wings, such- as thosegenera.

Epiphragma, show this pattern on the wing
pad in the older pupae, and this helps to confirm their identity.
of the genera Discobola and

The abdomen

The abdomen consists of apparently nine segments, the last two being
approximated or fused to form the cauda. The segments arecloselyusually plainly divided into a broad dorsal (tergal) and ventral (sternal)

and a much narrower and more restricted lateral (pleural) area.
The segments beyond the first are variously subdivided into rings, or
annuli, by false constrictions, there being usually one, but in some cases
region,The Crane-Flies of New York— Part757

II

two or more, basal rings and a much broader posterior ring. The segments
In the Limnobiini and someare variously armed with setae, or spines.
Hexatomini these occur on the basal annulus, and consist of a transverse
welt which is margined (as in Antocha and EUiptera) or covered withabimdant microscopic chitinized points.
In some Pediciini the setae
occupy both the basal and the posterior rings of the intermediate abdominal
segments.
In the Eriopterini and most Hexatomini, as well as in thetipuline forms, the setae occur on the posterior ring, just before the caudal

margin, and consist of a single transverse row of small spines.

In theCylindrotominae the segments are often armed with prominent elongate
body projections (Liogma, Triogma). In Phalacrocera elongate lobes
In the Ptyare confined to the posterior segments of the abdomen.
chopteridae the segments are densely covered with transverse rows oflong tubercles which are replaced by chitinized plates

margin of the segments.

near the posterior

In Bittacomorpha these tubercles are stelkite

at their tips.

The dorsum of the cauda in most crane-flies is armed with four, five,which are often spinous at or near their tips.
These represent the lobes that surround the spiracular disk of the larva,
and their number generally corresponds to the larval condition. Thus,or six conspicuous lobes

in the Ptychopteridae there is a single lobe, in the Pediciini there are two,and in the Hexatomini and the Eriopterini there are four or five. In thethere are often but foiu- evident lobes, the dorsal pairof the larva not being represented.
In some generahzed groups, as
the Limnobaria, the Rhamphidaria, and others as indicated elsewhere in this paper, the larval spiracles persist in a more or less functional
Tipulinae

condition.

Abdominal spiracles.

— In the Eriopterini and the Hexatomini thereisa distinct and apparently functional spiracle on each pleurite of the second
to the seventh abdominal segments, located near the base of the posterior

In the Eriopterini these spiracles are small and, in the cast pupal
appear tubular or elongate. Smaller vestigial spiracles are evident
in most genera of the Tipulidae.
As has been mentioned earlier in this paper, in certain primitive genera
ring.

skin,

of Tipulidae— such as Rhamphidia, Ula, Epiphragma, and to a
— there aratherextent those of Limnobariaon the dorsum of the eighth segment.

ispair oflesserlarge spiracles

In Dactylolabis cubitalis the lateral

Charles Paul Alexander

758

abdominal spiracles are protuberant, notably the pair on the second
segment.

—

The genitalia.
The genital sheaths conceal the adult organs within.
In the male the valves are usually subequal or the sternal valves arealittlethe longer.

In the female the sheaths, or acidothecae, of theovipositor usually have the dorsal valves considera]:)ly longer than thesternal valves and distinctly upturned at their tips.
In the few groups
with fleshy ovipositors, the female cauda is very difficult to distinguishfrom the cauda of the male. The valves are variously armed with spines

or lobes, at or before the tips.

internal morphology

The internal morphology and anatomy of the crane-fly larva are here
The two genera that have been studiedin greatest detail by the writer are Eriocera and Antocha.
The most
considered in general terms only.

important literature on the various systems of the body is cited in ordertomake reference to it more easily available.
The digestive system

The most important literature on the digestive system is as follows
Ptychoptera. Grobben, 1876; Van Gehuchten, 1890.
Chionea. Brauer, Egger, and Frauenfeld, 1854:613-614, pi. 11, fig. 4.
Dicranota. Miall, 1893:242-245.
Phalacrocera.
Miall and Shelford, 1897:347-351; Bengtsson, 1897.
Ctenophora. Anthon, 1908:546-551, figs. 12-35.
Holorusia. Kellogg, 1901 a; Comstock and Kellogg, 1904:56-57.

Herbivorous larvae (Tipula, Holorusia, Ctenophora, Phalacrocera)
In the larva of herbivorous species the alimentary canal extends theby the coiled,
The esophagus is slender, opening

length of the body as an ahuost straight tube inclosed

perforated sheets of adipose tissue.
into the hypopharynx.

The proven triculus has a large esophageal invagiThe ventriculus has at its anterior end four

nation at its anterior end.

elongated ventricular caeca, these being of two distinct lengths in pairs(didynamous) in Ctenophora, and all four alike in Holorusia.

In Cteno-phora there arc four small, pocket-like gastric caeca near the posterior endof the ventriculus, which are not mentioned as occurring in Holorusia.

At the junctui'e of the ventriculus and the small intestine are the fourThe Crane-Flies of New York
coiled malpighian tubules.

— Part II

759

Behind the^ventriculus lies the small intestine,with a caliber much smaller than that of the ventriculus. At the unionof the large and the small intestine is a prominent diverticulum of very-

Ctenophora three times as large as in Holorusia). The
form the rectum.
Phalacrocera has the ventriculus without caeca and the hind intestine

large size

(inlarge intestine gradually dilates behind towithout a diverticulum, altho it is decidedly a herbivorous genus. The
In Phalacrocera, and probably
condition in Chionea is almost the same.
in most other larvae, a peritrophic membrane (a secretion of the epithelium

which incases the food and keeps it from direct contact with the epithelium)ispresent.

There are four malpighian tubules in all the Tipulidae that have beenthoroly studied (Chionea, Eriocera, Dicranota, Phalacrocera, Ctenophora,

In the Rhyphidae, likewise, there
and adult stages. In the Ptychop-

Holorusia, Tipula, and a few others).
are four tubules in both the larval
teridae,however, there are five tubules, as in the Culicidae, the PsyThis might indicate some basis for placing

chodidae, and related families.

the Ptychopteridae in close proximity to the Culicidae, as has been doneby Lameere(1906).

The salivary glands consist of two large coiled tubules which pass intoa collecting duct and unite to form the common collecting duct that opensat the base of the hypopharynx.

Carnivorous larvae (Dicranota, Eriocera)

The ahmentary canal in the carnivorous type of crane-flies is a short,straight tube, with a minimum of appendages such as caeca

In Eriocera the esophagus

ticula.

isand diver-very long, and the ventriculus is

The malpighian tubules are yellow at their
The intestine is shortorigin, soon passing into an orange-brown color.
short

and

and withoutstraight,

caeca.

without a diverticulum.

The salivary glands in the newly killed larva of Eriocera spinosa arevery conspicuous by their opalescent color, which persists for a day ormore after death. These glands consist of two elongate-oval or cylindricalglands, of the opalescent color just mentioned.

From the anterior endof each of these glands passes the long, slender, salivary collecting duct,

which unites with its fellow to form a conmion duct opening at the mouth
From the caudal end of each gland is a flattened, almost hyahne,

cavity.

760Charles Paul Alexanderelongate lobe whichisindistinctly

pseudo-segmented, in

its

posteriorpart being in intimate connection with the abundant fat tissue.

Respiration and circulation

The most important literature on the respiratory and circulatory organsisas follows:
Ptychoptera. Grobben, 1876.
Dicranota. Miall, 1893:245-248.
Phalacrocera.
Aliall and Shelford, 1897:351-356.
Liogma. Miiggenberg, 1901.
Ctenophora. Anthon, 1008:551-554.
Kellogg, 1901 a; Comstock and Kellogg, 1904:57-58, 00.
Holorusia.
Tipula maxima. Brown, 1910.
Tipulinae, Hexatomini (especially Tipula paludosa and Limnophila punctata)

Gerbig,.

1913.

General articles.

De Meijere, 1895; Viallanes, 1880.

The most important works on the structure of the tracheal system withspecial reference to the mechanics of the spiracles, are those by Miiggen-berg (1901), De Meijere (1895, 1902, 1916), Brown (1910), and Gerbig
Miall and Shelford (1897:351-356) discuss in much detail the(1913).
structure of the heart in Phalacrocera.

In Eriocera spinosa, which may be considered as typical for this groupof crane-flies, there are two principal tracheal trunks which lie in a dorsal

positionand run almost the lengthof the body.

They are connectedacross by a very delicate, unbranched, simple, tracheal commissure,send ofT branches laterally to supply the various organs of the body.

and
Near

the posterior end of the body they are approximated, and just in front of
the spiracles they expand into the tracheal chamber.

Directly cephaladchamber the first lateral branch passes off, numbering from theposterior end forward.
Branches 2 to 8 are large and much forked.
Just after leaving the main tracheal trunks, each of these sends off aventral branch which supplies the alimentary canal and the fat tissue.
Branch 3 supplies the region of the malpighian tubules; branch 4, theof thisposterior part of the stomach; branch 5, the anterior part of the stomach;branches 6 to8,

the esophageal region— branch 6 supplying the pro-

ventriculus, branch 7 and part of branch 8 the esophagus, and the remainderof branch 8 the pharyngeal region.

The mainpart of each lateral branch continues laterad, at(anterior) fork sending a

its

first

branch forward to the next branch, so as toThe Crane-Flies of New York— PartII

761

form a complete but much-branched lateral trachea. The branches arevery diffuse and abundant and the skin is well supplied. As alreadystated, the cross-commissures are very delicate and are unbranched ornearly so, the dorsal regions of the body being supplied by branches fromthe lateral supply.

The condition of the tracheae in Antocha, the only apneustic crane-fly
larva among the species here considered, may be outlined as follows:
The usual two principal trunks are present, joined across on eachsegment by delicate cross-commissures which send off two approximated
Laterad and ventrad of the
These are joined to
midwaybetween the
branches
insertedabout
the main trunks by finelying nearer
posterior
end
of
the
body
towardthedorsal commissures,
branches except on the eighth segment.

principal trunks are the delicate lateral tracheae.

the posterior commissure than the anterior one.

At the ninth segment each trunk sends off a branch from its proximalside, these branches being connected by a long commissure and supplying
The commissure is about as long as that partthe posterior pair of gills.
Immediately caudad of, orof the branch between it and the trunk.
just at, the fork, but

the anterior
finally

gills;

on the ventral side, a subcqual branch passes intoat thesame point the delicate lateral tracheal trunk
Caudad of this triple forking the mainends in the main trunk.

trunk gradually widens out into a cylindrical structure which is truncatedapically, at the tip passing out into

two small elongate branches, near

the base with about three delicate branches, two being dorsal in positionand one more lateral.
lo])es,All of these tracheae, to the gills and to the caudal

send off many branched capillaries at frequent intervals, and thecaudal lobes unquestionably function as tracheal gills.

A comparison of Antocha with Dicranota as described and figured
by Miall (1893:245-248) shows, in the latter, distinct spiracles and thegills

similar but much smaller.

The tracheal arrangement differs in thata single branch on either side supplies both gills of that side, while thecaudal lobes are tracheated l)y a branch that leaves the main trunk close
to the spiracles. (Plate XII, 2

and 3.)

The arrangement of the tracheae at the base of the wing pad is describedand figured for Bittacomorpha by Dr. Chapman in Comstock's The
Wings of Insects (1918:36-37).

Charles Paul Alexander

762

The nervous system

The most important literature on the nervous system is as follows:
Ptychoptera. Grobben, 1876.
Dicranota. Miall, 1893:241.
Phalacrocora.
Miall and Shelford, 1897:350; Bengtsson, 1897.
Ctenophora. Anthon, 1908:556-557.
Holorusia.
Comstock and Kellogg, 1904:58-59.

In Holorusia the brain, or supra-esophageal ganglion, is composed of
two principal lobes united posteriorly and lying above the esophagus.
Beneath the brain and on the under side of the esophagus lies the subesophageal ganglion, which is connected with the anterior end of thebrain lobe by the circum-esophageal commissures.
The above-named
organs form a complete ring, or collar, around the alimentary canal. The
ventral nervous system leads backward from the subesophageal ganghon
on the ventral surface of the body wall. In the thorax there are four
closely approximated ganglia representing the thoracic and the firstabdominal segments. Beyond these and located in the abdomen are
six abdominal ganglia.
The ganglia send off four large nerve trunks.
The condition is similar in Ctenophora and in Tipula, there being seven
abdominal gangha, of which the first is located in the thorax and the last
six in the abdomen.
In Phalacrocera there are eight abdominal ganglia,
the first being usually found in the first abdominal segment and theeighth in segment 10. The head ganglia are all independent of one
another, not fused into a complex.
The longitudinal commissures between
the first five ventral ganglia are double, and those between the eight
abdominal ganglia are simple (Bengtsson). In Dicranota there are likewise eight abdominal ganglia, the first being located in the first al^dominal
segment and the others in segments 5 to 10, there being two close together
in the tenth segment.

The reproductive system

The most important literature on the reproductive system is as follows:
Ptychoptera. Grobben, 1876.
Dicranota. Miall, 1893:248.
Phalacrocera.
Miall and Shelford, 1897:356-357.
Holorusia. Comstock and Kellogg, 1904 58.
:

The reproductive organs appeiar early in the larval development.

In

Dicranota the testes lie in the ninth segment and form elongate capsules,

The Crane-Flies of New York— Part II

763

when sufficiently advanced giving off the vasa deferentia from the innerEven in larvae not fully matured, Miall foundside of the hinder end.
ripe

No division of the testes
spermatazoa may

spermatazoa mixed with spermatoblasts.

into follicles

isapparent.

In Phalacrocera,also, ripe

be found in the older larvae.
In Phalacrocera the ovaries form a pair of cylindrical bodies tapering
to each end, lying on either side of the beginning of the intestine.

Theovaries are very peculiar in structure, seeming to be adapted to the almost

simultaneous discharge of all the eggs.

The muscular system

The most important literature on the muscular system is as follows:
Body musculation
Ptychoptera. Grobben, 1876.
Dicranota. Miail, 1893:241.
Ctenophora. Anthon, 1908:.545.
Holorusia.
Comstock and Kellogg, 1904:59-60.

Musculation of the mouth parts
Phalacrocera.

Bengtsson, 1897.

The fat bodies, or adipose tissue

The most important literature on the adipose tissue is as follows:
Dicranota.

M^all, 1893:241-242.

Anthon, 1908:554.

Ctenophora.
Holorusia.

Comstock and Kellogg, 1904 56.
:

The imaginal disks, or histoblasts

The most important literature on the imaginal disks is as follows:
Phalacrocera.

Ctenophora.
Holorusia.

Miall and Shelford, 1897:357-358.

Anthon, 1908:555.
Kellogg, 1901 b; Comstock and Kellogg, 1904:61.

PHYLOGENY
The

origincrane-flies

adult

flies

which

in

is

and phylogenetic development
stillof thelargely problematical.

most generalizedof the various groups of

The evidence supphed by theliving groups points toan ancestor

many respects resembled the recent Mecoptera, or scorpion

764Charles Paul Alexander

That this resemblance is presumably not fundamental is suggestedby the immature stages of these same generalized forms, which show
flies.

but few features that can be reconciled with those of a panorpid-like
Unfortunately the fossil record helps but little. At the begin-ancestor.

ning of the Tertiaries, crane-flies were already numerous both in species
and in individuals, but in most cases they are clearly referable to existing
It is evident, then, that the investigator must go still further
genera.
back for his evidence, and the fossil crane-flies of the Upper Mesozoic

The phylogenetic developare, unfortunately, still largely unavailable.
ment of the group may be considered from two standpoints, the geologicalrecord and comparative morphology.
THE GEOLOGICAL RECORD

The most important works on

fossilcrane-flies

are those of

Loew(1850), Scudder (1894), Handlirsch (1906-08, and 1910, a and b), Meunier(1906), and Cockerell (1910, 1913, 1916, and 1917, a and b).

The writerisindebted to Professor Cockerell for assistance in determining the ageofmany of the fossil-bearing beds.

Excellent accounts of the various

deposits may be found in Cockerell (1913) and in Tillyard (1917).

The Tipuloidea, representing the superfamily of cratie-flies, is herein
as being constituted of six families, of which two, theEoptychopteridae and the Architipulidae, did not survive the Mesozoic
period.
The Eoptychopteridae are known only from the Mecklenburg

consideredTheknown from the same formation and include eightspecies arranged in three genera.
The other four families contain recent
Lias (lowest Jurassic), and include but three monotypic genera.

Architipulidae areforms and are included in this paper.

The North American Eocene and Miocene, as represented by the WhiteRiver and Green River beds and the Florissant shales, respectively, give
evidence of having had a northern fauna, especially in the Eocene. This
shown by the great development of the Cylindrotominae, which
White River basin almost dominated the crane-fly fauna during
the Eocene. It seems probable, moreover, that the group was forced
is

well

in theinto colder regions of the globe during the Oligocene,

when the tropicalNo group of crane-flies that can beconsidered as being tropical has yet been found in the Florissant.
On theelement reached far to the north.

other hand, the European Oligocene, as shown by the Gurnet Bay beds

The Crane-Flies of New York— Part II

765

and the lower Oligocene Baltic amber, has a considerable tropical element
as, for instance, Styringomyia, found in both formations,and Trentepohlia, Polymera, and Brachypremna, the last two being
amber records. In connection with these two last-named genera, asfossils known only from the Baltic amber, it should be noted that each
is found living to-day only in America, where they are confined to theapparent—

austral and tropical regions.

The Tanyderidae are represented by the Oligocene Macrochile (amber)and Etoptychoptera. At present this is still a very small group, including
but two genera and nine species.
The Ptychopteridae include a Ptychoptera from the Bohemian Krottensee (probably upper Oligocene) and a Bittacomorpha from theFlorissant.

The Rhyphidae (Trichocerinae) are represented by two species fromthe Baltic amber and one or two additional species from the Miocene.
The Tipulidae may be considered in general terms only. But one
In
tipulid is hsted by Handlirsch as being known from the Mesozoic.
the earliest Tertiaries, however, a variety of species is found in this family

which almost parallels their recent development.
The Limnobiini first appeared in the Eocene (Dicranomyia, Geranomyia). In the lower Oligocene, numerous species of Dicranomyia andDicranomyia was common, tho
probably not so rich in species as at present. Rhamphidia is found inamber, and Antocha has been described from the Florissant Miocene by
Scudder; the latter record, however, seems very doubtful to the writer,
judging from Scudder's figure and description.
The Hexatomini are represented by Ula, Limnophila, Eriocera, andPolymera in amber, and by Limnophila in the Florissant shales.
There is no record of the Pediciini being preserved as fossils.
The Eriopterini are well represented in the Baltic amber by Erioptera,
Gnophomyia, Gonomyia, and related genera, and also in the Miocene by
Gonomyia, Cladura, and others. Toxorhina, an apparent relative of
Rhipidia occurred.

In the Miocene,

Elephantomyia, occurs in the Baltic amber.

The Styringomyiini are represented by the only living genus, St5n-ingomyia, a fly of uncertain affinities, in the Gurnet Bay Oligocene (Cockerell,
1917 c and 1919) and in amber, reappearing in the Pleistocene African
copal.

766

Charles Paul Alexander

The dominance of the Cylindrotominae (genus Cyttaromyia) in the
Eocene and Miocene of the North American fauna has already been
mentioned. No records of this group from the European Oligocene
The recent species of the subfamily are practically all
forms belonging to cold, temperate regions, the few Oriental species of
Stibadocera coming from mountains at considerable altitudes.
The Tipulinae have been found as far back as the Mesozoic, but therecords are not entirely satisfactory. In the lowermost Tertiaries, however, undoubted tipuline forms occur.
Species occur in the Green River
shales of Colorado (Eocene).
The group was well represented in the
Oligocene (Baltic amber, Tulameen beds of British Columbia, Krottensee,
are available.

and Gurnet Bay), and was very common in the Miocene (Radoboj, andespecially in the Florissant of Colorado, where some twenty-five speciesof Tipula and closely allied genera or subgenera have been described by
Scudder and Cockerell).
Tipulidae of the Pleistocene are not numerous, only a few having been

made known from the refuse of lake dwellings in England(Dicaera,

apparently related to Ctenophora), and in the African copal, including
such genera as Styringomyia, Elephantomyia, and Toxorhina.

comparative morphology

The morphology of the various stages of crane-flies has been detailed
elsewhere in this paper and need not be repeated here.

PHYLOGENETIC' CONSIDERATIONS

The cucephalous familes of crane-flies are undoubtedly lower, phylothan the Tipulidae, and the latter have been derived fromThe generalized type recurs in all three subfamilies of the
Tipulidae, and it is uncertain which of these three should be placed
lowermost. Presumably all three groups arose from an immediate common ancestor, or the Tipulinae and the Limnobiinae arose from one point of
the tree, the Cylindrotominae developing from the Umnobiine stem at asomewhat later period. The accompanying phylogenetic tree (Plate XII, 4)

genetically,

the former.

The Limnoshow but little deviation from the fundamental type. From thelevel of the lowermost Hexatomini (Ula and Epiphragma), in close

graphically illustrates this apparent evolution of the group.
l)iiniproximity to the Limnobiini, the remaining groups of crane-flies can be

The Crane-Flies of New York— Part II

767—

the Pediciini and the Eriopterini on the one hand, the hexatomine divisions on the other. The highest levels of development of theirrespective types are apparently Diotrepha (Limnobiini), Dicranota

derivedGonomyia (Eriopterini), and Hexatoma (Hexatomini).
The immature stages of the Tipulidae of the antipodal regions (Australia
and Chile) are entirely unknown, and their discovery may bring to light(Pediciini),

striking novelties thatThe

may well upset the present ideas of classification.

largest remaining gaps in the present

knowledge of the Nearctic

fauna relate to the genera Atarba, Toxorhina, Cryptolabis, Phyllolabis,and Megistocera, but it is not expected that any major groups will haveto be created for their reception.

WITH KEYS AND DESCRIPTIONS OF
THE SPECIESLIFE HISTORIES OF CRANE-FLIES,

Various classifications and arrangements of Diptera have been proposed
and the principal evolutionary changes in the entomologist's

in the past,

conception of classification are herein indicated.
The groupings in the superfamily Tipuloidea have been very diverse.

Brauer (1883) separated the eucephalous famiUes from the Tipulidae(Polyneura).
Osten Sacken (1893) placed the Tipulidae in the Netnocera
Coquillett (in Howard,vera, the Rhyphidae in the Nemocera anomala.
1912:85-86) included the Tipulidae in his superfamily Tipuloidea, but
separated the Rhyphidae, placing the latter in Bibionoidea. Lameere (1906)
gave a classification that had little basis in fact, the Rhyphidae being
considered by him as Brachycera and the Ptychopteridae being placedwith the Culicidae. Knab (1915), working on the respiratory systemsof the larvae, demonstrated the hitherto unnatural groupings of theof the Nemocera, and arranged the crane-fly famihes, alongwith a few others, in the group Polyneura, a group coordinate in rankwith the previous conception of the Nematocera. The latest grouping,that of Malloch (1915-17 b), was based on a broad knowledge of both thefamilies

adult flies and the immature stages; and here, for the first time, one getsthe inclusion of thethe true conception of the superfamily hmits—

Ptychopteridae, the Tipulidae, and the Rhyphidae in a single major group.
Malloch's arrangement is herein adopted, with the following exceptions:(1)

the Limnobiidae are not held as constituting a separate family, but

areunitedwith theTipuUdae;(2)

the family Tanyderidae has been

Charles Paul Alexander768

erected, to include the generalized

Tanyderina which have hitherto beenplaced with the Ptychopteridae.

The immature stages of the four families of the Tipuloidea are readily
The larvae of the Tipulidae can be confused only with thoseseparable.

of certain low brachyccrous forms, as, for example, the Leptidae.

In theBrachycera the mandibles work vertically and parallel to each other;in the Nematocera, including the Tipuloidea, they operate transversely
or obliquely against the teeth of the mentura and the hypopharynx.

The presence of fleshy lobes surrounding the spiracular disk is a character
The larvae of a few groups of brachycpossessed by almost all Tipulidae.
erous Diptera, such as certain Leptidae, Sciomyzidae, and other families,
possess entirely similar caudal lobes but are readily recognizedby thesmall and very reduced head capsule.

The eucephalous famiUes of the Tipuloidea may be distinguished bymeans of the characters indicated in the following keys:
Larvae1.

2
Body eucephalous, head non-retractile; amphipneustic or metapneustic
Tipulidae (p. 791)
Head incomplete behind, retractile; not amphipneustic

2.

Caudal end of body prolonged into a slender breathing tube; metapneustic
Caudal end of body not prolonged into a breathing tube; amphipneustic.

3.

Breathing tube stouter, non-retractile; gills large, pinnately branched; punctures of head
Tanyderidae, supp. (p. 769)
multisetose; found in wet decaying wood
Breathing tube slender, completely or partly retractile; gills slender, cylindrical,
unbranched; punctures of head with simple or plumose hairs; found in wet earth.

3

Rhyphidae (p. 787)

Ptychoptebidae (p. 772)
Pv-paeL One of the pronotal breathing horns greatly elongated, much longer than the body, the(Family Ptychopteridae, p. 772)
other breathing horn very short, abortive.
2
Breathing horns short, or, if elongated (some Tipulinae), not longer than the body3
and the difference in size not so apparent
Ptychopterinae (p. 773)
2. Tarsal sheaths lying side by side, parallel
The fore tarsal sheaths overlying the middle pair, the four middle and hind tarsi the
Bittacomorphinae (p. 779)
longest, parallel
Rhyphidae (p. 787)
3. Tarsal sheaths overlying one another in pairs
Tarsal sheaths lying side by side
Tipulidae (p. 791)

The pupae of the Tanyderidae are still unknown.
Itispossiblethat Bittacomorphella

horns; in this case this genus would run

by the arrangement
couplet 2.

(page 779) has short breathing

down to couplet 3 above, butof the tarsal sheaths

itruns out as indicated in

I

The Crane-Flies of New York— Part II

769

Family Tanyderidae
Larva (supposition).'— Body eucephalous, metapneustic.

Integument smooth, shiny white.

Last two segments of abdomen produced into a stout, non-retractile breathing tube, whichisdistinctly five-lobed at the tip.

Tracheal gills two, very large, pinnately branched.

Headwith scattered punctures, which are multisetose; lateral plates of head united across venter.
Mandibles opposed, narrow, tridentate. Maxilla with palpus two-segmented, the outer

Labrum small, semi-circular, with two punctures; clypeus withlobe digitiform, pubescent.
four punctures.

Pupa.

— Unknown.

The Tanyderidae comprise a group of primitive crane-flies including
but three recent genera, with ten species. Of these the genus hereinconsidered, Protoplasa, with three known species, is found in the Northern

Hemisphere.

All

that

is

known concerningthebiology of supposed

species of this group pertains to Protoplasa fitchii and is discussed below.

Genus Protoplasa Osten Sacken (Gr. first + to form)
1859
1878

Protoplasa O. S.
Idioplasta O. S.

Proc. Acad. Nat. Sci. Phila.,Cat. Dipt. N. Amer., p. 222.

p. 252.

The genus Protoplasa is confined to temperate 1^5'orth America.
are three known species, of which P. vipio 0. S.
are v/estern in their distribution, while

There
and P. vanduzeei Alex,P. fitchii O. S.

iseastern.

The

general characters of the supposed larva are given above; the detailed

account in connection with the species P. fitchii follows.
Protoplasa fitchii 0. S.
1859 Protoplasa fitchii 0. S.

Proc. Acad. Nat. Sci. Phila., p. 252.

The remarkable insect Protoplasa fitchii is oneof the rarest of the

There are scarcely a score of specimens extant in thevarious collections of the country, most of which are from the mountainous
section of North Carolina. The fly has not been reared, but the writer has

local crane-flies.

in his possession a larva that he refers with much confidence to this species.

It is one of themost remarkal)le dipterous larvae that have ever come to

the writer's notice, and, whether or not it belongs to Protoplasa, it should
certainly be called to the attention of entomologists in the hope that

may some day be bred and its identity confirmed or ascertained.

it

These

peculiar dipterous larvae were discovered by H. S. Barber, C. T. Greene, and770Charles Paul Alexander

R. C. Shannon, on the Potomac River near the mouthof Dead Run, Fairfax County, Virginia. They were found during thelatter part of May, 1916,
in a much-decayed maple log, where they
were associated with the 'larvaeof the syrphid fly, Temnostoma bombylans (Fabr.)(Barber, 1913).
Greeneattempted to rear the larvae procured at this time, but
did not succeed.
In May, 1917, he sent the writer one of thepreserved larvae for study.
Later Dr. Viereck was interested in the matter, andon May 27 he procured

one fine, healthy larva, which was sent to the writer
It arrived safely on the 29th, and was at onceat Ithaca, New York.

placed in rearing.

nately this larva died the day after it came into

Unfortu-

the writer's possession,and this remarkable insect still remains unreared.

Associated with thisdecaying maple in which it was shipped,
werelarvae of Temnostoma and a larva and a pupaof the tipulid Epipiiramna
larva in thepieces ofsolatrix.

Theevidences that this larva

It belongs,

is that of Protoplasa arenumerous
without doubt, to the Nematocera, and theimmature stagesof all the remaining families of thatdivision have been made known
Many features in this larva point strongly to the condition in otherfamiliesof crane-flies.
The eucephalous condition of the head, and the elongate
breathing tube, are suggestive of thePtychopteridae; the five-lobed
spiracular disk, the anal tracheal gills, themetapneustic respiratory

and other features, are very similar to conditions
in certain
Tipuhdae. However, there are conditions obtaining
here that are foundnowhere else m the Diptera, so far as is known tosystem,the writer, such as the
combination of a eucephalous head of primitive
organization, a stout
non-retraetile breathing tube, the large,
pinnately branched anal. gills'
the multisetose punctures on the head,
and the details of structure of themouth parts.

The multisetose punctures are suggestive of the branchedor plumose hairs of Ptychoptera, and givea possible hint of the origin
and ancestry of this condition in the latter group.
The writer cannot
but regard it as suggestive that the larvae arefound in direct association
with those of Epiphragma in saturated decayingwood.

He has mentioned

in other papers (Alexander, 1910:254,
and 1919 d: 883, 915) the remarkablesuperficial resemblance that exists between
the adult flies of Epiphragma
fasapenms and those of Protoplasa, both forms
havingsomely banded wings of a pattern not found
fauna.

very hand-in other species in the local

The Crane-Flies of New York— Part771

II

The only observations on the adult flies that the writer has ever been
make were in reference to five specimens taken in New York

able toState in 1909 (Alexander, 1910:254).

These were swept from rich vege-

tation along the banks of the Sacandaga River in northeastern New York.

The adult flics have never been recorded from the vicinity of Washington,
but are regional, since the species ranges thruout the eastern UnitedStates as far south as Georgia (in the mountains).
Larva.

—Total length, 11.8 mm.
Length excluding breathing tube, 5.6-6 mm.
Length of breathing tube alone, 5.8-6.2 mm.
Length of ninth segment of abdomen (chitinized part of breathing tube), -3.4-3.6 mm.
Length of anal gills, 2-2.1 mm.
Diameter of body, 2.3 mm.
^

Coloration pure white, head and apical two-thirds of breathing tube pale brown, chitinized.

In the living larva there is a transparent, subcircular area on the dorsum of segment 7

thru which the beating of the heart can be clearly seen, but in preserved specimens this areaishard to perceive.

(Plate XIII, 5 )Body eucephalous, entirely smooth, shiny, dorsum of segments 2 to 9 each with a transverse group of tiny points.

Thorax with the first segment longer than the succeeding two

segments taken together, narrowed anteriorly, broader behind; mesothoracic segment about
half as long as the first and about twice as long as the very narrow metathorax.

Abdominal

segments gradually increasing in length to the seventh; segments 8 and 9 abruptly narrowed
into a stout, elongate breathing tube which is about equal in length to remainder of body;basal, or proximal, part of tube (segment 8) shorter

than terminal part (segment 9) andnot chitinized; segment 9 elongate, chitinized, with a deep transverse constriction a shortdistance before tip; this apical part,and the region

just before the constriction, smooth,the remainder of the last segment with numerous doHcate transverse wrinkles.
disk (Plate XIII, 6) surroundedby five

lobes,

one dorsal, twolateral,Spiracularand two ventral

in position; dorsal lobe the smallest, lateral lobes the broadest; lobes tipped withsharp,

chitinized spines, v/hich are continued for a short distance down the outer face of each lobe;
fringes of long hairs along margins of lobes; disk with a brownish tinge around spiracles
and an elongate-oval median mark between, and slightly below, the spiracular level; innerface of ventral lobes with indistinct, tiny, brown punctures.
Spiracles large, separated bya distance about equal to diameter of one. Anal tracheal gills two, one on either side of
the base of segment 8, very large and pinnately branched, there being about fifteen branches

to each gill; each of these branches constricted into four or five lobules; lobes at their tips

broadly obtuse (Plate XIII, 9).

Head short and broad, epicranium

chitinized.

Across ventral

face,

the sclerite firmly

united by a narrow transverse band; median part densely punctulate, this area delimited
laterally and posteriorly by an indistinct curved line.

and the clyptus distinct from the labrum.

On dorsum, the prefrons conspicuous

Head and mouth parts provided with numerous

setiferous punctures, each of which is multisetose (from five to eight setae to each puncture).

Chaetota.xy of epicranium (Plate XIII, 7) with three lateral groups of long

bristles,theCharles Paul Alexander

772

posterior group being the longest and most dorsal in position; on ventral face, four punctureson either side (Plate XIII, 8); prefrontal sclerite apparently lacking setae, but on the epicranium
proper, along margin of prefrons, a longitudinal group of six punctures on either side, theanterior pair the longest.
stiff

hairs;

dorsalLabrum semicircular in outline, outer margin fringed with long,
two transverse punctures.

surface with

with four setiferous punctures in a transverse row.

Clypeus narrowly transverse,

Dorsal median part of epicranium

forming a subquadrate lobe whose lateral angles bear brushes of hairs; laterad of these angles

a rounded hollo-w overlapped by a flat, bilobed operculum(thismay be some modificationof the antennae which should occupy about this position on the head).

Mandible elongate,strongly chitinized, tridentate, the apical tooth the longest and more acute than the others,

the middle tooth bluntly obtuse; inner posterior angle of mandible produced proximad

beneath outer lobe of maxilla into a bluntly rounded lobe; mandible with a group of curved
setae on scrobe near base,and a group of longer

setae on dorsal face of mandible.

bristles lying distad

and mesad of these

Maxilla with base, in a position of rest, concealed beneathhead-chitin; palpus stout, antenniform, two-segmented, the basal segment short and stout,

the apical segment much shorter and narrower, with a small, lateral, sensory papilla; outer
lobe of maxilla produced cephalad as a stout, digitiform lobe which is densely hairy.

Family Ptychopteridae—

Body eucephalous, metapneustic, long and slender, the caudal end prolonged
more or less completely retractile breathing tube bearing the spiracles at the tip.
Lobes surrounding spiracular disk indistinct. Anal gills two, elongate-cylindrical,
unbranched. Integument with tiny hairs (Ptychoptera), or with shght warty proLarva.

into atuberances (Bittacomorpha), or with conspicuous elongate tubercles (Bittacomorphella).
Pseudopods on abdominal segments 1 to 3 each bearing a curved claw. Head complete;eyespots distinct.

IMandibles opposed.

IMentum many-toothed (Ptychopterinae) or merely

bilobed (Bittacomorphinae).

— Usually with one of the two pronotal breathing horns greatly elongated, much

Pupa.'

body (this may not be true in Bittacomorphella); in Ptychoptera andBittacomorpha it is the right horn that is elongated, the left being degenerated; in Bittacomorphella the right horn is degenerated. Tarsal sheaths all parallel in the Ptychopterinae,
longer than thethe fore pair overlying the middle pair in the Bittacomorphinae.
setiferous tubercles arranged in transverse rows
in longitudinalrows on pleurites.

Abdomen covered withon tergites and stcrnites, and more or lessCauda with a powerful dorsal median lobe near base of

segment 8.

The family Ptychopteridae includes three genera falling in two tribes:
Ptychopterinae including the single genus Ptychoptera, and the
Bittacomorphinae including the " phantom crane-flies," Bittacomorphatheand Bittacomorphella.
inconnection

The habits of these species are discussed below
The following keys separate

with the various genera.

the subfamiUes of the Ptychopteridae:

Vhe Crane-Flies of New York — Part II773

LarvaeMentum wath outer margin finely serrated; mandibles with three large outer teeth; pseudopods small; coloration yellow or brown

Ptychopterinae (p. 773)

Mentum bilobed, not toothed; mandibles with a single large outer tooth; pseudopods promiiient,each with a conspicuous curved claw; coloration rusty red or black.

Bittacomorphinae (p. 779)

Pupae

M

with
branched
Fore tarsi lying above middle tarsi; wing pads withAll tarsi lying parallel; wing pads

M unbranched.

Ptychopterinae fp. 773)

Bittacomorphinae (p. 779)

The immature stages of this group of flies have long been known, having
Reaumur (1740), Lyonet (1832), and other early
workers.
The immature stages of the common American species
The unknown
Ptychoptera rufocincta are herein recorded for the first time.
Ptychoptera described by Malloch (1915-17b: 240-241) is evidently P.
been investigated byand is here considered as that species.

lenis O. S.

The most important literature on

the Ptychopteridae

Ptychoptera paludosa
Ptychoptera paludosa
Ptychoptera lacustris
Ptychoptera contaminata

Larva, pupa, general.
General

Ptychoptera albiniana
Ptychoptera alhimana
Ptychoptera sp. {lenis supp.)
Ptychoptera sp

GeneralLarva, pupa, general.

Ptychoptera sp
Ptychoptera sp

Larva, general
Larva, general

Ptychoptera sp
Bittacomorpha clavipes
Bittacomorpha clavipes.
Bittacomorpha clavipes
Bittacomorpha clavipes
Bittacomorpha clavipes

GeneralLarva, pupa, general.
GeneralGeneralLarva, pupa
GeneralVan der Wulp

Larva

as follows:

Beling, 1886: 171-172.
Van Gehuchten, 1890. (Histology of the alimentary canal.)

Larva, general

Cameron, 1917:65.
.

Larva
General(1857),

isReaumur, 1740, pi. 6.
Wesenberg-Lund, 1915:348-351.

.

.

Topsent, 1914-16.
Malloch, 1915-17b:240-24L
Leger and Duboscq, 1909. (Protozoan parasites.)
Grohben, 1876. (Morphology.)
De Meijere, 1916: 188-191, figs.
14-20. ( External morphology.
Lyonet, 1832.
Hart, 1898 [1895]: 189-195.
Needham and Betten, 1901:574.

Howard, 1912:9.5-96.
Malloch, 1 9 15-17 b: 239-240.

Weston and Turner, 1917:53.

Miall (1895), Griinberg (1910), and Brunetti(1912) also give general accoimts of the immature stages of Ptychoptera,for the most part taken from the earlier works cited above.

SUBFAMILY Ptychopterinac
Genus Ptychoptera Meigen (Gr. fold + wing)
1800
1803
1856

Liriope Meig.
Nouv. Class. Mouch., p. 14 {nomen nudum).
Ptychoptera Meig.
Illiger's Mag., vol. 2, p. 262.
Ctenoceria Rond.
Dipt. Ital. Prodr., vol. 1, p. 187.

Charles Paul Alexander

774
Larva.

— Head oval to subpyriform, broadest behind, narrowed

patterned above; setae of head branched or plumose.

infront,

not heavily

Submenturn large, usually but a little

narrowed posteriorly, the anterior lateral angles slightly produced; mentum broader thanPrementum with palpi rather small,long, outer margin with numerous teeth (18-22).
lying parallel, densely hairy on outer, or ventral, face.
Mandible with three powerful outerteeth

and aboutsix

smaller inner teeth.

Maxilla with the cardines elongate triangular,

with three setiferous punctures; maxillary palpi antenniform, cylindrical.
transverse, with dense tufts of hair beneath.

Labrumbroad,

Integument almost smooth, without prominent

warty elevations. Pseudopods low, with small claws.
Head .small, with a reduced cephalic crest. Sheaths of maxillary palpi elongate,
Pupa.
the tips projecting around knee joints of fore legs. Leg sheaths with all the tarsi parallel,—

not overlapping one another.

Wing sheaths with media branched.

Pronotal breathinghorns very unequal, one (usually the right) exceedingly elongate, longer than remainderof body, the other tiny, degenerate.

Abdomen set with transverse and longitudinal rowsof small setiferous tubercles on the segments.

Ptychoptera is a small genus including sixteen described species, almost
occurring in the Holarctic region, with a few species in India andFormosa. Two species are found in tropical Africa, and another, anallundescribed form, in southern South America; hence the genus is probalilyfound thruout the world in countries in the temperate zones, with theprobable exception of Australia.
The literature on the immature stages of this genus is summarizedunder the family account.
Topsent (1914-16) has furnished the best account of any species of
His observations refer to Ptychoptera albimana.
the genus Ptychoptera.
The following notes are extracted from Topsent's paper:
The eggs range in number from .520 to 587, averaging 554. They are pale yellow, slightly
arcuated, the surface curiously ornamented, the dimensions being 0.825 by 0.264 millimeters.
The duration of the egg stage is seven days. The newly hatched larva measures 3.85 milliThe adult larva measures 77 millimeters,
meters, the respiratory tube 0.99 millimeters.
The growth of the larva is rapid. Whenof which the respiratory tube is 20 millimeters.
When sixty-seven days old,
fifteen days old it measures, when extended, 25 millimeters.
The pupal stage is from ten to twelve days, averaging eleven
it measures 45 millimeters.
There are normally two generations a year.
days.

De Meijere (1916:188-191) gives a critical account of the structureof the larva of a European Ptychoptera.

It agrees well with the description

most conspicuous differences shown by
De Meijere's species being as follows: labium with the submenturn having
the sides straighter; mentum with the anterior margin evenly, but gently,of P. rufocinda given below, theconvex, with only eighteen evident teeth.

The details of the maxilla,

The Crane-Flies of New York
especially of the setae on the— Part775

II

are not the same.

cardines,

the generic characters given above include

all

However,the species to which thewriter has been able to refer.

Malloch (1915-17b: 240-241) describes and figures an American Ptychoptera which is presumably P. lenis 0. S.

forms in most respects.

It agrees

The author indicatesclosely with the otherthementum as having

median part being projected beyond the level ofMalloch's
the lateral parts and having more teeth (eight instead of six).
specimens were taken in the Yellowstone Park, Wyoming, in Augu.st,three distinct parts, the1890.
to theFrom the locality data it is almost certain that the larvae belongcommon Ptychoptera lenis of the western United States.

Tonnoir (1919) describes a curious sexual dimorphismin thegenus

Ptychoptera.
Ptychoptera nifocincta O.' S.
1859

The

Ptychoptera rufocinda 0. S.

species

Ptychoptera

and among open

Proc. Acad. Nat. Sci. Phila., p. 252.

nifocincta

The

is

common around wet

swaleson the wing from
May to early July, and again in late August and early September; they
may be double-brooded. In appearance they are conspicuously unlike
shrubl^cry.

adult

flies

are

their relatives of the Bittacomorphinae, rather resembling certain large

fungus gnats (Mycetophilidae).

The larvae live in situations similar to those frequented by Bittaco-—morpha, and often occur in the same associations
with larvae of
Limnophila (Lasiomastix) macrocera, Pilaria tenuipes, P. recondita, andErioptera chlorophylla, as well as with many Chironomidae, leeches,
nematode worms, and mollusks. In appearance they are strikingly
different from the larvae of Bittacomorpha, being pale, white or yellowish
white, with the body almost smooth, not covered with the dense transverse rows of setiferous tubercles found in Bittacomorpha, and with thepseudopods on the basal abdominal segments poorly developed. The
larvae of this species are smaller than those of Bittacomorpha, with thehead proportionately much smaller. In structural details, however,they are rather similar to the larvae of Bittacomorpha. The larvae of
Ptychoptera nifocincta feed on the decaying vegetable matter in theirhaunts.
When fully grown, the pupa forms within the last larval skin,and the elongate pupal spiracle is coiled about the mesothorax, as describedCharles Paul Alexander

776

for Biitacomorpha clavipes (Hart,

1898 [1895]: 191).

Careful breedings of1913 at Orono, Maine, placed the indoor pupal life at fourandays and eighteen hours, and that in nature at probably five days
this species in—

unusually short pupal duration.
Larva.

— Length, when

fully grown, 30-32 mm.; when fully extended, about 35 mm.
Diameter, 2-2.2 mm.

Head light reddish brown, not marked with darker spots as in Bittacomorpha; bodytomentum short, pale; seventh and eighth segments of abdomen,

whitish or pale yellow;

and breathing tube, light brown.

Body almost smooth, sparsely clothed with short, appressed hairs arranged in indistinct
transverse rows, on intermediate segments of body there being about twenty of these rows;

body not at all tuberculate, as in Bittacomorpha.

Prothoracic segment short, not so longas mesothorax; metathorax nearly as long as preceding two segments combined.

(In olderlarvae thsct are about to pupate, the right pupal breathing horn may be seen coiled under-

neath the skin of the mesothorax.) First five abdominal segments swollen posteriorly intoa ring that completely surrounds the segments; first three abdominal segments with low,
indistinct

pseudopods on either side of the median line, each with a small, recurved claw;

pseudopods after first pair more widely separated than, and not so well developed as, in
Abdominal segments 4 and 5 more elongate, swollen posteriorly
the Bittacomorphinae.
but not bearing pseudopods; segment 6 narrowed behind, with a few scattered, outspreadinghairs; segment 7 narrower than preceding and telescopic within itself, a.t about two-fifths
the length there being a transverse row of long hairs marking the limit of telescoping; when
fully extended,segment 7 a little longer than segment S; segment S a little narrower, andsegment 7. (The parts of segments 7 and 8 which are exposed intelescopic basally into

the retracted condition are brown and subchitinized, and bear scattered, outspread hairs

which are most numerous near the caudal end of the exposed part and here form transverse
rows; similar rows of sparse, setiferous punctures are on the dorsum of the swellings on thefirst five

abdominal segments.)

about one-fourth

Breathing tube (segment 9) retractile into segment 8 forwhencompletely concealing

Tracheal

gills

two, elongate-cylindrical, situated near base of segment 9; in normal position of rest,

gills

its

length;retracted,

gills.

usually projecting about one-half their length beyond end of segment S.

Apex of breathingtube truncated but without well-defined lobes.

Head small, proportionately much smaller than in Bittacomorpha;

oval tosomewhat

pear-shaped; narrow anteriorly, broadened behind, near posterior margin abruptly narrowed.

Prefrons bread, conspicuous.

(As noted by most earlier writers on the genus Ptj'choptera, thehead bears numerous punctures with setae which are plumoso or have a branched appearance
The writer has examined numerous specimens under high magnification[Plate XV, 20].

and is incUned to believe rather that in some cases several bristles arise from a single puncture
and are closely approximated basally, altho free distally, and that the plumose appearance
apparent.
In other cases, however, actual basal fusion has taken place. The
number of free tips from a puncture varies from three to six, five and six being common
numbers.) Labrum broad, transverse, on disk two large setiferous punctures bearing
is here merely

branched setae; on either side beneath, conspicuous tufts of long hairs, these tufts continued

The Crane-Fues of New York— Part777

II

obliquely proximad to near median line, where there is a median lobe densely covered with

Epipharynx small, subquadrate, margin almost transverse, with about sevenlong hairs.

Labium (Plate
XIV, 12) with submentum much narrowed on basal half, sides subparallel, thence expandedblunt teeth; sides of organ with long hairs which are directed backward.

so that cephalic end is about twice as wide as caudal end; anterior lateral margins produced
into blunt projections; mentum (Plate XV, I7j broadly subquadrate, anterior margin nearlytransverse and with from twenty to twenty-two teeth forming an irregular comb. (There isconsiderable variation in the shape of this comb and in the form of the individual teeth comprising it; usually the median third of the mentumparts, but this condition

acute.

isis

produced outward beyond the lateral

not always well-marked; the individual teeth may be blunt or

In the specimen shown the normal condition is illustrated; the central lobe includes

and each lateral part about seven teeth, the lateral teeth being usually a little thesome specimens the median lobe has the teeth very indistinct, while in others
there is a conspicuous tendency for the median pair to unite into a single broad tooth.)six teeth,

larger; inPrementum pale basally, indistinctly covered with pale papillae; palpi lying close together,subparallel, the lateral parts with short, dense hairs, the ventral face with shorter papillae;each lobe bearing at tip a small, blunt, cylindrical knob which is slightly chitinized.

Antenna

XV, 18) short, almost cylindrical but slightly narrowed basally, bearing on trimcate
apex about five sensory papillae of various diameters, one being much the largest, two others(Platebeing long and very slender; one of the papillae is bisegmented, the apical part being more

(De Meijere [1916, fig. 14] shows his European Ptychoptera

slender than the basal part.

as having the antennae two-segmented and with the sensory papillae very different fromthe condition found in P. rufocinda.)

Mandible (Plate XV, 19) strong and powerful; cuttingedge sub triangular, with three large outer teeth and about six or seven small inner ones;of the larger teeth the outermost is the slenderest, the third is the largest and stoutest; small

inner teeth subequal in length, the outermost stout, inwardly the teeth becoming
slender;moremandible on ventral face near margin with two powerful setae, the posterior one

often recurved, the anterior one directed forward.

Maxilla (Plate XV, 19) with cardines

roughly elongate-triangular, proximal angle acute, ventral face with three large setiferous

punctures bearing several bristles of unequal length (the writer has not been able to locate
setae on the middle puncture) outer edge of sclerite with a fringe of long hairs, longest at
narrow inner end of segment; stipites roughly triangular, with a group of short spines at;apex; palpi antenniform, stout, cylindrical, with sensory papillae at tip (one large papilla,

about three of medium size, and three or four small ones); outer lobe of maxilla with a small,
egg-shaped or subcylindrical, knob at its outer angle, below which the rounded lobe isdensely clothed with long, pale hairs; caudad of these, along margin, a row of from six toeight powerful spines and a few long hairs.

Pupa.— Total length, .34.3-40 mm.
Length excluding kreathing tube, 14.5-15.8 mm.
Length of breathing tube alone, 19.8-26 mm.

Width of body, d.-s., 1.6-1.7 mm.
Depth, d.-v., 1.7-1.8 mm.
Pronotal breathing horns reddish brown, dark brown at extreme base; thorax, wing
sheaths,

andleg she;iths

dark brown; abdomen whitish, with small tubercles

and broad

Charles Paul Alexander

778

chitinized apices to segments dark brown.

Pupa most readilystellate

abdominal tubercles.

(Platepupa
and the short, non-distinguished fromof Bittacomorpha by the venation, the position of the tarsal sheaths,

XIV, 13.)

Anterior cephalic crest small, deeply bilobed

by a broad V-shaped notch, the rounded
Dorsad of these, two smaller

lobules roughened and each terminated by a single long hair.

and slenderer, very widely separated, tubercles, each lying just inside antennal sheaths.
Antennal bases located on ventral side of head, between eyes; antennae bent dorsad andthence caudad around knee joints of fore legs, the tips lying just outside middle tibiae; apicalantennal segments showing distinctly thru sheaths.
tips curved over knee joints of fore legs.

Sheaths of maxillary palpi elongate,

Clypeus smaller and narrower than the conspicuous

Labium rectangular, each half with tips obliquely truncated.
XIV, 14.)
Pronotum with lateral ventral angles almost square, each with about two small setae.

labium, transversely wrinkled.
(PlateBreathing horns very unequal in length; the right one greatly elongated, much longer thanremainder of body, enlarged at extreme base, the outer part with numerous papillae which
are more numerous and more approximated toward end of organ, these papillae lying in asingle straight line which makes a long spiral around organ; left breathing horn very small

and degenerate, only a little longer than sheath of maxillary palpi, curved, with about adozen papillae which are closely crowded toward apex of organ. Mesonotum finely andtransversely wTinkled, prescutum with a very acute V-shaped dorso-median mark,

its

apex

Postnotum with
two small tubercles at about two-thirds its length, one on either side of a pale median Hne.
Metanotum short, sheath of halteres extending just beyond base of second abdominal
segment. Wing sheaths clearly showing venation, the branched media being characteristicLegof the genus; wing sheaths extending almost to end of second abdominal segment.
sheaths with tips of fore and middle tibiae enlarged, and with inner apical angle of eachproduced into long points for the long tibial spurs of adult flies. Leg sheaths extending tojust before end of third abdominal segment; all six legs lying side by side, not overlapping
directed backward; scutal lobes projecting, each with a few tiny hairs.

as in Bittacomorpha (Plate XV, 21).
First abdominal segment chitinized above, apical half with a few weak tubercles.

Dorsumsegments 2 to 6 with posterior margins each having a chitinized band set with about
twenty-five to thirty setiferous tubercles, the outermost ones being the largest; remainderofof dorsum of each segment with irregular transverse rows of scattered tubercles,on narrow,interrupted, chitinized bands; usually one of the bands, at about midlength of segment,

broader and more strongly chitinized than the others; these bands obliterated on posterior
segments, but caudal band here very wide; about fifteen of these rows on segments 2 andand from twenty to twenty-five rows on segments 3 to 5; these bands not regularly transbut anastomosing rather freely, net occupying more than half of abdominal surface;segment 7 with the broad caudal band only; tubercles rather short, with three or four short,

6,verse,

irregular spines at tip,

Abdominal

these not presenting a stellate appearance as in Bittacomorpha.

sternites similar to

dorsum, tubercles lacking where leg sheaths rest against

segments 2 and 3; segments 4 to 6 with transverse rows similar to those of dorsum but
weaker; caudal bands on segments 4 to 7 very broad, the caudal margin with tubercles,
there being about twenty tubercles on segment 4, the number gradually decreasing to seg-

The Crane-Flies of New York
ment 7, on which there are about fifteen.

— Part II

779

Abdominal pleurites with tubercles arranged inlongitudinal rows, there being about six rows on each of segments 2 to 6, the rows being

almost continuous for the whole length of abdomen; tubercles at caudal margin of eachsegment enlarged and powerful; on segment 6 the rows converging behind into a single
powerful tubercle on caudal margin of segment; a similar enlarged tubercle on caudal marginof seventh segment; segment 7 and cauda narrov.-ed.
Male cauda (Plate XV, 22) with aprominent dorso-median lobe projecting directly away from the body; dorsal sheaths short,indistinctly bilobed; ventral sheaths very elongate, divergent (Plate XV, 23).

(Plate XV, 24)

Female cauda

with the same prominent dorso-median lobe; acidotheca of ovipositor long,straight, beyond its midlength a blunt, conical tubercle directed laterad and slightly caudad;

sternum (Plate XIV, 15) with ventral lobe only about half length of tergal acidothecae, caudal
margin with three lobules.

———

Nepionohjpe.
Orono, Maine, June 24, 1913.
Neanotype.
With the larval type.
Paratypes.
Both larvae and pupae, June 24 to July 5, 1913.

SUBFAMILY Bittacomorphinae

The following keys separate the genera of the subfamily Bittacomorphinae:

LarvaeSize small (total length under 20 mm.); coloration black, breathing tube light yellow;
breathing tube entirely retractile; body covered with very long projections which areincased in a black, horny substance; mandibles with an inner comb of teeth.

Bittacomorphella Alex. (p. 779)
Size larger (total length over 40 mm.); coloration rusty red; body tapering gradually to thelong, slender, partly retractile, breathing tube; body covered with transverse rows of shorter,
stellate tubercles; mandibles without an inner comb of teeth.

Biitacomorpha Westw. (p. 783)

Pupae
Size small (length, excluding breathing horn, under 12 mm.); right breathing horn small,degenerate; abdominal tubercles weak, tipped with several strong setae.
Bittacomorphella Alex. (p. 779)
Size larger (length, excluding breathing horn, over 14 mm.); right breathing horn elongate,filiform, longer than the body; abdominal tubercles strong, elongate, crowned by a circlet
of four or five spines and tipped with a setiferous papilla. .Biitacomorpha Westw. (p. 783).

Genus Bittacomorphella Alexander- (Gr. diminutive of Biitacomorpha)
1916

Bittacomorphella Alex.

Proc. Acad. >Jat. Sci. Phila., p. 545.

Larva. — Body short, covered with very prominent projections which are longest on

and caudal parts of body.

Pseudopods prominent, with very large, curved claws.

tube short, entirely retractile within body.

lateralBreathing

Head subquadrate, the foramen ventral inhead unbranched. Mandible with an inner comb of teeth. IMentum
bilobed, cephalic margin untoothed.
Coloration black; breathing tube light yellow.
Pupa.'
Right breathing horn very short, degenerate. Fore tarsi overlying middle tarsi.
Tubercles on abdomen moderately elongated, not crowned by a circlet of spines as in Bittacomorpha, but tipped with a long seta.
position; setae of—Charles Paul Alexander

780

The genus Bittacomorpholla contains but two described species, thegenotype, B. jonesi (Johns.), and the larger B. sackenii (Rodcr) fromwestern America, the immature stages of which are wholly

unknown.
There is no published literature on the biology of this group of crane-flies.
Bittacomorphella jonesi (Johns.)
1905

Bittacomorpha jonesi Johns.

Psyche, vol. 12, p. 75-76.

Bittacomorphella jonesi is a curious little phantom crane-fly, not uncom-

mon in cold Canadian woods thruout the Northern States, where it isusually found near running water or springs and often in small, dark

The adult flies sometimes lurk beneathdark bridges and culverts, where they are often associated with
An account of the habits of thespecies of Dolichopeza and Oropeza.
adults of this species may be found in an earlier paper by the writer(Alexander, 1916b: 545-546).
The larval habitat is very different from that of other local species

ravines or along shaded runs.
low,of thefamily, which, as

a

rule,

preferopen swamps, swales, or wet

meadows. The larvae of this species live in rich organic mud in shaded
woods. They were first found on the Bool hillside at Ithaca, New York,beneath decaying beech leaves in wet or damp mud which was mixed
with old beechnuts,

and similar debris.

hulls,

acorns,butternuts,

broken decayed twigs,

The Bool area is a very steep hillside with a general

northern exposure, heavily shaded with tall forest trees.
it

In former days

extended far to the eastward and was connected with Slim Jim Woods,

near the second bridge in Forest Home.

The cut area is now a pasture,

On the shaded
skunk cabbage occupies pockets or level areas where the soil
The forest cover consists of beech, hard maple,
is largely calcareous.
but patches of skunk cabbage and cat-tails still persist.
hillside thebasswood, yellow birch, red oak, butternut, elm, a fev/ large alders, a
few aspens, and on the surrounding hillside a thick stand of hemlock.
The shrubs include Ribes fioridum L'Her., Cornus alternifolia Linn, f., andThe herbage at this season is of the dominant skunk
similar species.
cabbage, young seedlings of Impatiens hiUora Walt.,

and Cardamine Douglassii (Torr.) Britt.

Geum rivale Linn.,

In places there are thick matsof mosses, Brachythecium rutabulum (Linn.) B.

Amblystegium on decaying prostrate limbs.

& S., on the limy soil, andThe Crane-Flies of New YorkThe

— PartII

781

cuiious larvae of Bittacomorphella were here found associatedfollowing crane-fly larvae and pupae: Dia-anomyia stulta,
Limnophila adusta, L. (Dicranophragma) fuscovaria, Ulomorpha pilosella,Penthoptera, MolophiJus hirtipenm's, Erioptera megophthalma, Ormosia
innocens, 0. nigripila, Tipula collaris, T. oropezoides, T. cayuga, andIn addition numerous other natural associates were found, suchothers.

withtheas the larvae of a carabid beetle, Nehria sahJbergi Fisch., sow bugs, mollusks,

and a great variety of other organisms.

On July 10,

1914, adults of

Bittacomorphella were not uncommon in the same association, at whichtime they were flying with other crane-flies such as Dicranoptycha germana,

Molophilus hirtipemiis,

M. puhipennis,

Erioptera vespertina, E. venusta,

and Gonomyia blanda.
The first larvae of Bittacomorphella were found on May 11, and atthat time were almost fully grown. They present a very remarkableappearance, being black or very dark in color and covered with numerous
long projections. The breathing tube, which is capable of entire retraction
within the body, is very short, and is light yellow, in contrast with theremainder of the body. The larvae are, as a rule, very slow and sluggish
in their movements, but when disturbed they become more active.
Largeand small larvae, of two distinct sizes only, were often found in the same
situations at the same time.
A fully grown larva was placed in rearingon June 11, 1917, and emerged as an adult male on June 24. This provides for a pupal duration of not more than thirteen days, but the pupal
stage is undoubtedly much shorter.
Larva.

— Total length, 15-17 mm.
Length exclusive of breathing tube, 12.5-14 mm.
Length of breathing tube, 2.4-3 mm. to base of;gills,

gills,

L3-L8 mm.; beyond

Ll-1.2mm.

Greatest diameter across body, L7-2 mm.
Coloration dark brown to almost black, breathing tube light yellowish; in young individuals and occasional older specimens, coloration more rusty.

Body short, stout, cylindrical but appearing depressed, covered with numerous elongate
projections.

Breathing tube very short and capable of complete retraction within body
Body appearing proportionately broader, and tapering more abruptly to(Plate XVI, 25).

breathing tube, than in other species of the family herein discussed, this appearance being

due to great length of lateral body projections.

The most notable single feature of larva consisting of the numerous projections frombody (Plate XVH, 33), these being incased in a blackened, horny substance which is somewhat brittle. Usual shape of body extensions cylindrical, slender, and generally simple

Charles Paul Alexander

782

except for those near end of body, which are asymmetrically once-forked.
jections heavily chitinized, but distal end almost transparent.

Basal half of pro-

Entire surface of projections

beset with numerous transverse rows of short hairs, usually about nine to twelve hairs in eachrow, nine and ten being conmion, these hairs doubtless serving to hold the blackened, horny
covering of the projection.
jections, eachSensory papillae (Plate XVII, 34) borne at or near apices of pro-with a long bristle; these papillae narrowed at base, thence enlarged to forma head, on which bristle is inserted; usually one or two bristles to each projection, but
Dorsaloccasionally an additional one present, which is much smaller and degenerate.
body projections occupying transverse rows across segments, those near lateral and caudal
parts of body being long and powerful, those on median region being short and degenerate;

segments ofthorax andabdomen subdivided into false segments,

these transverse rows

occupying caudal margins of these segments, there being usually from four to six of the

weak projections between the powerful lateral ones. In addition to these projections, sparsebranched hairs lying in the same transverse rows (Plate XVII, 35). Projections at end of
body surrounding base of breathing tube all long and powerful, and, as stated above, someweakly bifurcate.
Ventral body projections similar to those of dorsum but relatively smaller.
Abdominal pseudopods feebly chitinized at tips, with very large, slightly curved claws
which are but little shorter than the pseudopods themselves (Plate XVII, 32). Breathingtube short and stout, surface before apex transversely wrinkled. Papillae rather numerous,
bearing sense hairs scattered over surface of tube, those just back of apex short and spinelike, those farther back long and slender, very like and homologous to the bristles terminating
Gills two, stout, about one-third length of terminal
the body projections, as discussed above.
section of breathing tube.

Body

projections incrusted with a black, horny substance, as

discussed above; on lateral projections, apical setae likewise incased, at least basally, pro-ducing a bilobed or even a trilobed appearance; this black corneous incrustation brittle andeasilyremoved, leaving projection and bristles intact.

Head subquadrate, sides nearly parallel, occipital foramen ventral in position (Plate XVI,
head covered with numerous small, chitinized points which are longest on
anterior ventral angle slightly produced; two setiferous punctures onventral surface, one on either side just behind anterior angles, the other closer to foramen.

2G); surface ofposterior angles;

Dorsum with setae arranged as shown in Plate XVI, 27; setae simple. Labrum (Plate XVII,
31) with four dorsal setiferous punctures along anterior margin, themedian pair closely

approximated; another powerful seta occupying each lateral angle on dorsal

side;

dense

brushes of long hairs on either side beneath. Epipharynx (Plate XVII, 31) supported by two

powerful chitinized arms connecting across midventral region and then extending lateradand expanding outwardlj^ to form posterior margin of labrum; surface of epipharynx with
Labium (Plate XVI, 28) with cephalic margin of mentum almost
closely appressed teeth.
entire, the broad median part produced cephalad and feebly bilobed; palpi with a dense
Antenna (Plate X\'I,
fringe of long hairs around base and with sensory papillae at tips.
30) short, cylindrical, somewhat globular or barrel-shaped; about four elongate papillae
and two or three shorter ones, these papillae terminal in position, the largest one occupyMandible (Plate XVI, 29, 30) with the outer angle a powerful hooking the inner side.
bearing smaller teeth on ventral face at about midlength; inner angle flattened, and, besides
terminal blade, bearing a comb of about five teeth, the innermost being the longest andThe Crane-Flies of New York— Part II

slenderest; a dorsal line of strong setae extending from base of outer783

hook inward; outeredge of mandible with two strong setae, these protected at their bases by small dorsal ears,
or projections, from the mandible; dorsal face of mandible with a powerful hinged prostheca;
near base of mandible on dorsal face a curious five-lobed sensory organ. Maxilla (Plate

XVI, 29) with the cardo triangular, the ventral or outer face with two closely approximatedsetilerous

punctures;

triangular,

stipesantenniform, cylindrical, rather

elongated,strongsetae near palpus; palpus

w4th

three

and

with about six

sensory

papillae at tip,

one being much longer than the others; outer lobe of maxilla with cephalic margin blackened
and chitinized, inner angle with a dense brush of long hairs.

Pupa.

— Length excluding breathing horn, 9.5 mm.
Width, d.-s., 1.7 mm.
Depth, d.-v., 1.4 mm.

The following description is taken from the cast pupal skin of the onlyspecimen that was reared:
Antennal bases approximated on front between eyes. Clypeus bluntly rounded at apex,
Lobes of labium (Plate XVII, 36) broad, rounded apically; maxillary
palpi short and stout, broad at base, gradually narrowed to the short tip, which is nottransversely wrinkled.

recurved.

(The structure of the head and the eyes indicates some peculiar characters notpossessed by the pupae of related genera, but the cast pupal skin is insufficient for accurate
diagnosis.)Two bristles below eye and just above base of palpus, and a longer and more

slender seta farther laterad.

Sides of head behind antennae appear to be produced laterad

Right breathing horn small, degenerate, much curved. (In the single
pupal skin available, it cannot be determined whether the left horn has been broken off or isintoblunt points.

undeveloped.)

Just laterad of each breathing horn a small tubercle bearing a long seta.

Scutal lobes with about four stout seta*.

Tarsal sheaths of fore legs, as in the subfamily,

overlying the middle pair but shorter (Plate XVII, 37).

Abdomen with chitinized bands extensive, as in Bittacomorpha.
about as in other species of family.

Arrangement of tubercles

Tubercles shorter and weaker than in Bittacomorphaand not crowned by a circlet of spines, each being tipped with one or more (four or five) longsetae (Plate XVII, 38); the pleural tubercles the longer and many of them multisetose.
Cauda of male, in general features, similar to that in Bittacomorpha, the dorsal median
lobe (Plate XVII, 39) stout, the ventral horns (Plate XVII, 40) short and powerful,directed laterad; horns on dorsal lobes apparently lacking.

———Nepionotype.
Ithaca, New York, May 30, 1917.
Cast pupal skin, reared June 23, 1917.
Neanotype.
Paratypes.
Topotypic, May 15 to June 10, 1917.

Genus Bittacomorpha Westwood (Gr. Bittacus + shape)
1835
Larva.

Bittacomorpha Westwood.

London and Edinburgh Phil. Mag., vol. G, p. 281.

— Form elongate, body gradually narrowed behind into the partly

retractile

Integument with transverse rows of tubercles. Pseudopods on abdominal
segments 1 to 3 prominent, with large curved claws. Head subpyriform, dorsum with rows
breathing tube.

Charles Paul Alexander784

of prominent black spots converging behind.

tooth.

Pupa.

Mandible stout, with a single powerful outer

Mentum bilobed, anterior margin not comblike.

Color of body, rusty red.

— Right pronotal breathing horn very elongate; thesubdegenerate.

Fore tarsi overlying middle tarsi.

leftvery small and short,

Tubercles on abdomen very long, locatedon broad transverse bands of chitin, each tubercle with a star of four or five spines surrounding
the apex, which bears a long seta.

The genus Bittacomorpha, as here restricted, includes but two species
the genotype, B. clavipes

America,(Fabr.),

and B.

occidentalis—

Aid. of westernconcerning the biology of which nothing has been recorded.

The literature on the immature stages of Bittacomorpha clavipes is summarized under the family account (page 773).
Bit'acomorpha clavipes (Fabr.)
1781

1835

Tipuln clavipes Fabr. Spec. Ins., vol. 2, p. 404.
Bittacomorpha clavipes Westw. London and Edinburgh Phil. Mag., vol. 6, p. 281.

Bittacomorpha clavipes,

the " phantom crane-fly,"isacommon andwidely distributed species thruout North America east of the Rockies.
It is easily recognized by the black-and-white-banded legs, with their"and swollen metatarsi. The species is very
While incharacteristic of alder swamps and the wet margins of ponds.
male
trailingonfemaleahead,
thefly,
thecopulation the insects oftenalight
on
plant
the
When
they
a
stem,
behind like the tail of a kite.
female is invariably uppermost, the mafe often hanging free with noneof its feet on a support. 'The swollen metatarsi are almost completely
filled by the tracheae, and these serve to buoy the insects as they drift
conspicuously

enlarged

about in the wind.
in detail.

Brues (1900) descril^es these peculiar tracheal dilations

He says, in part:

When flying, Bittacomorpha uses the wings scarcely at all, relying in great measure upon
wind currents for transportation. The legs are exceedingly light, as the exoskeleton is thinand dehcate, and encloses practically no tissue which can serve to increase their weight.

In a letter from Dr. J. G. Needham, dated September 27, 1917, valuable

data on this habit of drifting are furnished, as follows:
Yesterday while crossing the Fall Creek bridge near my home on Cornell Heights, T made anobservation on Bittacomorpha that interested me greatly. A breeze was blowing up the gorge,
and on the breeze a Bittacomorpha was drifting rapidly upward in the usual flight attitude,with broadly outspread legs, the swollen metatarsi hanging vertically,allphantom-like in

It came up from below the level
slenderne.ss and in strongly contrasting black and white.
face, and passed on upward with the breeze untilof the rail, swept past within two feet of
Since
lost to view, perhaps 100 feet higher than the bridge, and much farther upstream.

myThe Crane-Flies of New York— Part II

785

the creature can fly only very slowly and here was moving several times faster (I could not
Perhaps this is asee whether it was using its winf^s), it was obviously drifting in the wind.
normal function of the expanded metatarsi.

The larvae are usually abundant in dccaj'ing vegetable matter in rich
mud about ponds and in swamps. The writer has found themespecially numerous in the Basin Swamp at Orono, Maine (in 1913),
and near Round Pond at McLean, New York. At Orono they wereorganic

associated with larvae and pupae of Phjchoptera rufodnda, Limnophila
macrocera, Pilaria tenuipes, P. recondita, Erioptera chlorophyUa, a varietyof chironomid larvae, numerous larvae of Trichoptcra in their cases,
nematodes, and leeches. Needham and Betten (1901:574-575) give asummary of the larval habitat of this species as they found it in thenorthern Adirondacks. Weston and Turner (1917:53) have recorded

the larvae as being scavengers and thus serving as important factors
in the elimination of sewage in the Coweeset Stream near Brockton,

Massachusetts.

The immature stages of this interesting crane-fly have been well conby Hart (1898 [1895]: 189-195), whose account has been briefly
summarized by Howard (1912:95-96). The larvae are found in shallow
sideredHere they Hve inwater that is filled with decaying vegetable matter.
mat of dead stems of rushes, grasses, and willow leaves, in semi-

thestagnant or slowly flowing water.

The larvae are elongate-cylindrical,
They are deep rusty red

with a long, partly retractile breathing tube.

or brown in color, quite distinct from the pale whitish larvae of Ptychoptera
They feed on decaying vegetableor the black larvae of Bittacomorphella.

matter, diatoms, and mud that is filled with organic matter.

They rest

beneath the surface of the water, with the tip of the extended breathingtube at the surface film or just beneath the surface, in the latter case
When about to pupate,
breathing by means of the small tracheal gills.
the very long, coiled breathing tube of the pupa is wound around thethorax beneath the larval skin.
out into a very long,

stiff,

On pupation the tube soon straightens

bristle-like structure.

Like the larvae, thepupae rest beneath th(^ surface of the water, with the tip of the breathingtube projecting above the surface film. The pupal duration is apparently
about a week.
Larva.

— Length when

fully extended, about GO mm.
Length of breathing tube, about 20 mm.
Diameter of body, about 2.G to 3 mm.

Charles Paul Alexander

78G

Body tapering gradually at either end, posterior end prolonged into breathing tube.
Usual color pale rusty brown, but the writer has found a few nearly full-grown specimens
which were as pale in color as the larvae of Ptychoptera.

Body covered with numerous
Head broadly ovate,transverse rows of small tubercles, or papillae, which bear short setae.

convex above, where it is conspicuously marked with rows of black spots, these interrupted
lines

converging behind.

IMouth parts in general similar to those in Bittacomorchella, themain points of difference being as follows: mandible (Plate XVIII, 42) shorter and stouter,ending in a powerful outer tooth, the comb of inner teeth being reduced to about eight small
tubercles, the two bristles on outer margin of mandible not overlapped by projecting" ears ";labium (Plate XVIII, 41) shorter and stouter, with a different arrangement of papillae; epiphar-ynx long, narrowed behind, distinctly bilobed, each half with parallel rows of long, combliketeeth projecting proximad; anterior comb of epipharynx with the anterior teeth the largest,

the teeth gradually reduced in size behind; posteriorcomb with the rows of teeth widely

separated anteriorly, approximated behind so as to be contiguous' or nearly so at their ends;

space between these rows filled with long hairs; anterior teeth small and feebly chitinized,
First three abdominal segments bearing conspicuous pseudopods,
each terminated by a sharp, slender claw which fits into a groove on the face of the pseudopod.

posterior teeth stronger.

Pupa.

— Total length, 40-GO mm.
Length excluding breathing tube, 1.5.5-25 mm.
Length of breathing tube, 25-35 mm.
Degenerate breathing tube, length 2 mm., diameter 0.2 mm.
Width, d.-s., 1.8 mm.
Depth, d.-v., 2.6 mm.

Breathing tube light brown; wing sheaths brown; leg sheaths hght brownish yellow anddark brownish black, alternated, corresponding to the leg markings of the adult fly. Abdomen
pale yellow, rather uniformly covered with abundant brownish tubercles and transverse,
chitinized plates, these brown areas scarcer on pleura and not especially abundant on apicalmargins of segments.

Pupa somewhat similar in general structure to pupa of Ptychoptera.

Anterior cephaliccrest small, lobules rounded, each tipped with a long, stout seta; immediately behind anterior
crest, a similar blunt, bilobed projection of front; laterad of crest, a slender, elongate tubercle

on either side, immediately behind untennal sheaths, each with a long seta; two other setiferous
tubercles on head behind antennae and maxillary palpi.
Antennal bases approximatedbetween eyes. Sides of head, laterad of eyes, with a small setiferous tubercle. Maxillary
palpi not recurved at tip, as in Ptj'choptera, ending opposite knee joint.

Clypeus elongate,

gradually narrowed toward apex, transversely wrinkled; two hairs toward base near inner

marginof eye.

Eachhalf of labium broad, roughly subquadrate,

tips

broadened andobliquely truncated (Plate XVIII, 43).

Breathing horns almost as in Ptychoptera. (Nearly always it is the right horn that is elongated, but in about ten per cent of the specimens the left horn is elongated while the rightis degenerated;

Hart records one specimen in which both horns were developed, but unequally,

the right measuring 23 mm. and the left 13 mm.; some of the specimens recorded by Hart
are larger than any that the writer has ever seen.)of fore tarsi; media unbranched.

Wing sheaths ending almost opposite tips

Leg sheaths (Plate XVIII, 47) with fore tarsi much shorter

The Crane-Flies of New York— Part II

787

than the others and lying directly over middle tarsi;

tarsi of hind and middle legs paralleland extending beyond tips of fore tarsi. Scutal lobes each with about four sctiferous tubercles.
Thorax and first al)dominal segment transversely crenulated.
Abdomen with transverse bands of chitin much broader than in Ptychoptera, so that they
cover almost the entire abdominal surface; these bands with about twelve tubercles on
segments 3 and 4, about ten on segment 5, and from six to eight on the posterior segments;
tubercles of various sizes, small and somewhat degenerate ones being interspersed withlarger ones; tubercles long and slender, each crowned by a circlet or star of from three tostout spines (Plate XVIII, 44-46), a setiferous papilla arisingfrom the center of this circlet of spines; spines on pleura longer than those on remainder ofsix (usually four or five)

abdomen, but not arranged in distinct longitudinal rows as in Ptychoptera, being usually more
irregular, in some cases showing three or four more or less distinct rows; these pleural chitinized areas usually bearing from two to four tubercles, which are closely approximated basally
so as to present a somewhat branched appearance.
IMale cauda (Plate XVIII, 48) as in
Ptychoptera, but dorsal median lobe very short and stout; tubercles on segment immediately
before cauda long and slender, similar to those on remainder of abdomen.

— Ithaca, New York, May 1917.
— Orono, Maine, June 24, 1913.
— With the type pupa.

Nepionotijpe.

15,

Neanolype.
Paralypes.

Malloch's figure of the pupa (1915-17 b: pi. 35, fig. 6)
It

was probably made from a femaleisdiagrammatic.

individual, the antennal sheaths

being shorter in this sex than in the male.

Family Rhyphidae
Larva.

— Bodyconstrictions.

Spiracuiar disk surrounded by two or five lobes

lobes (Trichocerinae)

Pupa.
tal

Mandibles opposed.
Eyespots distinct.
Thoracic and abdominal segments divided by false

eucjphalous, amphipneustic.

Spiracles on sides of prothorax.

—Head

,

with a bilobed setiferous cephalic crest.

breathing horns(Rhyphinae) or by fouror unprovided with lobes (Mycetobiinae).

short, notprominent.

Tarsal

Palpi stout, straight.

Prono-

sheaths lying in pairs, one above

another, the fore legs lying on the middle legs, and these latter on the hind legs.

Lateralabdominal spiracles small but distinct.
r."

The family Rhyphidae
of genera

which areincludes an apparently heterogeneous groupin reality

very closely related.

The adults are ofdiverse appearance, but the immature stages are exceedingly similar to

one another and undoubtedly
family are closely allied.

allthree of the groups included in the

The immature stages of the Rhyphinae (Rhyphus) have been discussed
by many entomologists, among others by Johannsen (1910:35-36,

Charles Paul Alexander

788

Rhyphus pundatus) and by Malloch (1915-17 b: 243, jR. pundatus). The
larvae are often handsomely banded and mottled with brown or purplish.
Johannsen and other authors describe the cauda as ending in two shortThe

general structuralcharacters are those described above for the family.

The larvae occur

but Malloch mentions

lobes,

fivesuch lobes.

in decaying vegetable matter, in manure (especially horse and cow dung),
in sewage,

and in similar material.

The Mycetobiinae are represented by Mycetobia, a curious fly which
superficially resembles a mycetophilid rather than a crane-fly.
Long ago
Lyonet, Dufour, Guerin-M(§neville and others described and figured thelarva of Mycetobia and noted the eucephalous condition of the head andthe amphipneustic spiracles.

Osten Sacken

affinities of this genus with Rhyphus.

(1863)

firstsuspected theMore recently, work by Johannsen

Malloch (1915-17 a, and 1915-1713:244-245), Edwards
and others has definitely settled the relationshipThe larvae and the pupae agree
insect with the Rhyphidae.

(1910:31-32),(1916),of thisKnab(1916),The larvae
wood and about fermenting sap in wounds of trees.
The genera Ditomyia Winn, and Symmerus Walk, are now placed in aclosely with the general family characters discussed above.

occur in decaying

separate family from Mycetobia, the Ditomyiidae (Keilin, 1919).
Until recently, the Trichocerinae have been considered as being members of the family Tipulidae.

They include only the genus Trichocera,with about twenty-five nominal species, and, presumably, Ischnothrix
From the general
Bigot, represented by a single species from Cape Horn.

appearance of the adult, these flies have usually been referred to the tribe
Brunetti (1912)
Limnophilini, in a position near the genus Limnophila.
referred them to the Pediciini, and most other recent workers have accorded

themtribalinfluencedor

subfamily rank in the Tipulidae.

by the workof

Keilin(1912),

referredBezzi(1914:214),

Trichocera to the

Rhyphidae, but later (1918a: 20) placed it back in the Tipulidae (as LimMalloch (1915-17b:234) likewise places Trichocera with thenobiidae).
Tipulidae, but mentions the close resemblance of the larva to that of
Rhyphus. The best discussions of the morphology of the larva and thepupa are those by Keilin (1912) and De Meijere (1916:191-194), bothof whom were strongly impressed by the striking resemblance of the larva
to that of Rhyphus.

In the present paper, the Trichocerinae is the only

group considered in detail.

The Crane-Flies of New York — Part II

789

SUBFAMILY Trichocerinac

Genus Trichocera Meigcn (Gr. hair + horn)
1800
1803

Petaurista Meig.
Trichocera Meig.

1911

Paracladura Brun.

Larva.-

Nouv. Class. Mouch., p. 15 (nomen nudum).
Illiger's Mag., vol. 2, p. 262.

Rec. Indian Mus., vol. 6, p. 28G.

— Body eucephalous, amphipneustic.

Thoracic segments divided into two annuli.

Spiracles on lateral margin of posterior ring of prothorax.

Abdominal segments divided

Cauda ending in four lobes, ventral lobes the longer and more slender;
Eyespots distinct. Laterallobes bearing numerous stout hairs near tips on outer face.
into three annuli.

plates ofhead widely separated on midventral line.

Mandible with prostheca distinct.
Clypeus short; labrum dumbLeg sheaths lying in pairs above one— Cephalic crest small, lobes with stout setae.

Pupa.-

bell-shaped; palpal sheaths stout.

Antenna elongate.

another, gradually lengthening, fore pair the shortest, posterior pair the longest.

breathing horns short.

Abdominalspiracles small,

Pronotalbut distinct and functional.

The small winter gnats of the genus Trichocera are rather familiar,
months in cellars or even insince they are not rare during the winterthe open on

swarms.

warm days,

occurring in sunlit places in small, dancing

They are abundant during fall and spring.

They occur also

shady places in summer, but are less in evidence at this season.
Trichocera is found somewhat commonly and regularly in mines, often

in cool,

at very considerable depths.

Boheman (1850) records specimens of T.

mines 600 feet below the surface, and Lampa (1890) also
Dr. H. B.
records the species as being found at considerable depths.
Hungerford found numerous adults of a species of Trichocera in theAmeth^yst silver mine near Creede, Colorado, in 1914. Specimens that
he obtained were taken at the sixth level, but the miners said the insects
were to be seen in all parts of the mine; along the laterals at the sixth
It is supposed thatlevel they were noted 7000 feet from the entrance.
these individuals breed in the animal waste which naturally accumulates
Trichocera is also a characteristic inhabitant of cavernsin such places.
and grottoes, all stages being found in such situations (Schmitz, 1909:80;
regelationis

inBezzi, 191 1-12 :46-i7, 49,

and 1914:214).

The swarming and mating of these flies is well known. It has been
ably described by Ainslie (1907), and is here discussed only in generalterms. The insects swarm commonly in the autumn. Sometimes theswarms include but comparatively few individuals, but at other times
many thousands participate. They swarm usually from five to twentyfive feet above the ground, all facing in the same direction, that is,Charles Paul Alexander

790

toward the wind or breeze. Mating takes place in the air, and united
The swarms arepairs then fly away or drop to the ground beneath.
often very dense, and individuals come in frequent contact with one
When the breeze shifts, the swarm immediately readjusts its
another.
position and direction.
The immature stages of Trichocera are spent in decaying vegetable
matter, beneath dead or decaying leaves, in debris, in fungi, and insimilar situations.
Sometimes the larvae and pupae are rather numerous
in stored roots and tubers, especially potatoes, in which cases they may
assume an economic importance (Johannsen, 1910:34-35; Carpenter,
The specimens used by the writer for study are part of Johannsen's1912).
The taxmaterial, determined as T. regelationis from Patten, Maine.
onomic condition of the group is such that no specific identification of
the adult flies can be attempted at this time. The immature stages of thegeneralized subgenus of Trichocera, Diazosma Bergroth, are unknown.
The most important literature on the genus Trichocera is as follows:
Trichocera regelationis
Trichocera regelationis
Trichocera rcgelalionis

GeneralLarva, general
Larva, pupa

Dufour, 1840: 161.
Schniitz, 1909:80, pi. 8, fig. 3.
Johannsen, 1910:34-35, figs. 51-

Trichocera regelationis
Trichocera hiemalis
Trichocera hiemalis
Trichocera fuscata
Trichocera sp
Trichocera sp
Trichocera sp
Trichocera sp

Pupa

De Meijere, 191G: 194.

Larva, general
GeneralGeneralCurtis, 1846b.

57.

Trichocera sp
Trichocera sp

Larva
Larva
General

Larva
Larva, pupa

Larva

Cameron, 1917:63.
Carpenter, 1912.
(Damage.)
Bremi-Wolf, 1846: 175.
Perris, 1847: 37, pi. 1, fig. 3.
Bezzi,1911-12:4()-47, 49.
De Meijere, 1916: 191-194, figs.
21-23.
Keilin, 1912.
(Morphology.)
Malloch, 19 15-17 b: 234-235, pi.
26, fig. 1; 306.

Trichocera regelationis, supposition.
Larva.

— Length, 8-9.5 mm.
Diameter, 1 mm.

Coloration pale brown in preserved material, whitish in fresh specimens.

Body rather short, cylindrical to slightly depressed (Plate XIX, 49). Pseudopods lacking.
Head complete, non-retractile, strongly chitinized; lateral plates of head widely separated
on midventral line, connected only by a narrow bridge posteriorly; chaetotaxy as shown
Mandibles opposed, of three parts, principal segments bearing(Plate XIX, 52 and 53).
on inner side near base an apparently movable appendage (prostheca) which has, besides theLabrum bluntly rounded, with long hairs. Epilarge apical tooth, three smaller teeth.

pharynx with lateral combs of about six blunt teeth.

Antenna two-segmented; basal segment

The Crane-Flies of New York— PartII

791

very short, disk-shaped, inserted on a large brown-margined plate which is part of the headsecond segment much narrower, egg-shaped; in addition to this segment there arechitin;several small sensory papillae on the end of the first segment.

Segments of body divided into secondary annuli, thoracic segments with two such rings,
abdominal segments with three; annuli bearing transverse rows of short setae. Anterior
spiracles on posterior ring of prothorax near lateral margin conspicuous, smaller than posterior
but constructed on same general principle.

Spiracular disk surrounded by four
more slender than dorsal pair, inner face narrowly chitinized,
outer face densely clothed with abundant short yellow hairs (Plate XIX, 50 and .51);
dorsal lobes shorter and blunter, with short hairs on apices of outer face.
Spiracles large,
spiracles

lobes; ventral lobes longer andat base of dorsal lobes.

Pupa.

— Length, 7.5-7.S mm.
Width, d.-s., 1.4 mm.
Depth, d.-v., 1.4 mm.

Coloration whitish; head, thorax, and sheaths of appendages brown.
Anterior cephalic crest small, lobes widely separated, each tipped with a long, stout seta
directed ventrad; a tiny seta just behind each anterior lobe.
slightly

Antennal bases above andbetween the ej'es, bent dorsad and thence caudad, passing behind joints of legs, in
Frontal region between eyes slightly

the female attaining to about one-third length of wing.

tumid, somewhat shiny.

Clypeus short, the sides

parallel,broad, dumb-bell -shaped, the caudal margin concave.

ending before knee joints of fore legs (Plate XIX, 55).
base of antenna.

Mesonotum

the apex U-shaped;

labrumMaxillary palpi very short and stout,

A small tubercle just laterad of

Each cheek produced into a long, blunt, wrinkled tubercle.

(Plate XIX, 54) strongly gibbous, pale medially, narrowed in front, anterior

margin truncated and sending a sharp median carina cephalad; sides of mesonotum opposite

wing root with four small setae, in two slightly separated groups.

notum with a short bristle.

Anterior angles of pro-Breathing horns small, short and almost straight, broad basally,

apical half narrow, inner face fused or closely

approximated with pronotum, apex cleft.
Leg sheaths with

Wing sheaths attaining level of tips of fore tarsi; venation rather distinct.
fore legs stout; forehindtarsi;

three

false

tarsi

overlying middle tarsi

terminal segments ofannuli;caudal(Plate XIX, 55);

middle tarsi overlying

Abdominal segments divided into about
each segment fringed with short hairs. Tiny
Female ovipositor (Plate XIX, 56) with the dorsal

tarsi swollen.

margin ofabdominal spiracles on pleural segments.

valves short, widely separated, acutely pointed; ventral acidotheca elongate, approximated,bent slightly ventrad.
Larvae and pupae.

— Patten, Aroostook County, Maine, May 3 and 23, 1907.
Family Tipulidae

The family Tipulidae is the largest group of crane-flies, and possibly
the only one to which thename is justly applicable.

It includes

a vastnumber of species (nearly three thousand), arranged in about one hundred

Charles Paul Alexander

792

and forty genera. The species are found in most parts of the world, being
Crane-flies require moisture
restricted onl}' by intense heat and cold.
in order to complete their development, and, as a consequence, are almost

always found in the neighborhood of flowing or stagnant water.
speciesknown

to the writer are inhabitants of desert

conditions,Notheapproach probably being some Eriopterini, such as Helobia,
Trimicra, and other genera.
The immature stages frequent very wide ranges of habitat, which are
They are readily separalile from otherindicated elsewhere (page 716).
The
related species by the characters outlined on pages 744 to 758.
following
keys:
subfamihes of Tipulidae may be separated by thenearestLarvae1.

Body provided with elonga,te spiues or leaf like projections.
Cylindrotominae, pars (p. 959)

2
Body without distinct spines
2. Form depressed, with more or less distinct lateral tubercles; terrestrial on spermatophytic

Cylindrotominae, pars (p. 959)

plantsForm terete; if depressed, without tubercles
3. Spiracular disk surrounded by six or eight lobes
Spiracular disk not as above

3

Tipulinae (p. 974)
4

5
surrounded by two, four, or five lobes
7
Spiracular disk with three lobes or without distinct lobes
Head capsule massive, the hypopharynx a flattened plate with few teeth; size large
6(aberrant Tipulinae)
Head capsule massive or dissected, if the former the hypopharynx not as above; size
Limnobiinae, pars (p. 793)
usually small
Genus DoHchopeza Curt. (p. 981)
Spiracular disk with five lobes; lives in moss
Spiracular disk with four slender, hornlike lobes; lives in earth.
Tipula selene Meig. (p. 1016)
Size large (30 mm. or over); form very stout, terete; head capsule of the tipuline type;
Genus Tanyptera Latr. (p. 988)
lives in wood
Size small (20 mm. or under); form slender, terete; head capsule of the limnobiine type.
Limnobiinae, pars. (p. 793)

4. Spiracular disk

5.

6.

7.

Pajme
1.

Basal abdominal segments unarmed with teeth or spinous projections before posterior
2
marginBasal abdominal segments armed with a transverse row of usually small teeth or chitinous
projections before posterior margin.3

Last larval skin adhering to posterior end, attaching pupa to a plant stem or a leaf;
Cylindrotominae, pars (p. 959)
coloration bright green
Limnobiinae, pars (p. 793)
Not as above
3. Maxillary palpi curved or recurved at tips; size large, usually 12 mm. or over.
Tipulinae, pars (p. 974)
4
Maxillary palpi not recurved at tips
4. Maxillary palpi long, slightly or decidedly curved at tips; size large, length usually 12 mm.
5or over
Maxillary palpi short, straight; size small, usually 10 mm. or under.
Limnobiinae, pars (p. 793)

2.

The Crane-Flies of New York — Part II5.

793

Dorsal abdominal segments with two slender spines before margin.
Cylindrotominae, pars (p. 959)
Dorsal abdominal segments with four or more teeth or spines before margin.
Tipulinae, pars (p. 974)

SUBFAMILY Limnobiinae

The subfamily Limnobiinae
small crane-flies.
other

principal

includesa vast assemblage of usually

Only a few genera approach the ordinary size of thesubfamily,

theTipulinae, such genera being Limnobia,

Psaronius, Limnophila, Eriocera, Pedicia,

and a few others.

The writer has endeavored to key the immature stages of tribes, subAs has been stated elsewhere, the keys are based
tribes, and genera.
almost entirely on material seen by the writer, and additional specimens
undoubtedly modify the arrangement very conhowever, that the keys ai given will at least
The charactersfurnish suggestions or a basis for succeedinj work.
given in the keys, in so far as is possible, are tho33 that can b3 seen without making a detailed dissection of the specimen. However, for most
species it is necessary to study the larval head, as already outlined (pageThe character of " head massive " or " head rodlike " can often741).
be detected thru the larval integument without dissection. The spiracular disk is usually studied without especial difficulty.
The immature stages of the majority of the species are spent in moistofother species willsiderably.

It is believed,

earth, usually near water.
Elliptera,

Some are nearly, if not quite, aquatic (Antocha,

some Dicranomyia);

some Limsome
Elephantomyia, andothers are fungicolous (Ula,nobia); several live under the bark of trees (some Dicranomj'ia,

Rhipidia,

Discobola,

Gnophomyia,

Teucholabis,others); one, at least, mines in the leaves of plants (Dicranomyia).

The habits of the various genera and species are discussed in greater
under the respective titles. The following kej's separate thedetailtribesand the subtribes of the subfamily Limnobiinae:
Larvae

1.

2.

3.

_

Spiracular disk provided with two long ventral lobes
2
Spiracular disk not as above
3
Spiracles lacking or vestigial; mentum not completely divided medially; hypopharynx achitinized double comb; species aquatic, in silken ciises.
Limnobiini, subtrlb:; Antochiri^ (p. 79.))
Spiracles large, prominent, exposed; mentum completely divided medi.ally; hypophcrynx
rediciini (p. S''4)
labriform
Spiracular disk surrounded by four or five lobes
419
Spiracular disk surrounded by three lobes or without distinct Lbcs

794Charles Paul Alexander

4.

Head capsule massive, compact, the posterior incisions usually shallow
Head capsule of four or six slender rods, the posterior incisions profound•5.

Mentuin completely divided, a toothed plate on either side; abdominal segments without

5
15
6

distinct creeping-welts

Mentum, if present and chitinized, not completely divided; abdominal segments with
8
bi.sal creeping-welts
Eriopterini (p. 908)
Spiracular disk squarc-ly truncated, surrounded by five lobes
7
8piracular disk with four lobes
7. Each mental plate four-toothed; hypopharynx labriform.
Pediciini, subtribe Add phomy aria (p. 895)
Each mental plate with seven or eight teeth; hypopharynx a comhlike ring.
Hexatomini, subtribe Pseudolimnophilaria (p. 848)
98. Spiracular disk with five lobes
12
Spiracular disk with four lobes
9. Anteimae almost globular, with two conical apical papillae; lives in fungi.
Hexatomini, subtribe Ularia (p. 838)
10
An+ennae elongate-cylindrical
10. Abdomen with dorsal and ventral creeping-welts; menium with more than five teeth.
Limnobiini (p. 795)
11
Abdomen with six ventrrJ welts only; mentum with live or fewer teeth
Limnobiini, subtribe Rhamphidaria (p. 830)
11. Mentum five-toothed; lives in earth
Hexatomini, subtribe Epiphragmaria (p. 843)
Mentum three-toothed; lives in wood
12. Antennae almost globular, with two conical apical papillae; lives in fungi.
Hexatomini, subtribe Ularia (p. 838)
13
Antennae elongate-cylindrical
13. Abdomen with dorsal and ventral creeping-welts, the latter naked; mentum not threeLimnobiini, subtribe Ellipteraria (p. 806)
toothed; forms aquatic
Abdomen with ventral creeping-welts only; mentum with only three primary teeth;
14species not aquatic
14. Form long, slender; skin naked, shiny, transparent; apical segment of antennae elongate,
as long as, or longer than, basal segment; mentum with a smaller tooth on either side;
Limnobiini, subtribe Dicranoptycharia (p. 828)
lives in earth
Form short, stout; skin white, opaque; apical segment of antennae short, hemispherical;
mentum without small lateral teeth; lives under hark.
Hexatomini, subtribe Epiphragmaria (p. 843)
16
15. Blades of maxillae not produced; form long and slender
Blades of maxillae produced into flattened elongate appendages, the tips of which protrude from the thoracic orifice when the head is completely retracted; form short17
and stout
16. Spiracular disk squarely truncated, surrounded by five lobes which are fringed withnumerous, usually short, hairs; esophageal region not conspicuously grooved.
Eriopteri7ii subtribe Eriopteraria (p. 911)
Spiracular disk surrounded by four lobes, each ventral lobe with a single elongate bristle;esophageal region elongate, grooved; lives under bark.
Eriopterini, subtribe Elephantomyaria (p. 952)
17. Mental region a narrow, transverse, chitinized bar.
Hexatomini, group Limnophilae (p. 858)
18
Mental region not chitinized
18. Mandibles hinged; maxillae and labrum densely hairy; dorsal plates of head capsule
Hexatomini, group Ulomorphae (p. 869)
united into a spatula
Mandibles not hinged; maxillae and labrum not densely hairj'; dorsal plates of headcapsule separated
Hexatomini, subtribe Hexatomaria (p. 876)
20
19. Head capsule massive, compact
Head capsule of six slender rods; lives under bark.
Eriopterini, subtribe Eriopteraria (p 911)
C.

,

The Crane-Flies of New York20.

— Part II

795

Mental plates not completely divided; abdominal segments with basal creeping-welts
on both ventral and dorsal surface; spiracular disk indistinctly four- or five-lobed.
Limnobiini (p. 795)

Mental plates completely divided; abdominal segments without welts; spiracular disk
Eriopterini (p. 908)

obliquely truncatedPupae
Pronotal breathing horns eight-branched; forms entirely aquatic.
Limnobiini, subtribe Aniocharia (p. 799)
2
Pronotal breathing horns simple, unbranched
2. Rostral sheath elongated; lives in wood. .Eriopterini, subtribe Elephantomyaria (p. 952)
3
Rostral sheath not longated
3. Pronotal breathing horns very minute, conical, visible only with a lens.*
Limnobiini, subtribe Dicranoptycharia (p. 828)
4
Pronotal breathing horns larger, not microscopic
5
4. Dorsal spiracles on eighth abdominal segment large and functional
7
Dorsal spiracles on eighth abdominal segment small or lacking
5. A large circular spinous area on abdominal pleurites; cephalic crest chitinized, acutely
pointed; pronotal breathing horns directed ventrad; lives in decaying wood.
Hexatomini, subtribe Epiphragmaria (p. 843)
1.

;

Not as above(>

Pronotal breathing horns large, flattened, the tips yellow; abdominal tergites with
Hexatomini, subtribe Ularin (p. 838)
shagreened crossbands
Pronotal breathing horns slender, cylindrical; abdominal tergites with tran verse rows of
Limnobiini, subtribe Rhamphidaria (p. 830)
small spines
7. Abdominal pleurites with circular areas set with numerous microscopic spicules; pronotal
breathing horns short, usually truncated at tips, which are margined with the breathing
Pediciini (p. 894)
pores
Abdominal pleurites not as above, if with spines these large and few in number; pronotal
8
breathing horns long, cylindrical
8. Abdominal segments with broad transverse bands or welts on basal rings of third to
Limnobiini (p. 795)
seventh tergites
Abdominal segments with basal ring unarmed as above, posterior ring before margin
9with a transverse row of spines or stiff setae
9. A distinct crest on mesonotal prescutum armed with tubercles, spines, or setae; size
Eriopterini (p. 908)
small (usually under 9 mm.)
No distinct crest on mesonotal prescutum (scutellum armed in some Eriocera); size
10
large (usually over 10 mm.)
10. Leg sheaths very short, barely exceeding wings; lives under bark.
Eriopteriin, genus Gnophomyia (p. 934)
11
Leg sheaths longer, extending one or more segments beyond tijis of wings
n. Size small (under 6 mm.); abdominal armature weak, lacking on segment 7.
6.

Pediciini, subtribe Adelphomyaria (p. 89",)
Size larger; abdominal armature stronger, spinous; if small in size (Dicranophragma),basal armuli of abdominal segments armed with naked tubercles. .Hexatomini (p. 835)

Tribe Limnobiini

A large group of crane-fli(^s, arranged in a few often extensive genera,
comprise the tribe Limnobiini.

At first sight the tribe appears to be a

5 In thegenus Cladura (Eriopterini), reared while this paper was going thru the press, the breftthing
pores are likewise microscopic, being entirely sessile (page 949).

Charles Paul Alexander

796

heterogeneous assemblage, but in reality it constitutes a natural group.

The tribe as herein arranged includes the old group Limnobiini, with theaddition of several genera that were formerly distributed in the Antochini.

The divisions of the tribe as now constituted are as follows:— including the old
Limnobiini.
— including the genus
This
preceding subtribe and may be a group belonging to
Antocharia — including Antocha and presumably
Orimargula, Orimarga, Diotrepha, and possibly
Rhamphidaria — including Rhamphidia andDici^anoptycharia — including Dicranoptycha only.
1.

Limnobaria

2.

Ellipteraria

tribe

Elliptera.

isclose to theit.

allied genera, as

3.

others.

4.

its allies.

5.

These groups are not far removed, phylogenetically, from the lowermost*divisions of the Hexatomini, such as the Ularia and the Epiphragmaria,
and the two tribes are unquestionably closer together than their arrangement on paper would indicate. The separation of the two major groups
was made largely on the characters of the imagines.
The larvae have the bod}'' terete, moderately elongate or very long andslender (Dicranoptycha).
The abdominal segments are subdivided intoa basal and a posterior ring, the former with transverse welts of microscopic chitinized points or hooks.

In the Limnobaria and the Antocharia

these welts occur on both the dorsal aiid ventral surfaces in the form of

microscopic hooks; in the Ellipteraria they are on segments 3 to 9 on thedorsal surface only, being indicated on the ventral surface but naked;

Rhamphidaria the welts are ventral in position on segments 2 to
Dicranoptycharia they are similar, on segments 2 to 8. The
body in Dicranoptycha is entirely glabrous.
The head capsule is of moderate to large size and is massive and compact, consisting of a narrow dorsal plate which is usually indented behind,and two broad mussel-shaped lateral plates which are connected anteriorly across the venter to form the mental plate.
The mental plate
consists of an outer plate which usually terminates in a single medianpoint, and behind this an inner plate which contributes additional teeth
to the mentum. ,In Dicranoptycha there is but one subequal tooth oneither side, with an additional much-reduced tooth; in Rhamphidia
there are two teeth, and in the other groups there are usually four or five.
The hypopharynx is usually a double plate united at the ends to formin the7; in theacollar,with the anterior margins finely toothed.

The maxillae

areThe Crane-Flies of New York— PartII

797

and simple in structure; the cardo and stipes are large; the palpus is
The antennae have the apical segment or papilla ranging from
elongate, in Dicranoptycha, to very flattened and disklike, in Limnobia
and its allies. The mandibles are usually of simple structure, with one or
two dorsal teeth and from three to seven teeth in the ventral cutting
large

flattened.

row.

The spiracles are lacking in some species, at least, of Antocha. The
is surrounded in Rhamphidia by five subequal lobes, in
Dicranoptycha by four slender, naked lobes; in many Limnobaria the lobes

spiracular disk

are lacking or indistinct.

The larvae of many of the species are able to spin silken cocoons or
tubes in which they live.

These tubes are open at both ends, and areusually covered exteriorly with particles of extraneous matter gatheredinthe larval haunts.

The pupae usually lack a distinct setiferous cephalic crest, altho one
The pronotal breathingis present in Rhamphidia and in Dicranoptycha.
horns are usually large, and are either subcircular, or wider than long (mostLimnobaria), rarely elongate (Rhamphidaria), very large, earlike, andcontiguous or practically so on the median line (Ellipteraria), or branchedin theDicranoptycharia, howThe abdominal segments on the basal ring
often show a transverse welt of small hairs or a double convergent row of

into eight long filaments (Antocharia)
ever,

;

they are microscopic.

chitinized hooks; in Discobola,

Rhamphidia, and Dicranoptycha, how-ever, this is apparently not the case, the abdominal armature being more

The eighth abdominal segment often bears a pair of dorsal spiracles; these are apparently lacking
in some species (Antocha saxicola) and are small in most Limnobaria, but
are large and functional in Rhamphidia.
The following keys separate the subtribes of the tribe Limnobiini:
eriopterine or hexatomine in appearance.

Larvae1.

Body ending in two long vertral lobes; spiracles lacking or very reduced; forms strictly
aquatic

Body not as above; spiracles large
2. Body with ventral and dorsal welts on abdominal segments
Body with ventral welts only
3.

Antocharia (p. 799)
2

3
4Spiracular disk surrounded by four lobes which are provided with long fringes of hair;dorsal welts microscopically spiculose; ventral welts naked; species aquatic.
Ellipteraria (p. 806)

Spiracular disk not as above; dorsal and ventral welts alike

Limnobaria (p. 808)

Charles Paul Alexander

798
4.

Body moderately elongated, covered with a long, dark pubescence; abdomen squarely
truncated at end, surrounded by five lobes, presenting an eriopterine appearance;
mentuin conspicuously five-toothed
Rhamphidaria (p. 830)
Body very long and slender, glabrous; abdomen surrounded by four narrow, glabrous
lobes; mentum indistinctly five-toothed
Dicranoptycharia (p. 828)

Pupae
Pronotal breathing horns branched; forms entirely aquatic
Antocharia (p. 799)
Pronotal breathing horns simple
2
2. Pronotal breathing horns very tiny, microscopic, conical
Dicranoptycharia (p. 828)
Pronotal breathing horns large, conspicuous
3
3. Pronotal breathing horns large, earlike, contiguous basally; forms aquatic, in silken
Ellipleraria (p. 806)
cocoons
Pronotal breathing horns not contiguous basally
4
4. Cephalic crest small or lacking; pronotal breathing horns short and broad, rarely elongated;a pair of small spiracles on dorsum of eighth abdominal segment. .Limnobaria (p. 808)
Cephalic crest large, setiferous; pronotal breathing horns long and slender, cylindrical;
a pair of large spiracles on dorsum of eighth abdominal segment.
Rhamphidaria (p. 830)
1.

.

The most important literature on the tribe Limnobiini is as follows
Antocha saxicola
Anlocha sp

GeneralElliptera omissa

Thaumastoptera calceati
Limnobia quadrimaculata
Limnobia quadrimacidata

Larva, pupa, general.
Larva, pupa
Larva, pupa
Larva, pupa, general.
GeneralLarva, pupa

Limnobia bifasciata

Larva, pupa, general.

Needham, 1908a: 205.
Malloch, 1915-17 b: 236-237.
Mik, 1886 b.
Griinberg, 1910:31-32. (Copy.)
Malloch, 19 15-17 b: 226-227.
Lenz, 1920a.
Von Roser, 1834 (as annvlus)
Beling, 1873b: 590-591 (as annu-

Larva

Elliptera omissa
Elliptera omissa

.

.

.

.

lus).
..

.

Bremi-Wolf,

1846 (as xanthop-tera).

Limnobia bifasciata
Limnobia bifasciata
Limnobin decemmaculata
Limnobia decemmaculata
Limnobia flavipes
Limnobia inustu

Larva, pupa, general.
Larva, pupa
GeneralGeneralLimnobia sexpunctala

Larva

Limnobia nubeculosa
Limnobia obscuricornis.
Limnobia tripiinctata
Limnobia triocellata
Limnobia triocellata
Limnobia immatxira.

.

Loew, 1873:41.
Verrall, 1912.

BeHng, 1886: 202.
Belmg, 1886:202 (as macro-

Larva
General.

.

GeneralLarva, pupa
Larva, pupa
Larva, pupa, general.
Larva, pupa

Paste jrfk, 1909 (as xanthoptera)

De Meijere, 1916: 198-201.

.

.

stigma).
Beling, 1879: 54-55 (as nigropunctata).
Beling, 1879:56.
Beling, 1879: 55-56.
Beling, 1873 b: 591 -592.
Johnson, 1906:2.

Pupa

Malloch, 1915-17 b: 215-216.
Malloch, 1915-17 b: 216.

Libnoles perkinsi

GeneralPerkins, 1913: clxxxii (as Limno-

Discobola caesarea

Pupa

Dicranomyia trinotata
Dicranomyia trinotata
Dicraiiomyia dumetorum

Larva, pupa
Larva, pupa
GeneralMik, 1884.
Thienemann, 1909.
Grunberg, 1910:29.

bia).

Winnertz, 1853.

(Copy.)

The Crane-Flies of New York
Dicranomyia dumelorum
Dicranomyia dumelorum
Dicranomyia dumelorum
Dicranomyia sp
Dicranomyia foliocuniculalor
Dicranomyia folincuniculator
Dicranomyia umbrala
Dicranomyia simulans
Dicranomyia simidans

GeneralGeneralBeling, 1879 56.
Beling, 1886:201-202.
Schubart, 1854.
Swezey, 1913.
:

GeneralGeneralLarva, pupa

Swezey, 1915:87.

De Moijere, 1916: 197-198.

Larva
Larva, pupa, general.
Larva, pupa

.

.

Pupa

Needham, 1908 a: 214-217.
Malloch, 1915-17 b: 213-214.
Beling, 1873 b 592.
Beling, 1879:52-53.
Beling, 1879:53-54.
:

Larva, general
Larva, general
GeneralLarva, pupa
GeneralGeneral

Rhamphidia longiroslris
Rhamphidia longiroslris
Rhamphidia flavipes
Rhamphidia flavipes

799

Belinp;, 187.3 b: 592.

Larva

Rhipidia maculata
Rhipidia macidala
Rhipidia uniseriala
Rhipidia domestica
Dicranoplycha winnemana— Part II

Johnson, 1910:704.
Alexander, 1919 b.
Gercke, 1884.
Griinberg, 1910:30.
(CopJ^)
Hart, 1898 [1895] 197-199.
Malloch, 1915-17 b: 231-232.

Larva
Larva

:

Subtribe AntochariaThe

subtribe Antocharia inchides the genus Antocha and probably

three or four related genera, such as Diotrepha, Orimarga, and Orimargula.

The group is well-definedin all stages, so far as these are known, thelarvae presenting a curious superficial resemblance to those of Pedicaria,

while the pupae introduce a novelty of structure of the breathing horns,
which is discussed in detail elsewhere (page 805). It is probable, however,that these peculiarities of larval and pupal structure are largely the resultof habitand habitat, and a critical survey of the structure shows a close

relationship with the other subtribes herein recognized.

Genus Antocha Osten Sacken (Gr. close approximation)
1859
Larva.

Proc. Acad. Nat. Sci. Phila., p. 219.

A7itocha O. S.

— Body slender, tapering behind, ending caudally in two elongate ventral lobes

which bear a few hairs at their tips and at intervals along their length.
2 to 7 each with a swollen area

Abdominal segments
on basal ring densely covered with microscopic hairs.

Tracheal gills four in number, large, constricted into three or four lobes.
or rudimentary.
split behind.

Head capsule moderate in size.

Maxilla conspicuous, consisting of two subequal lobes which are provided with

dense brushes of hairs.

Pupa.
in front,

Spiracles lacking

JNlentum with nine or ten teeth, deeply— Anterior end

Hypopharynx with chitinizod teeth.
body large, tapering behind. Head with a small median lobeofon either side of which is a small

tubercle;-

genae gibbous.

horns large, flattened, the margin branching into eight long filaments.

Pronotal breathing

Abdominal segments

on basal ring with a double transverse row of small hooks which converge at the ends to incloso
an oval depressed area; last segment of body terminating in two strong, recurved, chitinizedhooks.

Charles Paul Alexander

800

Antocha is a small genus of crane-flies (about sevenspecific limits arc

still

not well ^understood.

tributed thruout the Northern Hemisphere.
itive organization,

in

many

whose

species)

The species are well disThe adult flies are of prim-

but the larvae and the pupae are highly specialized

respects.

The only previous record of the immature stages of any member of thisgroup is the unknown Limnobiine No. 2 (Malloch, 1915-17 b 236-237) which
The main point
surely refers to an Antocha, possibly A. monticola Alex.
Mallochandthe species
bydescribed
thespecies
betweenof difference:

described in detail hereinafter
description of the formerisspecies.

,the small spiracles mentioned in the

A. saxicola lacks

spiracles,

since

it

has no us3 for them, being confined to submerged cases often many feetbelow tho surfaco of the water.
Altho nothing is known concerning the immature stages of the genera
Orimargula, Orimarga, and Diotrepha, the writer believes that thesegenera will b3 found to have larvae of this same general type, since fromthe structure of the adults they are obviously derived from the Antocha

The larvae are curiously suggestive of the Pediciini (as comparedwith Dicranota), but the structure of the mouth parts, the smooth pseudostem.

and indicate that the similarities
The pupae are unlike those of any crane-fly as yet

pods, and the cauda, are quite different

are analogous only.

made known, in the very remarkable breathing horns; but this is possibly
a condition brought about by the habitat rather than a fundamental
feature.
The apparently very different pupae of Elliptera are closely
related to Antocha.

Antocha saxicola 0. S.
1S50

Antocha saxicola. 0.

S.

Proc. Acad. Nat. Sci. Phila., p. 219.

Antocha saxicola has one of the most interesting life histories of any of
the crane-flies yet discovered, not only because of the larval and pupal
habitat, but also because of the peculiar structures that appear in thelarva and in the pupa and have been found nowhere else in the immature
stages of the family, sd far as is known.

The larvae simulate strikingly

the same stage in the Pedicaria, but are apneustic, entirely lacking functional spiraclestion.

and depending wholly on tracheal gills for their respira-The pupae have the pronotal breathing horns split into eight long

filaments, so that they bear a curious superficial reseml lance to the pupae

The Crane-Flies of New Yorkof the black fly (Simuliidae),— Part II

801

Both larvae and pupae spend their entire

lives in cases on stones in water— usuallyin running, well-aerated water,and often in the most rushing torrents.

The larvae, as already stated, lack spiracles, the entire respiration being
on thru tracheal gills, four in number, and the rich tracheal
development in the elongate caudal lobes. Thus the tracheal system istruly closed, and represents the maximum of specialization in the reduccarriedtion in sizeand final loss of the spiracles.

In air-breathing, terrestrial

forms, the spiracles are large and situated comparatively close together,

gradually becoming smaller and more removed from one another as thecreature becomesgills

more and more dependent on blood gills or tracheal

for respiration.

This is the only truly closed tracheal system known

to the writer to occur in the Tipulidae.

It should be noted that the lossof the spiracles is accompanied by great enlargement of the gills and thetaking on of the gill function by the two caudal lobes.

The haunt of thelarvae is in silken cases on rocks, often in the swiftest part of the stream,

where it is impossible to breathe thru spiracles and where the associatedlife (Ephemerida, Plecoptera, Trichoptera, and Diptera) all,or practically all, depend entirely on gills for respiration.
As a rule, the larval cases are made on rubble or rounded stones, acrevice or a groove caused by inequalities of the rock surface covered
forms ofover being the simplest and commonest place chosen.
in theis

The inequalities

rock are bridged over by a silken, mud- or silt-coverod case, whichvery delicate and laterally fimbriated with the young larva but becomesmuch

thicker, and more compact with the older larva and
The larval case is open at both ends and the larva passes backward and forward freely, showing considerable agility when disturbed.
firmer,pupa.

When the larva is still small, the case isinsignificant;correspondingly

but the case of the matured larva

is

small

and

conspicuous, measur-ing from 4 to 5 centimeters in length and about 1.2 centimeters acrossthe lateral " wings," or fimbriations.
The insect moves freely alongthe

tube butisvery loath

to

leave

it

unlessactually ejected.

Asstated above, the late larval and pupal covering is very different from theyoung larva, being smooth, compact,
The pupa has two powerful hooks
at its caudal end, enabling it to fasten to the case.
In most cases the pupa
hangs with the current, head downstream, like the pupa of Blepharocera
flims3%silt-covered tube of thehard, and often covered with pebbles.

802

Charles Paul Alexander

and unlike the somewhat similar-appearing pupa of the Simuliidae,

which
head upstream, against the current, the pupal case being
open at the cephalic end only.
These curious larvae were first noted at Ithaca, New York, in midrests with the

They were common in situations such as
Larvae were found thruout most of April, May, andJune; they would probably be seen in somewhat fewer numbers thru
April,

inCascadilla Creek.

described above.

most of the summer season, since the adult flies have a long seasonal
ajjpearance.
The first pupa was found on May 15, 1917, but the season
that year was very backward and undoubtedly the species pupatesearlierin more nearly normal seasons.
In Cascadilla and Fall Creeks, at Ithaca, the immature stages of Antocha
are usually associated with a fauna of rapid-water

following being the(lotic)forms, themore notable and constant:Planarians.

Planaria sp.

Ephemeridae.

Nymphs of Baetis, Leptophlebia, Ephemerella, Ecdyurus,

Perhdae.

Nymphs of Pteronarcys,

Epeorus,Trichoptera.

Lepidoptera.
Coleoptera.
Diptera.

Iron,

Heptagenia, Chirotenetes, and others!
Perla immarginata Say, Acro-

neuria, Neoperla, and others.
Larvae and pupae of Helicopsyche (abundant), Hydropsyche, Hydropsychodcs, Ithytrichia, Rhyacophila, Leptocerus, Polycentropus, and others.
Larvae and pupae of Elophila.
Larvae of Psephenus.
Larvae and pupae of Blepharocera, Simulium, Orthocladius, Tanytarsus, and others.

Early in spring the rocks are plastered with dense coatings of Diato-

maceae (Navicula, Synedra, Meridion, and other genera), which laterin the season become much rarer or disappear entirely.
It is often impossible to tell the case of Antocha from that of some very
similar caddisworm cases, especially some of the glossosomatine Rhyacophilidae. Dr.
Noyes found larvae of Antocha in a small, rapid-flowing stream near
Ringwood Hollow, and here the cases were covered with tiny pebbles
and it was quite impossible to distinguish them superficially from associated caddis-worm cases.

In Cascadilla Creek the little cases of Helicopsyche often plaster the upper surfaces of submerged rocks, and thewriter has found tubes of Antocha that were almost buried beneaththesecases.

The Crane-Flies of New York
Antochais— Part II803

by no means confined to rapidly flowing streams, altho
The writer has foundthe insects arc very often found in such situations.

adults in his Umt traps set over the Sacandaga River, in Fulton County,

New York, where the water was very quiet and at least ten feet deep.
Needham(1908a: 169-170, 205), similarly, found adults in numbers in
New York,

his tent traps set over Beaver Meadow Brook at Old Forge,inAugust, 1905, but did not locate the larvae.

In rapid-flowing streams

the insects sometimes occur just at the surface in a few millimeters of

The immature stages seem adapted to live
water, or at greater depths.
under almost any conditions of current, from moderate pressure to situations where the water rushes by in torrents and where but few of theusual lotic organisms, such as Blepharocera, Simulium, Psephenus, andothers, can exist.

From Clemens' studies (1917:14-23) it is evident thatthe current is much more rapid just beneath the surface than at various

lower depths, so that at a depth of one foot the current velocity is only
about two-thirds of that at the surface. Thus these aquatic organisms are
not constantly and entirely subjected to such tremendous pressures as
on first sight they appear to be. Many of the larvae and pupae perish
from desiccation, due to the lowering of the stream level and the consequent exposure of the rocks on which their homes are made.
Copulation between the adult flies takes place on the exposed rocks
in and along the margins of the streams where the larvae live (Osten
Sacken, 1869:127).
life,

The eggs are deposited in the water, and the entire
The

until the emergence of the adult fly, is spent beneath the water.

whole life cycle may require a year, altho the species is possibly doubleAt a single time, and even on a single rock, larvae of various
sizes, from very small ones to those almost fully grown, may be found,brooded.

and this probably explains the long flight-period of the adult.

That thedouble-brooded remains to be proved.
The food of the larva consists of microscopic plant organisms in thespeciesisthe curious maxillae, with their dense brushes ofundoubtedly being an adaptation for this type of food.
water,

Larva.

long hairs,— Total length, 9.5-10.5 mm.
Length of caudal lobes alone, 1.1-1.2 mm.
Diameter of body, 1-1.1 mm.

Coloration of living larva, light greenish brown above, clearer greenish ventra'ly; contents
of ahmentary canal showing clearly thru the thin skin; on segments 7 and 8, two paired, brightCharles Paul Alexander804

orange bodies which are very conspicuous, these possibly being fatty in nature; welts on theabdomen dark brown; in preserved specimens general coloration fading to a dull yellow.

Form elongate(Plate XX, 57), tapering behind.

Surface with a dense, appressed pubes-

cence and scattered erect hairs. Prothorax long, narrowed in front, anterior orifice marginedwith dense, fine pubescence; sides of prothorax with numerous long, erect, pale hairs.

Mesothorax and metathorax

indistinctly

anterior annulus with a few lateral setae.

as follows:

dorsal,divided into two approximately equal annuli;First abdominal segment short, the setae arranged

two small grouped setae near posterior margin, laterad of each of these,

but in alinement, a solitary stout seta, proximad of each of these a stout seta, and cephaladof each of these another strong seta, the principal setae of each side thus forming a roughtriangle; a long, delicate pleural seta on each side; ventral, two small groups of setae, withan arrangement similar to that of the grouped s°tae of dorsum. Abdominal segments 2 to 7
each indistinctly divided into two annuli by a transverse constriction, the anterior ring about
half the length of the posterior ring and bearing medially a transverse elongate-oval (dorsal)
to short-oval (ventral)welt, covered with microscopic points; ventral welts very

convexand swollen; pleura with a long seta; posterior ring with setae arranged as described above
for first abdominal segment; setae of posterior segments of body longer, but occupying same
relative position; small solitary inner seta of dorsal posterior line becoming large and prominent on seventh segment; segment 8 with six powerful setae in alinement on ventral surface,situated at base of gills, two being pleural and four ventral in position; dorsum of segment
8 with a rounded median lobe. Gills four, long, delicate, divided into lobes by constrictions
Caudal(the two caudal lobes, as well as the gills, have taken on a respiratory function).
ventral lobes two (Plate XXI, 68) very long, parallel, with scattered setae arranged as follows: at tips, sixer seven; atabout two-thirds length, three; at about one-third length, a tuft
of from fifteen to twenty long and short setae on la,teral and dorsal faces; a small solitary seta,
dorsal in position, at base of lobe; a similar bristle on ventral face at about midlength of lobe.
Like remainder of body, Cauda covered with an abundance of delicate appressed pubescence;on dorsal side, at base of each lobe, a rounded spot, and just inside this a narrow, longitudinal
Only dorsal lobe the median one of eighth segment,
line which is destitute of pubescence.
mentioned above.

Spiracles lacking.

Head capsule (Plate XX, 58) moderate in size.
longest, outer

posterior

one forming mental plate.

Lateral plates thin, double, inner one theDorsal plate broad in front, narrowed behind,margin bluntly notched; in front of this plate, two broad plates, rather widely
Labrum broad, cephalic margin andseparated medially, presumably belonging to clypeus.
ventral face with transverse rows of short hairs.

Mentum (Plate XX, 69) conspicuous, deeply

but not entirely divided as in the Pediciini; an outer flattened, circular median
lobe whose outer face is covered with small, scalelike roughenings; behind (dorsad of) thissplit behindouter lobe thementum proper, roughly triangular in outline, conspicuous, margin with abroad, blunt, median tooth which is sometimes (Plate XXI, 66) bilobed to form two subequal
apical teeth; besides this median tooth, four lateral teeth, the outermost one broad with its
lateral angle rounded.

Hypopharynx (Plate XXI, 64) forming a ring into which ducts of sali-vary glands open; anterior part, somewhat resembling

mentum in shape, a narrow bladewith anterior margin having about eight teeth; posterior part a transverse, arcuated bandwith anterior margin having about twenty teeth.

(In the figure, the two parts of the hypo-The Crane-Flies of New York— Part II805

pharynx are shown diagrammatically and separated; in some specimens the lateral teeth are
more acute, in others they are more rounded.) Antenna (Plate XXI, 65) elongate, cyhndrical,
chitinized, apex pale, with two long, sensory setae and a few papillae.

IMandible (Plate XXI,

63 and 67) strong, flattened, with two powerful bristles on back, or scrobal region, nearbase; inner face concave, tip ending in a long tooth, dorsad of apex a single smaller tooth,

ventral cutting edge with four gradually smaller teeth, beyond the last of which the margin
iscrenuiated into four or five indistinct carunculations; viewed from inside, lateral teeth

appearing blunt.

Maxilla (Plate XXI, 6.3)

large, consisting of two elongate-oval lobes, theinner one densely hairy; palpus, borne at tip of outer lobe on ventral face, shaped like onehalf of a cylinder split lengthwise, several tiny hyaline sense pegs at apex; laterad of palpus

and nearer base of outer lobe, a small elongate sensory tubercle with hairs at apex; inner
lobe of maxilla subequal in size and length to outer lobe, but more densely hairy; on its ventral
face, four or five long sensory tubercles which are expanded at their ends into setiferous heads;
at base of maxilla, a long, slender arm with three setiferous punctures at apex and anotherpuncture at about two-thirds length; setae of this arm very long and delicate. (A dorsal
view of the larva is shown in Plate XX, 57.)
Pupa.

— Length to

tip of cephalic crest, 6.2-6.8mm.

Width, d.-s., 1.4-1.5 mm.
Depth, d.-v., 1.1-1.2 mm.

Head, thorax, and sheaths of appendages dark brown in fully colored individuals; abdomen
pale yellowish white; terminal hooks of abdomen heavily chitinized.

Head on margin above eyes with a blunt median lobe and on either side a small but promCompound eyes large, semicircular in outline. Front between

inent tubercle; gena gibbous.

eyes with margins almost parallel.

Labiallobes

broad,

Labrum with apex truncated or indistinctly bilobed.

appearing subtriangular.

Sheaths of maxillary palpi not greatly

elongated, slender, cylindrical.

Pronotal breathing horns (Plate XX, 61) flattened at base, each arcuated basally behind,bending laterad to form a concave hollow in front; base dark brown, chitinized, branched
into eight long, pale filaments which are grouped more or less in pairs; the two ventral andthe four dorsal filaments arising from a short common base, the other two being separate
for their entire length; these filaments as long as, or longer than, antennal sheaths, varying
in length from rather short to a longer type.

Thoracic dorsum broad, ample, feebly wrinkled
Leg sheaths (Plate XX, 60) with all the tarsi very long and slender, reachingalmost to end of fifth abdominal segment. Wing sheaths comparatively narrow, reachingbase of third abdominal segment; anal angle sharp; venation fairly distinct.
Abdomen pale. Intermediate abdominal segments divided into two annuli, the posterior
ring much the larger; dorsa of segments 3 to 6 (Plate XXII, 71), and sternum of segment
6, each with basal annulus tumid and with two transverse rows of small hooks converging attransversely.

the ends to inclose a linear depressed area; these areas capable of contraction, so that thehooks of each row are united or approximated with those of the opposite row; from thirty
to thirty-five hooks in each row, anterior row with hooks directed backward, posterior row

with hooks directed forward; on seventh segment, only the anterior row of hooks present, veryslightly arcuated, the lateralhooks smaller than those near middle of row; caudad of thisCharles Paul Alexander806

row on segment 7, a darkened, transversely rectangular area bearing setiferous puncturesin two broken rows, the posterior row the more complete; posterior rings of other segmentsof dorsum bearing setae in somewhat the same arrangement.

2 to 8 with a long, delicate seta on each annulus.

Pleural area pale, segments

Dorsum of segnient 8 (Plate XXII, 70) witha large setiferous tubercle on either side, this tubercle densely covered with hairs that are
longest behind and shorter in front; caudad of these large, blunt knobs, a slender, setiferous
tubercle.

Last segment with tergal valves chitinized, elongate, extreme posterior marginrounded medially and feebly bilobed, lateral angles produced caudad and dorsad into powerful curved, heavily chitinized hooks; a few setae at. about midlengthof these hooks(PlateXX, 62).

Sternal valves shorter, slightly bilobed medially.

———

Nepionotype.
Ithaca, New York, June 4, 1917.
Neanotype.
With the nepionotype.
Paratypes.
Topotypic, May 1 to June 10, 1917.

Subtribe EllipterariaisThe present knowledge of the immature stages of the genus Elliptera
due entirely to the work of Mik (1886 b). From his rather detailed

description and figures, it certainly appears that the group should receive

coordinate rank with Antocharia, Limnobaria, and other divisions herein

The genus Elliptera shows peculiarities of structure in all stages,
but many features of its organization remind one forcibly of species of
Dicranomyia (such as D. simidans and D. trinotata) on the one hand, andof Antocha on the other; and it may be that the genus Elliptera stands incloser relationship to Dicranomyia than is now believed.
created.

Genus Elliptera Schiner (Gr. I omit, or ellipse + wing)
186.3

1913

Elliptera Schin.
Wien. Ent. Monatschr., vol. 7, p. 222.
Ellipoptera Bergr. Ann. Mag. Nat. Hist., 8th ser., vol.

11,

p.

576 (correctspelling).

Elliptera is a small genus, including but five species which have a dis-continuous range, two being found in Europe and three in western North

America.

The commonestof theNorth American

species,

E. clausa

O. S., was found oh wet moss in the spray of Vernal Falls, Yosemite Valley,
California (Osten Sacken, 1877:198).

The only information available on

the immature stages of a member of this genus is that furnished by Mik(1886 b) on the European species E. omissa Egg. (quoted

subsequently

and by Malloch, 1915-17 b: 226-227). The
description and account as given below are based entirely on Mik's

by

Griinberg, 1910:31-32,

paper.

The Crane-Flies of New York— Part807

II

ElUpfera omissa Egg.
1863

Elliplera omissa Egg.

Verb. Zool.-Bot. Ges. Wien, vol.

13, p.

1108.

Specimens of ElUptera omissa were found by Mik (1886 b) along waterfalls in mountainous regions.
The adult flies werenoted as late as September 10, swarming about the waterfalls.
Larvae and pupae were found on July 30 and August 17 near Salzburg,
upper Austria, living in elongate and somewhat depressed cocoons about
10 millimeters long and 4 millimeters broad which were arranged in longiThese cocoons, which weretudinal rows with short spaces between.
placed with the current, occurred on the wet walls of wooden chutes or
runways and also on dripping chalk cliffs. The immature stages spend
When ready totheir existence in these small cocoons of mud and silk.
emerge as an adult, the pupa makes its way thru the end of the cocoon
away from the current, leaving the cast skin attached to the opening. The
margins of the large pronotal breathing horns of the pupa are finely serrated and are presumably used in making this opening thru the cocoon.
The young larva probably creeps about on the floor of the runway, feedingon algae growing in the same situation. When nearly full-grown, thelarva crawls to a less exposed place and spins its cocoon. Many larvaeand pupae are killed by the drjdng-out of their haunts when the watersupply becomes insufficient to cover them.
courses and near

Larva.

— Length, 7 mm.
Diameter, 1.5 mm.

Body clearly depressed (Plate XXIII, 72), greenish white, scarcely shiny, with delicate
appressed grayish hairs which are thicker at the two ends of the body, especially on last

segment, where they become almost villous.

Integument very transparent, so that inteson segment 6 and thru to segment 8,tine and contents show thru, the intestine narrowing

where it broadens out and almost entirely fills the ninth and tenth segments.
prothorax a delicate, long, pale hair; on remaining segments two such hairs.

On sides of
On each ofabdominal segments 3 to 9, on dorsum near anterior margin, a low transverse ridge whichisthickly set with short, blackened points; on sternum of each of same segments, a similar

welt which is destitute of points.

In male larvae, clawlike appendages of genitalia of adults

showing thru skin on ventral side.

Head capsule (Plate XXIII, 73 and 74) massive, sHghtly longer than broad, black, somewhat shinj'j all the sclerites compact and closely united; anterior projecting part of capsule with
margins transparent, rust-browTi; median part with two small knobs, laterad of which are
two larger projections which are crowned with short points; capsule weakly keeled behindon dorsum (Plate XXIII, 74), anterior to which are two swollen elevations; on hinder margin of clypeus a styliform, bristly lobe, easily broken off, which is presumably the antenna.

Charles Paul Alexander

808

Labium strongly chitinized, triangular, split longitudinally.

Mandible (Plate XXIII,

75)

clawlike in appearance, a little smaller than either half of labium, on inner face with a chiti-

nized projection which is serrated.

Maxilla indistinct, the palpi coroniform.

Spiracular disk

XXIII, 77) with four lobes whose inner faces are narrowly lined with black chitin;lobes provided with lashes of long gray hairs; dorsal lobes the shorter and broader, and bearing on their inner face two elongated stigmata which are margined with pale rust-yellow.
(PlatePupa.'— Length, G.5 mm.

Pronotal breathing horns (Plate XXIII, 76) large, ear-shaped; bright yellowish white incolor, in contrast to dirty yellowish brown skin of head, thorax,

and appendages; each hornand homogeneous; and the ventral
side, with two longitudinal furrows converging toward apices, and with abundant elongate
tubercles, under low magnification this part appearing pitted because of the spaces between
Alargin of breathing horn chitinized and very finely notched. On outerthese tubercles.
basal part of each ear a parchment-like lobe, which joins ear to side of prothorax; in addition
to this, each ear at base is drawn out into an almost rectangular lobe which is closely approximated to pronotum. Leg sheaths extending about to base of fifth abdominal segment.
Abdomen distinctly depressed, greenish white in color; segments 3 to 7 on both dorsum andconsisting of two parts: the dorsal side, appearing smooth

sternum near base with a double cross-row of spicules which present a comblike appearance
Female pupa with
(Plate XXIII, 78), those on dorsal segments being somewhat stronger.
acidothecae grown together on inner face, at outer angle of each a chitinized hook which iscurved upward. Male pupa with ventral side of last segment produced into two chitinized
points which are bent toward each other and almost touch, these inclosing clasping organsof adult male and hooks of male larva as described above.

Subtribe Limnobaria

The subtribe Limnobaria includes about ten genera, which are very
closely related to one anotherdefined.

and whose limits are as yet not clearly

Many of the generic distinctions are based on male charactersof wing form, venation, or antennal structure.

The genera are often very

and it is difficult to give satisfactory characters to separate theirimmature stages. The keys to the genera, given below, will unquestion-

large

ably needmuch revising when a larger number of forms are studied.
Larvae1.

Form stout; teeth of mandibles and of mentum usually more numerous; mentum morepointed anteriorly

Limnobia Meig. (p. 809)

Form usually more slender; teeth of mandibles and of mentum usually fewer; mentum
transverse or subtransverse
2.

2

Mandibles very broad, flattened, with three ventral cutting teeth; mentum about transRhipidia Meig. (p. 825)verse, with nine or eleven teeth
Mandibles more slender, with usually four or five cutting teeth; mentum usually a litkle
Dicranomyia Steph. (p. 819)
pointed anteriorly, with about eleven teeth

The Crane-Flies of New York— Part

II

809

Pupae
1.

The five basal abdominal segments on both dorsum and venter with a comb of small,blunt teeth; wing sheaths showing an ocellate pattern; pupae h\ing beneath bark of
Discobola O. S. (p. 815)
coniferous trees
Abdominal segments provided with basal transverse welts of microscopic points on seg2
ments 3 to 7 wing pattern not ocellate
Pronotal breathing horns long and narrow, about three times as long as broad.
Rhipidia Meig. (p. 825)
Pronotal breathing horns short and broad, length and breadth not greatly different. ... 3
Size large (usually over 10 mm.); breathing horns often broader than long.
Limnobia Meig. (p. 809)
Size smaller (usually under 10 mm.); breathing horns usually as long as broad.
Dicranomyia Steph. (p. 819)
;

2.

3.

Genus Limnobia Meigen (Gr. swamp + I live)
Nouv. Class. Mouch., p. 15 {nomen nudum, preoccupied in

Amphinome Meig.

1800

Annelida).

1803
1818
1818
1856
Larva.-

Limonia Meig.
Limnobia Meig.

Illiger's Mag., vol. 2, p. 262.
Syst. Beschr. Zweifl. Ins., vol. 1, p. 116.
Syst. Beschr. Zweifl. Ins., vol. 1, p. 116.

Unomyia Meig.
Limnomyza Bond.

Dipt. Ital. Prodr., vol. 1, p. 185.

— Form stout, terete.

Abdominal segments with dorsal and ventrai transverse
Spiracular disk surrounded by indisHead capsule large, massive, the dorsal plate narrowed

welts covered with chitinized points on basal rings.
tinct lobes, the spiracles large.

behind and more or less bifid at its tip. Labrum transversely oval, with sensory bristles
Mandible blunt, with from four to seven cutting teeth. Maxilla simple,

near margin.

cardo and stipes large.

Antenna with apical papilla button-like. Mentum broad, marginHypcpharjaix a circlet of two chitinized plates, eachwith from eleven to thirteen teeth.
with from twelve to fifteen teeth.
Pupa.-— No cephalic crest.

broader than, long.

Pronotal breathing horns flattened, earlike, as broad as, orMesonotum unarmed.

Abdomen with transverse welts of fine hooks

on basal annuli of tergites 3 to 7 and sternites 5 to 7.
eighth abdominal segment.

Two tiny;piracieson dorsum of

Limnobia is a rather small genus (comprising about forty-five species)
and handsome flies. The species are most numerous
thruout the Holarctic and Ethiopian regions. The immature stages haveof usually large

a wide range of habitat.

Of the European
Meig.]

isspecies,Limnobia

hifasciata

Schr. [= L. xanthoptera

characteristically fungicolous, the larvae occurring in various

and related genera as stated by Stannius, Pastejrik,
L. decemmaculata Lw. occurs infungi (Daedalea and similar species), as recorded b}'- Loew (1873) and by
species of Agaricus

De Meijere, and other investigators.
Verrall(1912).

L. quadriinuculaia

occurs in tree fungi but

is(Linn.)[= L.

annulus Meig.] oftenThis species,

not confined to this habitat.

Charles Paul Alexander810

L. macrostigma Schum., and L. obscuricorms Bel. are often found in decaying, principally

Fabr.

deciduous,

wood.

[=L. ru'gropimctata Schum.],

Meig. are foundinL

tn'pundata

L. Uavipes

Fabr., L.

Fabr.,

humous earth and beneathsexpimdataand L. nubeculosa

leaves in woods.

The

pupal duration of L. quadrimaculata is from eight to twelve days, and thispupates in the ground, inclosed inwhich are covered with particles of earth and otherspecies, as well as others of the genus,

delicate silken cases

matter.

In America, L.

triocellata

O. S.

is

characteristically fungicolous.

L.

and presumably L. immatura 0. S., are found both in fungi
and in decaying wood.
L. indigena O. S. has been found in living tulip
roots from Greenville, South Carolina (Greene, ms.). L. fallax Johns.,
and presumably L. solitaria 0. S., live in organic mud near water. L.
parietina 0. S. probably has a similar habitat, since it was found in tent
traps set over Beaver Meadow Brook in the Adirondacks (Needham,cinctipes Say,

1908 a: 171).

Limnohia cinctipes Say
1823

Limnohia cinctipes Say.

Journ. Acad. Nat. Sci. Phila., vol. 3, p. 21, no. 4.

Limnohia cinctipes is one of the largest and commonest American species
It has been reared many times, some of theof the genus Limnobia.

records being as follows:

On July 23, 1883, Pergande collected larvae in an old fungus groA^ing
On July 20, 1886, the same species of larvae was foundon rotten wood.

constructing silken cases thru the fungus, and later in the ground for

pupation.

The pupae were active, and were able to draw back and forthin their tubes.

Adults began to issue on July 28, showing the pupal

stage in this case to be not more than eight days.

On April 25, 1912, a number of specimens of this species were received
from W. H. Shideler, of Miami University, Ohio. The specimens weretaken at Oxford, Ohio, on April 20, when several hundred larvae andpupae were found in an old dry log. The young pupae in the wood are
not covered with particles of debris, but the older pupae are inclosed ina case which is covered with wood fragments, only the top of the head andthe tip of the abdomen projecting beyond the case. When the pupae
are about to transform, the insect emerges to about half its length and theThe Crane -Flies of New York— PartII

811

skin splits down the dorsum, the pupal skin remaining in place after the

adults have emerged.

On September 15, 1912, many full-grown larvae were found in a fleshy
Fomes near Gloversville, New York, where they were assoUla elegans and a much
When about to pupate, the larva
lesser number of Limnobia iriocellata.
species ofciated with a much larger number of larvae of

becomes pale green in color and incases itself completely in a silken sheath
which is covered with particles of sand and other debris. As the pupa
grows older, the case becomes harder and more rigid. Numerous little
mites are to be found running up and down over these pupae, more
especially at the head end, and possibly seeking ingress into the insect.
One young pupa had a piece of cloth adhering to the side of its case. The
pupal stage lasts about five days.
Larva.

— Length, 18-22 mm.
Diameter, 2.5-3.2 mm.

Coloration light yellow to greenish, the setiferous transverse welts at base of abdominal

segments brownish.

Body terete, abdominal segments subdivided into two narrow basal rings and a broad
posterior ring.

Abdominal segments 1 to 7 with a broad basal welt on tergites and sternites,that of the first segment much smaller; these welts densely covered with microscopic hooks;

on the last two thoracic segments, welts indicated by very narrow lines. Cauda blunt,
obliquely truncated.
Spiracular disk (Plate XXIV, 83, and Plate XXV, 93) surrounded
by indistinct lobes, the ventral margin projecting far caudad and indistinctly divided into
two short lobes; lateral lobes very blunt; dorsal lobes short and blunt, often divided into
two smaller lobes. Spiracles oblong or elliptical, placed obhqucly. Gills four, blunt androunded, forme-;! for propulsion rather than for respiration.

Head capsule (Plate XXIV, 70) very much as in Antocha, the dorsal plate narrowed behindand somewhat bifid at apex; lateral plates shaped like a mussel shell, curved around to form
mentum. Labrum (Plate XXV, 86) distinct, oval, the anterior margin fringed with dehcate
hairs, the hairs at the lateral margins longer and coarser; on either side near anterior margin,
a blunt tubercle with three sensory bristles; just laterad of this a stout seta; along anterior
margin, four sensory setae which are subequally spaced. Epipharyn.x densely hairy.

Clypeus broader than labrum, with a seta at each outer anterior angle and two more on
Mentum (Plate XXIV, 81) elongate-triangular, not comeither side near posterior margin.
pletely divided into halves but deeply split behind, with an outer plate running cephalad intoa long, broad point; behind this another plate with the margins toothed, there being about
Hypopharynx (Plate XXIV, 80) with two
five or six long, acute teeth on either side.

rows of teeth forming a circlet, into which duct of salivary gland opens; anterior row having
about nine large, blunt teeth, with about six smaller teeth on either side, these latter sharperpointed and more crowded; posterior row having long, pointed teeth, about twelve in

number.

Antenna (Plate XXIV, 82) two-segmented, the basal segment chitinized, elongate-

Charles Paul Alexander812

shaped somewhat like a door knob; a few sensory
Mandible (Plate XXV, 87 and 88) powerful, produced into a strong apicalpoint, with about four or five blunt or irregular inner teeth and a strong dorsal tooth onouter margin before tip. Maxilla (Plate XXV, 87) large, the outer margin thickened, subchitinized; palpi at apex small, shaped Uke half a pill box, with a few sensory papillae
cylindrical, the secona segment flattened,projections.

at tip.

Pupa.

— Length, 18-20 mm.
Width, d.-s., 2.8-3 mm.
Depth, d.-v., 3.1-3.3 mm.

Entire head and thorax, including leg and wing sheaths, light brown, the thoracic dorsum
somewhat darker-colored, the wings more yellowish brown; abdomen pale light green, thesegments with the submedian brown band interrupted on pleural region; tip of abdomen
brownish, chitinized.

Form stout (Plate XXV, 89); body destitute of noticeable setae.
Head fattened (Plate XXIV, 84). Cephalic crest lacking; forehead with a shallow V-shaped
notch between antennal bases.

Eyes

of

male

large,

the front narrowed, with points oftentorium close to inner margin of eye; eyes of female more widely separated.
rather short, ending just beyond wing root.
ous,

Labrum short, obtuse.

divergent, blunt at their tips, posterior margin a little convex medially.

maxillary palpi large, subquadrate.

Antenna

Labial lobes contigu-

Lobes ofCheek with a large, flattened ledge overlying joint offore legs.

Thorax very gibbous.

A distinct anterior median carina between breathing horns.
XXV, 90) flattened, earlike, broader than long, directed

Pronotal breathing horns (Plateslightly proximad,

margin with a row of breathing tubercles, outer face wi'inkled. Wing
Leg sheaths reaching base of fourth

sheaths reaching base of third abdominal segment.

abdominal segment or a little longer; tarsi ending about on a level, or sloping gradually fromshort hind tarsi to long foie tarsi.

Abdominal segments indistinctly subdivided

into three rings;

ontergites 3 to 7,

andsternites 5 to 7, basal ring with a transverse welt which is densely covered with short hairs

or hooks, these welts tapering gradually to ends; sternites of segments 3 and 4 having incomplete welts on either side of leg sheaths; band on tergum of segment 7 not broken medially,

but a little constricted in some specimens; in older pupae the other annuli, especially the
Female cauda (Plate XXIV, 85)posterior one, variously darkened on dorsum and venter.
with the acidothecae short, the sternal valves the shortest, the tergal valves a little longer;

prominent lateral lobes at base of tergal valves, and a slightly smaller but very broad one
on each side of tergal valves at about midlength. Male cauda (Plate XXV, 91) similar tothat of female, but the dorsal lobes (Plate XXV, 92) much shorter, not longer than the ventral lobes,

and separated by a U-shaped notch; ventral lobes approximated, each ending
Two small circular spiracles on dorsum of segment 8, these a

in a small, blunt tubercle.
littlemore widely separated in male than in female.

—

Gloversville, New York, October 26, 1012.
Nepionotype.
Female pupa with type larva.
Neanolype.
Pnratypcs.
Several larvae and pupae with types and from
1912.

—

—

-

Oxford,Ohio, April

20,

The Crane-Flies of New York— Part II

813

Limnobia fallax Johns.
1909

Limnobia fallax Johns.

Proc. Boston Soc. Nat. Hist., vol. 34, p. 125.

Limnohia fallax belongs to the solitaria group and is apparently more
Austral in its distribution than the other members of this group (L. solitaria 0. S., L. hudsonica O. S.).

Larvae and pupae were found by Dr. Johannsen near Ithaca, New York,
July 20-26, 1905. They were wrapped in silken cases covered with earthy
matter, and were removed from the soil near a brook.
Larva.

— Length, contracted, 8-8.2 mm.
Diameter, 1.2-1.3 mm.

Coloration white.

Form stout and short, body terete. Transverse welts with chitinized points on abdominal
segments 2 to 7, those on dorsal surface broad, those on ventral surface narrower. Spiracularby four indistinct lobes, the lateral pair the largest and capable ofAnal gills indistinct.
Head capsule of usual Limnobia type. Labrum (Plate XXVI, 95) broadly oval, lateralangles and disk of epipharynx with tufts of long hairs; anterior margin fringed with shortsetae; about eight sensory bristles and papillae along anterior margin.
Mentum (Plate
XXVI, 96) large, triangular, running out into a long median apical point; lateral margins
with about six or seven flattened subacute teeth on each side. Hypopharynx as in this group:a collar formed of two chitinized, comblike plates; the first plate rectangular, its face covered
disk blunt, surroundedclose approximation, closing the large yellow spiracles.

with flattened scales, anterior margin with about fourteen acute pointed teeth, the twooutermost much the smaller, acicular; the second plate a narrow band of chitin similarly
toothed, the teeth at each end large, flattened, the next tooth very narrow, acicular, theremaining teeth, ten in number, flattened, acute, the middle teeth a little shorter and broader.

Antenna (Plate XXVI, 97) with basal segment elongate, cylindrical, and apical papilla or segment very tiny, disklike. Mandible (Plate XXVI, 98) large, moderately broad, with apicaltooth prominent, two large dorsal teeth, and a row of about five or six comblike teeth along
ventral cutting edge, the most basa! being short and blunt.
Maxilla (Plate XXVI, 99)
about as in this tribe, cardines and stipites large and simple; palpus large.

— Length of cast pupal skin, 10-13 mm.

Pupa.-

Labrum (Plate XXVI, 100) triangular, apex obtuse.

Labial lobes prominent, subquadrate.

Pronotal breathing horns (Plate XXVI, 101) flattened,
subcircular in outhne, with an outer marginal row of breathing tubercles.
Leg sheaths
Posterior margin convex medially.

ending just before apex of fourth abdominal segment; tips of tarsi about on a level, or thoseBand of spicules on seventh tergite, slightly constricted
Female cauda (Plate XXVI, 102 and 103) with tergal valves a little longer thansternal valves, and more acute at their tips; a small tubercle on outer margin of outer lobes
of fore legs a little the longer.

medially.

before tips.

I

I

—
—
—

Nepionotype.
Ithaca, New York, July 21, 1905.
Neanotyve.
Cast pupal skin with type larva.
Paratypes.
One larva and three pupal skins.

Charles Paul Alexander

814

Limnohia triocellata O. S.
1S59

Limnohia triocellata 0. S.

Proc. Accad. Nat. Sci. Phila., p. 216.

Limnohia triocellata is a common crane-fly in eastern North America.
It is closely allied to the European L. bifasciata Schr., the immature stagesof which have long been known.
Johnson (1906:2) found larvae of this species in a fungus at Riverside,
Massachusetts, on August 21, 1904, which pupated on the 22d and emergedon the 30th and 31st, thus giving a pupal duration of about nine days.
Malloch (1915-17 b 215-216) found larvae and cast pupal skins in an AgariThe writer found larvae
cus at Urbana, Illinois, in September of 1915.
Fomes,
associatedwith the larvaeof Limnohia triocellata in a species ofGloversville,
New
York,
on Septemberof L. cinctipes and Ula elegans, atFalls,
Great
Virginia,
on SepC. H. Popenoe found larvae at
15, 1912.
tember 8, 1912, in the fungi Hypomyces Lactifluorum (Schw.) Tul. andArmillaria sp., the adult flies emerging on October 7, 1912. Other specimens from the same place found on October 9, 1913, in a species of ClitoScores of specimens were taken incybe, produced adults on October 20.
Boletus felleus at Bradley Hill, Maryland, the flies emerging on July:16, 1914.
Larva.

— Length, 10-18 mm.
Diameter, 1.2-1.4 mm.

Coloration alittlefallax in all details.

more yellowish than that of Limnohia fallax.

Species very close to

Ventral welt on abdominal segment 1 well developed, but dorsal welt

lacking or very reduced.

Spiracular disk (Plate XXVI, 94) rather large, exposed; circularspiracles large, separated

by a distance less than the diameter of one; lobes surrounding

disk small and indistinct.

Pupa.

— Length, 12-15 mm. Not very

different

from other speciesofgenus describedherein.

—Great Falls, Virginia, September 28, 1913.
Nepionotype.
Paratypes.
With the type.

—

Genus Libnotes Westwood (derivation obscure)f 1876

Libnoks Westw.

Trans. Ent. Soc. Lond., p. 505.

is a small genus (about thirty-five species) of rather large
which are chiefly Oriental in their distribution altho three
The species Lihnotes perspecies occur in South and Central Africa.
kinsi (Grimsh.) has been considered as being a Limnobia, but it seems

Libnotes

crane-flies,

The Crane-Flies of New York— PartII

815

to the writer that the present p;en{u-ic reference is more nearlycorrect,
altho the (hstinctions between Limncjbia and some speciesof Libnotes
are very poorly marked.
L. perkinsi was bred from larvae in damp

moss

(Perkins, 1913:clxxxii, as Limnobia), and in a letter to thewriter Mr. O.H.

Swezeystates that he has reared this species from larvae indecaying
vegetation and in the accumulation of debris behind old leaf-sheathsonbanana plants (Musa, Scitamineae) in the Hawaiian Islands.

Genus Discobola Osten Sacken (Gr. discus + I throw)
1865
1869

Discobola O. S.
Proc. Ent. Soc. Phila., p. 226.
Trochobola O. S. Mon. Dipt. N. Amer., part4, p. 98.

Discobolaisa well-marked genus including about eight described

species, which are most numerous in the Australasianregion, two species
only being found in Europe and two others in America. The

adults ofthe commoner American species, D. argus (Say), are not rare.

They aremost numerous in late summer, and are often found resting on the stumps
and trunks of coniferous trees, especially white pine {Pinus
StrobusLinn.).

The immature stages of D. caesarea (O. S.) were found by Mik
in Austria, living inremoved.

(1884)

decaying pine stumps from which the bark had been

The following account is taken entirely from Mik's paper:

Male pupa.— Body cylindrical, slender (9.2 mm. long, 1.5 mm. indiameter). Head,prothorax, mesothorax, leg sheaths, and wing sheaths chitinized,
dark brown, shiny, thelast-named somewhat brigiiter than the others, the leg sheaths
somewhat darker at theirtips.
Eyes kidney-shaped, strongly shiny, blackish, between thema small,triangular,

blackishbrown spot.

Prothoracic breathing horns dull-colored, dark rust-brown atbase,'
becoming a bri,2;hter rust-brown more distally, compressed laterally,tuberculate, with marginindented.
Prothorax carinate, rust-yellow, margined on both sides by dull
reddish browntubercles.
In fully colored specimens, forehead and leg sheaths blackish
brown, wing pattern indicated on sheaths as somewhat diffused rings.
Leg sheaths reaching end of abdo^minal
segment 3, wing sheaths reaching end of abdominal segment 1.
Metathorax and abdomenthin-skinned, the former verdigris-colored, the latter whiteor somewhat yellowish green;
metathorax resembling an abdominal segment, but its posterior margin
is unarmed, wliile
the first to the fifth abdominal segments on both dorsum andventer bear a comb of very small,short, blunt teeth, which are closely approximated;these teeth chitinized and rusty brown attheir tips, giving to abdomen the appearance of
having brown incisions; these transverserows of teeth interrupted at pleura. Sixth abdominal segment palethruout and lacking
the comb. Seventh segment shorter and narrowed on dorsum,
pale, bearing on sternum
a rust-yellow chitinized plate which is narrowed anteriorly, leaving
an uncolored triangulararea on either side at base of segment. Eighth segment
swollen to include genitalia, theCharles Paul Alexander

816

two basal parts ellipsoidal, strongly shiny, rust-yellow, somewhat darker at tips, the apical
weak triangularpiece at its base; between apical parts of genitalia are inserted two small chitinized shields;
on venter, between basal parts, sheath of penis is inserted.
Female pupa.'
Body resembling that of male, but longer and somewhat stouter (length
Leg sheaths extending to just beyond midlength of abdominal
10.5 mm., diameter 1.8 mm).
segment 2. Seventh abdominal segment shortened and somewhat narrowed, on dorsum
largely pale, with a narrow chitinized margin only on lateral parts, so that the unchitinizedpart forms a triangle with the ape.x directed backward; on sternum this segment almost
parts small and knotlike, bluntly rounded; segment bearing on dorsum a—completely chitinized, rust-yellow, only a small triangular area at base on either side remaining uncolored; chitinized plate separated from plate of next segment only by an incomplete

segmentation, swollen, and bearing two longitudinal impressions.

Eighth segment bearing
on its dorsal surface the dorsal valves of ovipositor, fused at their base, chitinized thruout,
rust-yellow in color; segment bearing on its ventral surface a depressed conical chitinizedplate of a rust-yellow color, and with transverse impressed wrinkles; on either side a small,
Other characters as in male. (When the pupae
dark, chitinized, lower valve of ovipositor.
are placed in alcohol, the green of the metathorax and the abdomen disappears and is
replaced by a yellowish white color.)

Pupae were collected in large numbers in a pine wood near Hammern(upper Austria) in the latter days of August, 1882. The
pupae live in pine stumps, near the ground, where the bark has been
removed, more especially in situations where the wood is somewhat sappyand not yet completely decayed. Those found were not deep in the wood.
Their preseiice was discovered by finding the teneral adults on and neara stump, and many cast skins of the pupae projecting horizontally, thecaudal end of the body, up to the leg sheaths, adhering to the wood. No
emergence holes were found on the cut surface of the stump. The adults
at first have a very long, pale abdomen, which is of a verdigris color,most intensive at the base and paler toward the tip. The pupae thatwere found transformed as adults in from one to three days.

in Freistadt

Genus Qeranomyia Haliday (Gr. crane + fly)
1833
1835
1838
1865

Geranomyia Hal. Ent. Mag., vol. 1, p. 154.
Limnohiorhynchus Westw. Ann. Soc. Ent. France, vol. 4, p. 683 (spurious name).
Aporosa Macq. Dipt. Exot., vol. 1, part 1, p. 62.
Verh. Zool.-Bot. Ges. Wien, vol. 15, p. 597.
Plettusa Phil.

Geranomyia is a rather extensive genus including about eighty species,
which are most abundant in the tropics of America, Asia, and Australia.
On the African continent the genus is apparently less common. The
adult flies have an elongate rostrum which is used for sucking nectar

The Crane-Flies of New York— PartII

817

from tuliular flowers. The various species of the genus have been recorded
as feeding on a wide range of plant species, which have been indicated byKnab (1910) and by Alexander (1916 b: 486^93) and may be summarized
as follows:
Plants frequented

SpeciesGeranomyia canadensis

Compositae— Eupatorium, Solidago, Aster,

Silphium, Rudbeckia, Verbesina, Cacalia,

and similar species
Compositae
Solidago, Erigeron—— Daucus
Lauraceae — Persea
Compositae — Eupatorium, Solidago, Heli-

Geranomyia diversa

Umbelliferae

Geranomyia virescens
Geranomyia roslrata

anthusFor many years nothing was known concerning the immature stages
In 1917, J. R. Malloch found larvae andpupae of G. canadensis at Urbana, Illinois. Mr. Malloch and the writer
have in press a detailed paper on the immature stages of this species, andthe following brief account is abstracted from this paper and includedof any species of Geranomyia.

herewith in order to complete the data.

Mr. Malloch found the larvae on the grounds of the Floriculture Department of the University of Illinois. There is a small bubbling fountain
here, the waste

water from which flows along an open gutter.

gutter the immature stages of G. canadensis lived

In thisamong the vegetable

growth and diatomaceous ooze in the bottom of the trough. Mr. Malloch
and the writer found this same species in Union County, southern
Here larvae and pupae occurred on the face of rocks
Illinois, in 1919.
where the surface was continually damp with percolating water. A
railroad bank had been formed by piling up slabs of limestone to a height
In the irregularities and crevices of these pieces
of about four feet.
of limestone, the larvae of Geranomyia were living in delicate silken
tubes covered with a deposit of silt and diatoms. They emerged fromtheir cases to feed on the exposed surface of the wet rocks during twilight,
and even during the hours of sunlight, but upon being disturbed or alarmed
they retreated with great agility into their tubes. The pupae are foundin short, nearly vertical burrows in the same situations as the larvae;
here they rest with only the long, conspicuous breathing horns projectingfrom the entrance to the burrow. When transformation takes place,the pupal skin projects from the mouth of the burrow nearly to the ends

818Charles Paul Alexanderof the wing sheaths.

The number of larvae vastly exceeds the number

and this would seem to indicate that the pupal existence is ofvery short duration, else this stage would be found oftener.
of pupae,

Larva.

— Length, 12-12.5 mm.
Diameter, 0.8-0.9 mm.

Coloration grayish subhyaline; a large orange area on posterior lateral parts of prothorax;

abdominal welts dark brown.

Form moderately long and slender; thoracic segments gradually decreasing in length fromprothorax to mesothorax; abdominal segments gradually elongated to the fifth, thence short-ened to end of abdomen.
inal

Ventral surface of meso- and metathorax and of first eight abdom-

segments provided with a basal transverse welt which is densely set with microscopic

points; on dorsal surface thesebands smaller, occurring on metathorax and on abdominal

segments 2 to 8, not connected with sternal bands except on metathorax and on eighth
abdominal segment. Spiracular disk similar to that in Dicranomyia; the usual ventral
lobes represented only
oval, placed obliquelyby two small, dusky, setiferous areas. Spiracles large, elongateon the sides of a deep split and so capable of close approximation.

Anal gills four, each short, tapering gradually to the blunt tip.

Head capsiile compact, massive, as in tribe.

Labrum transversely oval, margin with shortyellowish hairs and a larger tuft on either side.

Antenna two-segmented, second segment
Mandible broad andrather stout, cylindrical, slightly arcuate; apical papilla small but high.
flattened, with a small dorsal tooth and a row of five ventral teeth.

JVIaxilla generalized in

Hypopharj^nx as in Limnobaria, consisting of a roughly circularchitinized collar provided with a crown of stout teeth.
Mentum broad, undivided, anterior

structure, as in tribe.

margin with eleven teeth.

Pupa.

— Length (including breathing horns), 8-9 mm.
Length of breathing horns, 1.2-1.3 mm.

Width of body, d.-s., 0.85-0.9 mm.
Depth, d.-s., 1-1.05 mm.
Pronotal breathing horns grayish subhyaline; head and thorax with sheaths dark brown;

abdomen whitish, hooks and spines brown.
Cephalic crest small, indistinctly bilobed, not setiferous; front long and parallel; rostral

sheath very long and narrow, subtended on either side by sheaths of paraglossae, the latterprojecting beyond tip of rostrum

and ending almost opposite end of wing sheath; margin
Antennal sheaths short, ending slightly beyond baseof wing pad.
Pronotal breathing horns very large and prominent, not contiguous basally;
about a dozen breathing pores along dorsal margin. Mesonotum unarmed; wing sheathsof cheeks flattened as in Limnobaria.

ending opposite base of third abdominal segment; leg sheaths ending opposite or slightly

beyond midlength of fourth abdominal segment; tarsal sheaths ending about on a level.
Abdominal segments 3 to 7 near base with two bands of chitinized hooks arranged in curved
transverse rows inclosing an oval transverse area.
Cauda chitinized, tergal region produced
into two parallel curved hooks bending strongly dorsad.
Found at Alto Pass, Union County, Illinois, June G, 1919.

The Crane-Flies of New York— Part819

II

Genus Dicranomyia Stephens (Gr. fork +fly)
Ryst. Beschr. Zweifl. Ins., vol. 1, p. lOG (nomen nudum).
Cat. Brit. Ins., vol. 2, p. 243.
Siagona Meig. Syst. Beschr. Zweifl. Ins., vol. 6, pi. G5, figs. 5-7.
GInchina Meig. Syst. Beschr. Zweifl. Ins., vol. 6, p. 280.
Numantia Bigot. Ann. Soc. Ent. France, ser. 3, vol. 2, p. 470.

Furcomyia Meig.

1818
1829

Dicranomi/ia Steph.

18.30

18.30

1854

—

Form slender. Body nearly glabrous, abdominal and thoracic segments withand ventral transverse welts on basal annuli. Spiracular disk small, the five lobes
Anal gills four, slender. Head capsule massive,
indistinct but indicated, spiracles large.
IVIouth parts almost as in Limnobia; ventral cutting edge of manof the Limnobia type.
dible with fewer teeth; mentum with anterior margin more transverse and with fewer
Larva.

dorsalteeth.

Pupa.'— Cephalic crest lacking.

Pronotal breathing horns broad, in D. simulans witha basal recurved hook on dorsal side, in other known species unarmed.
auuuli with transverse welts.

Basal abdominal

Dorsum of eighth abdominal segment with vestigial spiracles.

Dicranomyia is a very extensive genus including more than two hundred described species of usually small flies which are found in most partsof the world.

The immature stages, which are found in a variety ofis covered by the entire family of

habitats practically as extensive as
crane-flies,

range from forms that are almost strictly aquatic, thru species

living beneath the bark of trees, to still other species which are leaf miners.

In Europe, Dicranomyia trinotata (Meig.)of the hygropetric association,isa characteristic

memberthe insects living on rocks in streams,

where they are covered with a thin sheet of water and are usually associated with such insect forms as Beraea, Tinodes, Stactobia (Trichoptera),

Orphnephila testacea (Ruthe),Dixa maculata
The larva is cylinlength and from 1.5 to

Pericoma nuhila (Meig.),

Meig., Oxycera pulchella Meig., and other Diptera.
drical,measuring from 10 to 11 millimetersin

The dorsal surface is greenish mottled withdarker, the ventral surface brighter.
The larvae live in loosely spunsilken cases in which they pupate.
The pupae are about 10 millimeters
long, and live in cocoons which are almost horizontal in position.
The
2 millimeters in diameter.

mature pupa breaks thru the cocoon by means of its sharp-edged breathinghorns, the adult then creeping forth and leaving the cast pupal hull behind.
(Thienemann, 1909:64-65, and Grunberg, 1910:29.)
Dicranomyia dumetorum Meig. lives in decaying, principally deciduous,
wood. Winnertz (1853) found it in large numbers in a decaying beech
tree, associated with Bremia cilipes (Winn.),Charles Paul Alexander

820
V

A species of crane-fly doubtfully referred to Dicranomyia pilipennis
Egg. (Schubart, 1854) has been found in ditch water in Holland.

Dicranomyia umbrata de Meij., a Javan species, lives in the slimy green
algae floating in stagnant, as well as flowing, water.

The pupae live in
The larvae

cocoons in the algal sheath, with the cephalic end projecting.

are from 10 to 12 millimeters in length, cylindrical, about 0.6 millimeterinand of a yellowish color. The head is almost entirely
The body is almost smooth, having only an inconspicuous

diameter,retractile.

transverse welt on

margin.

the s$cond abdominal segment near theposterior

The caudal end is somewhat enlarged and is truncated behind.

De Meijere (1916:197-198)

supplies a good description of

the structureof the larval head capsule.

Dicranomyia foliocuniculator Swez., of the Hawaiian Islands, is theIt was found by Swezey mining
leavesofaspeciesof
Cyrtandra(Gesneriaceae) in the island of
in theOahu. Pupation of the species takes place within the mines (Swezey,
1913 and 1915).
In North America a number of species are known.
Dicranomyia
sim,ulans has been ably discussed by Needham (1908 a: 214-217) and laterby Malloch (1915-17 b [1917]). This species is, for the most part, amember of the hygropetric fauna, dwelling in usually lotic water where
Other conit is associated with a characteristic rapid-stream fauna.
ditions under which the species is found are discussed later.
D. hadiaand D. stulta live in and under saturated moss cushions. D. macateei
Alex, has been bred from larvae in decaying wood (Dr. W. G. Dietz).
D. rara O. S. -has been bred from larvae in a rotten willow, the larvae
being taken on Plummers Island, Maryland, by H. S. Barber on October 12, 1913, and emerging as adults on November 14.
only recorded leaf-mining crane-fly.

Dicranomyia simulans (Walk.)
1848

Limnohia simulans Walk.

List Dipt. Brit. Mus., vol.

1,

p. 45.

The best account of the life history of the common and widespread
Dicranomyia simulans is that by Needham (1908 a: 214-216), quoted
below
abundant on the piers along the west shore of Lake Michigan.

This piercovered outside with heavy plank.
About three feet of surface was exposed above the water at its normal stage. The planks
were old, and sheltered a scanty growth of short, stemmed mosses in the cracks, and bore
It is[at Lake Forest, Illinois] was built on heavy driven piling,

.

.

.

The Crane-Flies of New York— PartII821

a heavy fringe of Cladophora and other algae just below the water lino, with a film of " skinalgae " extending a little higher.
All over the sides of the plank, in either sun or shade, the adult simulans could be seen
I was first attracted
throuffhou't the summer months, sometimes in considerable numbers.
to notice them by their habit of running rapidly sidewise along the pier, and their resemblance
They run habitually sidewise, apparently rarely moving
to harvestmen (Phalangidae).
forward except to escape an obstruction, and very rarely appearing on the top of the pier.
They rest in an inverted position on the under surface of the overhanging plank on the top
They stiak to the surface so persistently that it is difficult to make one takeof the pier.
flight; they may be driven all about on the surface, or poked with a stick; they can fly wellenough when they will, but when induced to f'y they settle again almost at once, and withina few feet of their starting place.
They are associated upon the piers with Geranomyia canadensis and with numerous
midges and micro-caddis flies (Hydroptilidae) and a few larger caddis flies of the genus

Hydropsyche.
Males are more in evidence, but probably not more abundant in fact. The females comeout from their resting places only to lay their eggs, and are only to be seen when busily engaged
They stand on tiptoe, with the long ovipositor held inin the performance of this task.
vertical position at the tip of the deflexed abdomen, and they swing the body up and downin rapid shuttlelike vibration, freely rising and falling on the long and widely outspread
Thus the point of the ovipositor is driven against the wet surface of the plank, thrusting^legs.
almost as rapidly as the needlebar of a sewing machine; it is moved about over the surface,
as if searching for soft spots in the wood, and occasionally it makes a deeper thrust when asuitable place is found, and an egg is deposited.
The egg-laying process is often interrupted and is continuously interfered with by the
too importunate males. When a male in running about on the plank comes upon a female
ovipositing, he stands directly above her at the full upward stretch of his legs, while shegoes right along with her work; but the instant she ceases her vibrating and lifts her ovipositor, he is ready with his forceps, upturned and outspread at the tip of his decurved abdomen, to seize her. Usually she does not want to be interrupted and moves away, while he
Oftentries to run parallel and maintain all the while his position of vantage above her.
other males are encountered, and then the males engage in a rough and tumble fight. They
push and shove each other in a most ludicrous manner, reminding one of pigs fighting,and often an encounter of this sort enables the female to escape and go on quietly withher work.

The males have well developed eyes, but their sight must be very poor; for, while always
searching for females, they seem quite unable to find them by sight, often passing females
But their tactile sense seems more acute.
at work within a distance of a few centimeters.
V/hen a male in running to and fro had passed several times within six centimeters of afemale without noticing her, was deflected from his course towa,rd her by an obstruction
I purposely placed in his way, he instantly sprang toward her upon the slightest contact,even of tips of tarsi, but was quite unheeding until this contact occurred. If it did not occur
he would pass on, even by the narrowest margin.
All stages are found together on the piers.
The eggs are laid in the soft spots in the oldwood, where the surface of the pier is kept wet, but not continually covered by water, inthe zone of the " skin algae." The larvae live exposed or thinly algae covered, and crawl
about slowly over the wet surface. They are greenish in color and very inconspicuous.
In a cavity among the stems of the dwarf mosses (Bnjnm binum Schoeb. var. varium Lindb.
and Amhlijstegium orthocladon Lesq. and James) in a crevice at the upper limit of the
wet area the larva spins about itself a sheet of tissue and fastens bits of moss stems andleaves to its outside, and transforms inside the tube thus formed into a pupa.
The tube
is longer than its body, and the pupa moves in or out at will, doubtless by the aid of thehooks at the ends of its body.

Charles Paul Alexander

822

The following descriptions of the immature stages are adapted fromthose of Needham and Malloch:
Larva.

— Length 10-15 mm.
,

Diameter, 1.5-2 mm.
Coloration green, with distinct brown marks on dorsum and with an interrupted middorsal

row of alternating paler dots and crossmarks; dark area made up of closely placed

spinous hairs, clear areas for the most part devoid of hairs.

Body cylindrical, abruptly tapering behind on last abdominal segment.

Legs lacking, but

a scurfy, pubescent creeping-fold on under .surface of meso- and metathorax, a similar one on
first

abdominal segment, and much larger, transversely placed, fusiform creeping-ridges onventral surface of abdominal segments 2 to 7, on anterior ring of each .segment.

between dorsal segmentsofIncisionsabdomen margined with blackish spinules, which are slightly

stronger than other hairs of dorsum.

Spiraculardi.skvertically cleft, with sloping sides,

folded together when under water, border of aperture fringed with short hairs and destituteof fleshy lobes.

Spiracles oval.

Anal gills four, fleshy.

Head large, similar in general appearance to that of Limnobia, entirely retractile within
enlarged prothorax; head showing a broad, pale yellow, median band, sides black from baseof antennae backward.

Ivabrum transversely oval, with a margin of close-set, scurfy hairs;

clypeus one-fourth broader than labrum, yellow, with parallel sides but emarginate on front
for reception of labrum; three recurved, stout setae on lateral margin of clypeus on each side,

Mentum slightly convex in outline, median tooth muchone on each angle and two on disk.

second and third laterals as large as median tooth.
Antenna long, the shaft about three times as long as its greatest diameter. Maxillary palpi
short and inconspicuous.
longer and stouter thanPupa.

lateral,first— Length, 8-9 mm.
Diameter, 1.5 mm.

Body smooth and shiny, ends brownish.

Front of thorax upcurved dorsally.

Pronotalbreathing horns broad, laterally flattened, obtuse at apex, each with a basal recurved, sharp

hook on its dorsal side; breathing tubercles arranged in a semicircular row along obtuse tipof horns.
Dorsum of thorax with a faint fretwork of raised lines on surface. Abdomensmooth, with transverse lines of scurfy pubescence, terminating in a pair of stout, sharply
recurved hooks.

Malloch (1915-17 b,

pi.

33,

fig.

5)

has figured the peculiar pronotal

breathing horn of this species.

Dicranomyia stulla O. S.
1859

Dicranomyia stulla O. S.

Adults of Dicranortiyia

Proc. Acad. Nat. Sci. Phila., p. 210.

stultaare often exceedingly abundant,

flyingabout, or resting in close proximity to, rocky ledges or cliffs near streams.

They are found commonly in June, associated with such crane-fly species

The Crane-Flies of New York— Part II

823

as Geranomyia canadensis, Dadylolahis montana, Tipida ignohilis, T. api-

and similar forms. The larvae live in and beneath the saturated
cushions of moss (Amblystegium) that grow on the shale near the water'scalis,

edge.

Ithaca,The only associated crane-fly larvae found near Cascadilla Creek,
New York, where this species is common, were Tipula ignohilis,

the larvae of both species being exceedingly abundant.
Larva.

— Length, 10-12.2 mm.
Diameter, 0.6-0.7 mm.

Color pale whitish with a green cast; transverse abdominal welts dark brown.

Form rather long and slender. In addition to dorsal and ventral welts on abdominal
segments 2 to 8, a complete band at base of metathorax and ventral bands on mesothorax
and first abdominal segment; ventral abdominal bands larger and more conspicuous thanA few erect setae on body. Spiracular disk (Plate XXVII, 105) rather
deep vertical split, the large ovate spiracles capable of close approximation;

narrow dorsal welts.
small, with aventral lobes blunt, with a black spot on face, fringed with short black hairs and with two

sensory setae; each blunt lateral lobe narrowly lined with a black crescent; dorsal lobes verysmall, dusky; spiracular disk fringed with short, dark hairs.

Head capsule as in tribe.

Labrum(PlateXXVII,

Anal gills four, large and pale.

106) subtriangular; anterior

marginbroad, nearly straight across, with a dense fringe of hairs which are coarser at ends oflobe; near anterior margin of labrumlae;

two oval, hyaline areas, each with three short papil-

a few sensory setae along anterior margin.

Mentum(PlateXXVII,

107)

broad,

anterior outline triangular, running out into a rather long apical point, each side with about
five teeth.

Hypopharynx (Plate XXVII,108) as in this group of genera, consisting of acollar of two chitinized combs, each with about ten sharp teeth.

short, cylindrical, the apical papilla verysmall,

Antenna (Plate XXVII, 109)
Mandible (Plate

reduced to a tiny disk.

XXVII, 110) broad, flattened, with a blunt apical point which is only a little longer than
the teeth on either side of it; ventral cutting edge with about five blunt teeth,

which aregradually smaller from the outermost toward the base; inner face of mandible with a blunt
prosthecal tooth and an oblique fringe of coarse setae.

Maxilla (Plate XXVII, 111) withthe cardines large, with about three setiferous punctures; stipites short, cylindrical; outer lobe
fringed with long hairs and bearing the short, flattened, disklike palpus, which has five or sbc

hyaline pegs; inner lobe smaller, with dense, short hairs and a few sensory organs.

Pupa.

— Length, about 6 mm.

Labrum very obtusely rounded at apex, not bilobed.
slightly

convexXXVIII,112).

acrossposterior margin.

Pronotal breathing horns

Labial lobes straight across or very
Maxillary palpi narrowed toward tips (Plate(PlateXXVIII, 113 and114)

elongate-oval,

margin more bulging, the apex a httle narrowed but obtuse; a row ofbreathing pores along outer margin, beginning on lateral face near dorsal margin, these few
in number and widely separated, becoming more numerous toward apex of organ.
Legsheaths as usual in this group of genera, those of fore legs the longest, those of hind legs
the shortest.
Male cauda (Plate XXVIII, 115) with ventral lobes (Plate XXVIII, 116)
large, bluntly rounded at tips; two small, brown, approximated tubercles at base of split on
earlike, the ventral

Charles Paul Alexander

824

ventral side; two blunt tubercles on dorsal surface near base and close to median line (Plate

XXVIII, 117).

Tergal lobes at outer angles of a flattened plate, very short, triangular, eachwith two small hairs on caudal face before tip.

— Cascadilla Creek, Ithaca, New York, May 22, 1917.
— TypeJune
1917.
Paratijpes. — Abundant larvae and pupae with types, May 22 to June
Nepionoiype.
Nennotype.

locality,

G,

6,

1917.

Dicranomyia badia (Walk.)
1848
1859

Limnohia badia Walk. List Dipt. Brit. Mus., vol. 1, p. 40.
Dicranomyia humidicola O. S. Proc. Acad. Nat. Sci. Phila., p. 210.

Dicranomyia badia

isa very common species thruoiit eastern North

America, occurring in gorges and ravines and along streams. The adultIn
flies may be found resting on perpendicular cliffs near these haunts.

a position of rest they have all six feet on the support, a very different
resting position from that of the often-associated genera Oropeza andDolichopeza.

The larvae,asisfrequent in this genus, live in and beneath moss,

especially saturated cushions ofmoss growing in or near the margins ofThey are of a clear light pea-green color, and simulate the moss
to an astonishing degree. The writer found these larvae in Needham's
streams.

Glen, Ithaca, New York, on April 16, 1917, beneath wet cushions of
Amblystegium irriguum (Hook. & Wils.) B. & S., a moss that covers
In these moss cushions the larvae
all the rocks and stones near water.
were associated with tiny larvae of Tipula ignobilis and pupae of T.
When ready to pupate, the larva spins a small, silken, siltcollaris.
covered case, which is further protected by a covering of small pieces ofmoss stems adhering to its outside, this case being hung up in the moss
The pupal durationcushion, with the cephalic end of the pupa projecting.
(April
21
28
in
the
cases
observed).
The darksevendays
tois about
very
contrastingyellowbreathinghorns,
is
pupa,
with
its
coloredhandsome.
Larva.

— Length, 10.5 mm.
Diameter, 0.6-0.65 mm.

Coloration light pea-green, abdominal welts brown; after death general color fading tovery pale greenish white.

Form slender; body terete, dorsal and ventral transverse welts at base of abdominal segments conspicuous. Spiracular disk (Plate XXVIl, 104) with lobes indistinct, suffused withdusky, lateral lobes very blunt; disk surrounded by a fringe of short, dark-colored hairs.
Spiracles very large, ovate, dorsal ends close together.
Anal gills long and slender, pale.

The Crane-Flies of New York— Part825

II

Head capsule and mouth parts similar to those of D. slulla, already described, judging fromthe scanty material of D. badia available for study.

Pupa.

— Length, 8-8.5 mm.
Width, d.-s., l.l mm.
Depth, d.-v., 1.2 mm.

Head, thorax, and appendages dark brown; pronotal breathing horns light yellow; abdomencauda chitinized, light brown.

greenish, theLabrum very broad, indistinctly bilobed at tip.
posterior margin almost straight across.
119).

Labial lobes large, broadly transverse,

Maxillary palpi broad, tips truncated (Plate XXIX,

Lateral margins of cheeks flattened into ledges.

Pronotal breathing horns large, flattened, in lateral outline (Plate XXIX, 118) subcircular ornearly so, with a row of rather widely separated breathing tubercles along margin; as viewed

from above, horns directed proximad, so as to be contiguous at tips.
on mesonotum behind breathing horns.

A high median crest

Wing sheaths ending before apex of abdominal

Leg sheaths ending far before apex of abdominal segment 4; as usual in thishind legs a little the shortest, the fore legs a little the longest. Abdominal
segments with a distinct basal welt which is thickly margined with microscopic curved hooks.
Female cauda with sternal
Lateral spiracles distinct, but small and probably nonfunctional.
segment 2.

division, thevalves shorter than long tergal valves, the latter (Plate XXIX, 120) almost straight, each witha powerful, acute spine on lateral margin at about midlength, this directed dorsad.

Near themargin of segment 8, on dorsum, a pair of rudimentary spiracles.

— Needham's Glen, Ithaca, New York, April— TypeMay 1917.

Nepio7ioiype.

Neanotype.

locality,

16,

1917. (No. 5-1917.)

7,

Genus Rhipidia Meigen (Gr. a fan)
1818
1911

Rhipidia Meig. Syst. Beschr. Zweifl. Ins., vol. 1, p. 153.
Ceratostephanus Brun. Rec. Indian Mus., vol. 6, p. 271.

— Form rather stout, body terete.

Larva.-

Abdominal sternites 1

7 with narrow transverse basal welts of chitinized points.
lobes.

to 7

and tergites 2 toSpiracular disk with indistinct

Head capsule massive, not unlike that of Dicranomyia.

Labrum broadly transverse.

Mandible very broad, flattened, with only three ventral cutting teeth. Maxilla of simple
Antenna with apical papilla or segment very flattened, disklike. Hj^popharynx
structure.
of two chitinized plates,

each with about twelve comblike teeth.

Mentuni almost transverse

across anterior margin, with from nine to eleven teeth, the outermost fused.

— Pronotal breathing horns elongate for this subtribe, about three times as long as

Pupa.-

Abdomen with transverse bands of spicules on tergites 3 to 7 and sternites 5 to 7,
and on extreme lateral parts of sternites 3 and 4.
broad.

Rhipidia is a small to medium-sized genus (about thirty-five species)
having its center of distribution in the American tropics, with somespecies occurring thruout temperate Europe and America and a less number
The genus is based on a sexual charin Africa and the Oriental region.

Chaeles Paul Alexander

826

VLctvr, the pectinate antennae of the male, and several of the species
run inconveniently close to Dicranomyia.

The immature stages are spent beneath the bark of decaying trees ordecaying vegetable or animal matter. In Europe the genotype,Rhipidia maculata Meig., has been recorded as living in old cow-manure.
inBeling found the insects in such a situation, associated with the larvae
Rhyphus punclatus (Faljr.) (Beling, 1879:52-53) and a staphylinidofbeetl(>,

Platystethus morsitans Payk. (Beling, 1873 b: 592).

R. uniseriata

was found by Beling (1879:53-54) living in decaying beech wood,in company with larvae of Xylota lenta Meig. and X. segnis (Linn.) (SyrSchin.

phidae) as well as with larvae of a tipuline crane-fly, Ctenophora.

Thelarvae of this species, like those of R. maculata, live in thin silken cases,

open at both ends and covered with particles of wood and other debris.
In America, besides Rhipidia hryanti (which is discussed in some detail)the following records of the immature stages are available: Rhipidia
maculata, recorded in Europe as living in decaying organic matter, wasfound by Needham (1908 a: 170, 204) in tent traps set over the bed ofBeaver Meadow Brook in the Adirondacks, the insects presumably having emerged from the stream bed or from the thin layers of moss covering
the exposed stones. R. fidelis O. S. was reared from larvae in decaying woodnear Ithaca, New York, by Carl Ilg.
This species belongs to the same
suljgenus (Monorhipidia Alex.) as the European R. uniseriata, which hassimilar larval habits.
R. domestica O. S. has been bred, from larvae
obtained in fermented sap of the sour gum {Nyssa sylvatica Marsh.,
Cornaceae) at Clementon, New Jersey (Johnson, 1910:704). Males andfemales of this species were bred by Popenoe at Washington, D. C,
from larvae on more or less decaying roots of taro (Colocasia antiquorum
Schott, Araceae) taken at Gough, South Carolina, on February 1, 1911.
Specimens of R. domestica emerged on August 21, 1906, at Juneau,
Alaska, in radishes infested with H-ylemyia brassicae (Bouche)
Rhipidia (Rhipidia) hryanti Johns.
1909

Rhipidia hryanli Johns.
fig.

Proc. Boston Soc. Nat. Hist., vol. 34, p. 123-124, pi. 16,

20.

Rhipidia hryanti is one of the largest and handsomest species of thegenus, andiswidely distributed thruout the eastern United States from

Maine to Texas.

The innnature stages are spent beneath the decaying

The Crane-Flies of New York— PartII

827

The material studied was collected by J, R. Malloch in
Potomac Park, Washington, D, C, on May 11, 1913, and was rearedby R. C. Shannon (No. 35-1913), whose notes on the subject are as follows:
A small colony (about seven) of tipulids were found in a hollow of a tree behind the bark.
bark of trees.

The larvae were covered with slinie, whicli gathered debris as they moved about and so

Two larvae were boiled and precase.
TJie next morning one had pupated.
served in alcohol, while two otJiers were preserved in formal. As they move about theyOn May 14, the one that had pupated on the 12th
leave a path of slime behind them.
emerged. On May 18 another imago issued.

formed a

There is an obvious error in the duration of the pupal stage as given— only two days.

It is possil^le that a pupa was included in the material
and was overlooked in its case of debris. The date of emergence of thesecond adult would show a pupal duration of six days, which is probably

nearly correct.
Larva.

— Length, 1.3.2-14 mm.
Diameter, 1 mm.

Coloration white.

Form terete, rather stout.

Stornites 1 to 7 and tergites 2 to 7 with narrow transverse wolts

on basal rings; welts yellow, provided with long, transverse rows of microscopic roughened
points, those along margins of welts coarser, those in centers very tiny; these rows interrupted along pleura. Caudal end blunt, with lobes of spiracular disk indistinct. Spiracles
large,

capable of close approximation.

Head capsule massive, of the Limnobiatype.

Labrum (PlateXXX, 121) broadly transverse,

anterior margin almost straight across, near margin two oval areas which are provided withsmall sensory papillae.

Epipharyngeal region with abundant hairs.

Mentum (Plate XXX,

122) chitinized, the anterior margin almost transverse, with a large median tooth and about

three lateral teeth on either side, the outermost of these an evident fusion of about three lesser

Hypopharynx (PlateXXX, 123 and 124) asusualin this subtribe, a collar-like structurecomposed of two parallel combs united at the ends; the larger plate broadly elongate, surface
with abundant scalelike plates, anterior margin with twelve teeth, the lateral ones narrow,the teeth gradually enlarging toward the middle, the middle pair a little shorter; second plateof hypopharynx broadly transverse, narrow, likewise with about twelve teeth, the two outermost on each side long, subacute, the middle tooth shorter and more flattened; between thetwo middle teeth a small triangular or conical point. Antenna (Plate XXX, 125) short cylindrical, the apical papilla a very flattened disk or button which is much broader than long.
Mandible (Plate XXX, 126) very broad, flattened, with the apical tooth rather long and slender; two smaller teeth dorsad and three others ventrad, the dorsal teeth blunt, the ventral
Maxilla (Plate XXX, 127) of the simple
teeth truncated, the most basal tooth very broad.
generalized structure of this tribe; cardines large, with two setiferous punctures; stipites

teeth.

weakly chitinized basally; outer lobe of ma.xilla with the large flattened palpus at its tips;
palpus surrounded by numerous long setae and having several hyaline sense pegs at its
apex; inner lobe of maxilla smaller, with numerous long hairs, especially a slitlike brushnear margin.

Charles Paul Alexander

828
Pupa.

— Length, about 12 mm.

Labruin(cast skin).

rounded at apex.

broad,

indistinctly trilobed.

Labiallobes

broadlytransverse,

caudalmarginMaxillary palpi large, flattened, apex bluntly pointed; a rather angular

tooth on margin near base (Plate XXXI,
Antennae short and stout.

128).

Cheeks produced into flattened

ledges.

Pronotal breathing horns (Plate XXXI, 129) elongate for this subtribe, about three times
as long as the greatest diameter, flattened, apical half slightly expanded, margin with tiny
tubercles.

Wing sheaths attaining base of abdominal segment 3.

Leg sheaths attainingbase of abdominal segment 5; tarsi ending about on a level, or hind tarsi a little the shorterand fore tarsi a little the longer. Abdomen with basal bands of setae on tergites 3 to 7
and sternites 5 to 7, and on the extreme lateral parts of sternites 3 and 4; these bands thickly
margined with tiny, golden-yellow hairs or points, the median part of each band naked ornearly so.
Male cauda (Plate XXXI, 130) with dorsal lobes (Plate XXXI, 131) very small,more or less flattened, divergent, rather blunt at tips; ventral lobes elongate, contiguousalong inner face.

———

Nepionotype.
Potomac Park, D. C, May 11, 1913.
Neanotype.
Cast pupiil skin, with type larva. May 14, 1913.
ParaUjpe.
One larva with type.

Subtribe Dicranoptycharia

The subtribe Dicranoptycharia, so far as known, includes only thegenus Dicranoptycha. The division is close to the Rhamphidaria butiseasily separated from it in all stages.

Genus Dicranoptycha Osten Sacken (Gr. fork + fold)
1818
1859

Marginomyia Meig.

Syst. Beschr. Zweifl. Ins., vol.

Dicranoptycha O. S.

Proc. Acad. Nat. Sci. Phila., p. 217.

1, p.

147 (nomen nudum).

—

Larva.Form very elongate, terete. Integument smooth, glassy, transparent.
Abdominal segments 2 to 8 each with a basal transverse band or area of microscopic chitinized
points on ventral surface; segment 8 with a similar band on dorsum.
Spiracular disk surrounded by four lobes, the lateral pair more slender than the blunt ventral pair; dorsal lobe
very low or lacking; a triangular brown mark on disk between spiracles. Spiracles small,widely separated.

Anal gills a fleshy protuberant ring surrounding anus.

Head capsule compact, massive, the prefrons large with a few marginal punctures; externolateral plates very broad.
Labrum large, flattened, pale. Antenna two-segmented; apicalsegment almost as long as basal segment, gradually narrowed to the blunt tip.
with a blunt dorsal and two blunt ventral teeth.

]\Iandible

Maxilla generalized in structure.

Hjrpo-

pharynx a rounded cushion. Mentum deeply split behind but not completely divided,with three principal teeth and a small reduced lateral tooth on either side.
Pupa.
Cephalic crest low, depressed, setiferous. Labrum tumid. Labial lobes oval,contiguous.
Antennal sheaths ending opposite base of wing pad. Pronotal breathinghorns microseopic, represented only by tiny triangular tubercles. Meson otum unarmed.

—

The Crane-Flies of New York— PartWing sheaths ending opposite middle of third abdominal segment.

II

829

Leg sheaths ending

opposite base of fifth abdominal segment, the tarsi terminating on a level or nearly so.

Abdominal tergites and sternites each with four transverse rows of microscopic setae; lateralspiracles on segments 2 to 7; no apparent spiracles on dorsum of segment S.

Dicranoptycha is a principally Holarctic genus including twelve known
species, six of which

occur in the United States.

The flies are of medium

and of a dull, inconspicuous appearance, and are rather difficult oftaxonomic separation. The life histories of members of this genus have
only recently been ascertained. The following notes are taken from
more detailed accounts in earlier papers by the writer (Alexander, 1919,
a and b).
The habits of the immature stages of Dicranoptycha may be briefly
summarized as follows: The larvae and the pupae live in rich humous
soil overlain with a cover of leaf mold and other vegetable debris.
Theyfrequent open woods where there is more or less shubl^ery and tall herbage.
Running streams or rills are not necessary for the development of theimmature stages. The larvae live in the uppermost zone of the soil, where
they are associated with a rather characteristic group of animal forms,
such as dipterous larvae (Sciara), beetle larvae, and centipedes. They
are characterized by the exceedingly long, slender body and the shiny
glabrous skin, and may be confused with no other dipterous larva yet
made known. The glassy appearance of the body suggests the shiny
shell of a small univalve snail, the dead fragments of which occurred insome numbers in the same situations. The larvae of Dicranoptycha
winnemana Alex., an Austral species, attain a length of from 20 to 22
millimeters, with an average diameter of only 1 millimeter.
The larvae
When ready to pupate they incase themof the genus are herbivores.
selves in compact earthen cells, which are 10 x 3.5 millimeters in size,
firm in texture, and rather thick-walled but apparently without silk.
size

There is a small opening at either end. The pupal period is about tendays or possibly a little less. Pupation takes place in the relatively chy
soil that forms the larval haunt.
The pupa of Dicranoptycha winnemana measures from 9.1 to 12.8
millimeters in total length.
The width at the wing pad is about 1.7
The diagnostic features are given above at some length
millimeters.
for the genus and need not be repeated here.
For other notes the reader
isreferred to the papers already cited.

Charles Paul Alexander

830

The adult flies of the various species of Dicranoptycha are usually
They may be found resting on the upper

abundant where they occur.

surface of leaves of shrubbery andfromwat(>r,

which

is

tall

herbage in open woods, often far

not so necessary for development as with most

species of Tipulidae.

Subtribc Rhamphidaria

The division Rhamphidaria includes the genus Rhamphidia, with the
Rhamphidia is one of the con-possible addition of a few exotic genera.

stituent genera of the former group Antochini, but differs considerablyfrom all others whose life histories have been made known. The closest
relatives of Rhamphidia seem to be the Dicranoptycharia, on the one hand,
and the lowermost divisions of the Hexatomini, on the other. The
larvae and the pupae of Rhamphidia present a curious eriopterine appearance, but their structure indicates only a distant relationship with the
Eriopterini.

Genus Rhamphidia Meigen (Gr. rostrum)
Encyclop. Method., Ins., vol. 10, part 2, p. 585.
Encyclop. Method., Index, p. 831.
Heiius St. Farg. et Serv.
Leptorhina Steph. Cat. Brit. Ins., vol. 2, p. 24.3.
Rhamphidia Meig. Syst. Beschr. Zweifl. Ins., vol. 6, p. 281.

Megnrhina St. Farg. et Serv.

1825
1825
1829
1830
Larva.

— Bodyterete.

A transverse welt, covered with microscopic scabrous points, onventral surface of basal annuU of abdominal segments 2 to 7.

by

five

Spiracular disk surrounded

short lobes which are fringed with abundant, rather long, hairs.

Head capsule

Mandible short and stout, ending in two subequal blunt
Maxilla consLstrng of two rather short, densely hairy lobes. Antenna short.
teeth.
Hypopharynx chitinized, outer margin with about a dozen teeth. Mentum not completely
Coloration of body dark brown, produced by the dense appressed
divided, with five teeth.

massive, generalized in structure.

pubescence covering it.
Pupa.-— Cephalic crest double, the anterior part low, the posterior part the larger; lobes

setiferous,

divergent.

Two setae on front between eyes.

Pronotal breathing horns longand slender, slightly curved. Mesonotum convex. Wing sheaths reaching end of second
Leg sheaths ending just before posterior margin of fourth abdominal
abdominal segment.
Abdominal
segment; all the tarsi about equal in length, or those of fore legs a little longer.
segments with two narrow basal rings and a broader posterior ring having four narrow
Spiracles rather large ontransverse bands of spicules and comparatively few setae.
pleurites 2 to 7, and a large conspicuous pair on dorsum of segment 8.

Rhamphidia is a small genus of crane-flies (about thirty-five species),
widely distributed thruout the temperate and tropical regions of the world.
The larvae of the European R. longirostris Meig. were found by Gercke

The Crane-Flies of New York— Part II(1884) in submerged stems of the water dock,831

Rumex aquaticus Linn.

The eggs are described as l^eing rather long, white, and granulate.

It issuggested that the developmental stages may be associated with water.

The two local species have been reared and are discussed herewith.
Rhamphidia mainensis Alex.
1916

Rhamphidia mainensis Alex.

Proc. Acad. Nat. Sci. Phila., p. 498-499, fig. 14.

Rhamphidia mainensis appears to be a rather uncommon form, muchrarer than

R. flavipes, theother local

species.

Adult

flies

were notuncommon in the Basin Swamp, Orono, Maine, on June 12, 1913. Thisswamp is a low, sunken area surrounded on most sides by hills, opening
into the " Basin," an affluent of the Penobscot River.

Cold springs ofwater percolate down from these hillsides, and the soil is very wet, boggy,

and richly filled with organic matter. The chief floral constituents area few elms and white birches and an abundance of alders, Spiraea latifolia
Borkh., and S. tomentosa Linn. The herbage consists of ferns such as
Onoclea and Osmunda, patches of Iris, Impatiens biflora Walt., and many
Crane-flies associated with R. mainensis on therushes and sedges.
includedmentioned
thefollowing:date
Dicr-anomyia
haeretica,
Epiphragma fascipennis, Pseudolimnophila luteipennis, P. inornata,
Limnophila fasciolata, L. macrocera, Pilaria recondita, Uloniorpha piloseUa,
Tricyphona inconstans, Erioptera vespertina, and Tipula sidphurea, also

an abundance of Ptychoptera rufocincta and Bitlacomorpha davipes.
Larvae were first found on April 20, 1917, in the dark, cold swamp
known as Larch Meadows, south of Ithaca, New York. Here theyoccurred in the thick, black, saturated organic matter comprising thesoilof the swamp.

The vegetation consists of the dominant alder {Alnus

incana [Linn.] Moench.), the poison sumac (Rhus

Vernix Linn.), andthe marsh marigold (Caltha palustris Linn.), as well as an abundanceof other plant species in lesser

numbers.

The earthy material in which

the crane-fly larvae were found was full of the organic remains of plants,

such as ferns, leaves, alder catkins, and the like.

Associated with theselarvae at this time were numerous small hydrophilid beetles, and a greatabundance of larvae of a dascillid beetle of the subfamily Helodinae,
of various sizes and ages.
Numerous tabanid and stratiomyiid larvae,
and the larvae of the crane-fl}^ species Bitlacomorpha clavipes, Pseudolimnophila luteipennis, and Tipula dejecta, also occurred.

Charles Paul Alexander

832

The larvae were conspicuous by their dark coloration and, when placedby their active, snakelike movements, in this regard being very
different from somewhat similar larvae of certain Eriopterini which theyin water,resemble superficially because of the five subequal lobes surrounding the

The indoor pupal period is six days (May 8 to 14, 1917).

spiracular disk.
Larva.

— Length, 9-1

1

.

2 mm.

Diameter, 0.75-0.9 mm.
Coloration dark brown, sutures pale; pale spots on dorsum and on sides of bod}'; spiracular
disk pale, lobes marked with darker.

Form rather stout, terete (Plate XXXI, 132).
dark pubescence.

constriction;a faint

Body densely covered with a long, appressed,

Prothoracic segment narrowed in front, long, divided into two rings bymesothoracic and metathoracic

segments gradually longer.

First

abdominal segment shorter than last thoracic segment; abdominal segments 2 to 7 long,each divided into two narrow annuli by a deep constriction which is destitute of hairs but
has just before it a sharp, transverse ridge of stiff hairs; short, incomplete ridges of thesestiffhairs on dorsum of anterior ring; anterior ring about half as long as posterior ring;onventral side of each of segments 2 to 7 on anterior ring, a transverse swelling, these swellings

becoming more convex and prominent on posterior segments; swellings appearing almost
smooth, being covered only with microscopic, roughened points; posterior ring with a sharp
ridge of hairs at about two-thirds its length.

Chaetotaxy as follows: dorsal segments withshort setae at about midlength of posterior ring; a single rather stout, black bristle on pleuraof anterior ring, immediately

above transverse swellings; a similar seta on posterior ring;

two groups of very long, delicate setae on ventral face of posterior ring, one on each side of
the ridge of erect hairs; thoracic segments approximately similar to abdominal segments, but
Last segment of body elongated, thesternal setae at about midlength even more prominent.
spiracular disk (Plate XXXI, 1.36) surrounded by fiv6 lobes; dorsal lobe the smallest, rather
blunt, irmer face with a brownish, triangular-oval mark; lateral lobes of medium length,
inner face suffused with brown, which is darkest, almost black, on lower edge of lobe; ventral
lobes the longest, inner face of each with two broad, parallel, blackish lines, separated bya somewhat narrower pale line; entire disk fringed around with long, dark-colored hairs,
which are longest near tips of lobes, where they are strongly recurved, almost pencil-like;
fringe continuous between dorsal and lateral lobes, but between ventral and lateral lobes,
and between the two ventral lobes, hairs toward base of each lobe very short to lacking;ventral lobes just before tips with a single long sensory bristle.

Spiracles large, subcircular,

Anal gills
two pairs of pale, stout, cyHndrical lobes, tapering toward tips, before whichsituated at base of lateral lobes, dark-colored, narrowly margined with pale.
consisting of

there i-s a slight constriction.

Head capsule massive and compact, of the normal generalized limnobiine type.
large,

aLabrumconspicuous, transverse, densely fringed with long hairs; on epipharyngeal region,

large,

dense tuft of moderately elongate hairs on either side of medianline.

IMentum(Plate XXXI, 133) net completely divided, but with a very deep split behind, a large median
tooth,and two smaller teeth on either side; behind (dorsad of) elongate median tooth, awhose margins extend beyond those of outer tooth. Hypo-

slightly wider flattened lobe

The Crane-Flies of New York— Part833

IIpharynx a broad semicircular band of chitin whose anterior margin is provided with
about a dozen teeth, the intermediate ones more blunt and rounded, the lateral teeth longer

and more slender.

(The antennae of this species were not distinguishable in the specimens

available, but are undoubtedly similar to those of R. flavipes described hereinafter.)(Plate XXXI, 135) of the generalized linmobiine type, short

Mandibleand stout, terminating in two

blunt teeth; ventral cutting edge with about three teeth which are gradually smaller toward

base of mandible; a dorsal row of two teeth, of which the basal one is the smaller; base ofmandible on outer face (heel) prolonged into an acute flattened blade; a conspicuous tuftof hairs at prosthecal region.

]\Iaxilla consisting of

two rather short, stout lobes which

are shorter than the mandible, densely provided with short hairs; palpi large, shaped likea half of a short cylinder split lengthwise.
Pupa.'— Very similar to that of R. flavipes, as described on the following pages.

(Thewriter has only the cast pupal skin of R. mamensis, and it seems to agree very closely withthe pupa of R. flavipes except that the lobes of the labial sheaths are longer and more pointed,
and the labrum is a little longer to provide for the longer rostrum of the former species.)——

Nepionotype.
Ithaca, New York, April 20, 1917.
Neanotype.
Cast pupal skin, May 14, 1917.

Rhamphidia flavipes Macq.
1855
1856
1859

Rhamphidia flavipes Macq. Dipt. Exot., 5th supp., p. 17.
Rhamphidia prominent Walk. Ins. Saunders, vol. 1, Dipt., p. 435.
Rhamphidia brevirostris O. S. Proc. Acad. Nat. Sci. Phila., p. 222.

Rhamphidia flavipes is a common and widely distributed fly thruout
United States and Canada. The species is characteristicof cat-tail swamps and similar situations.
It has been reared from leavesof bur reed, Sparganium, brought in by C. H. Kennedy from Ringwood
Hollow, near Etna, New York, in September. These larvae were assothe eastern

ciatedwith larvae ofPrionocera fuscipennis, likewise a characteristic

inhabitant of open swamps.

The specimens here described were taken

in a small cat-tail swamp near Bool's hillside, Ithaca,

1917.

New York, in June,

Here they were associated with a number of larvae of characteristic

swamp-inhabiting crane-flies, such as Ptychoptera rufocincta, Pseudolimnophila luteipennis, Limnophila macrocera,cornis (in moss),

Pilaria recondita,

Liogma nodi-

Prionocera fuscipennis, and Tipula tricolor.

This is unquestionably the larva that was found by Hart and doubtfullyreferred ])y him to the genus Erioptera (Hart, 1898 [1895]: 197-199, also

Malloch, 1915-17

:

237).

Later, Mik (1898 62) doubted that this belonged:

to Erioptera because of the long lashes of hairs surrounding the caudal
lobes, a character notshown by typical eriopterine larvae.

The larvaethat Hart found were living among rushes and other vegetation floatingCharles Paul Alexander834

on the surface of the water in the Illinois River.

When submerged theselarvae were very active, quite as noted in the account of R. mainensis.

— In

Larva.'

allgeneral features like larva of R. mainensis (with the material available

the writer is unable to point out differences).

measuring from 12 to 13 mm.

Present species a little larger than R. mainensis,

and 1 mm. in diameter.

Mouth parts and headcapsule almost exactly like those of R. 7nainensis, already described.

Antenna (Plate XXXI,in lengthI'M) with first segment elongate, a little enlarged toward tip, and with an elongate, thimble-

shaped papilla at tip whose surface is delicately sculptured; surrounding this papilla atof basaltip

segment, a few microscopic tubercles and pegs; on face of basal segment on

proximal half, a circular porous plate.

Pupa.— Length, 7.8-8.8 mm.
Width, d.-s., 1 mm.
Depth, d.-v., 1 1-1 2 mm.
.

.

Coloration light brown; abdomen paler, trivittate with dark brown; a broad, dorso-medianline,

and narrower, somewhat interrupted, pleural stripes; breathing horns yellow, darker

at extreme bases.

Cephahc crest (Plate XXXII, 137) low, the lobes divergent, their lateral angles with a stout
Just behind the anterior crest a much larger, low, appressedwhich is transversely wrinkled, bearing on its side a seta directed dorsad. Eyes rather

seta which is directed cephalad.
lobe,large,

widely separated by front; two setae on front between eyes.

pointed at apex.

Labrum triangular,

Front a little elongated to provide for short rostrum of adult.

of maxillary palpi long, stout, almost straight.

Sheaths

Labial palpi contiguous, lying side by side

Antennae rather widely separated at their bases, ending just beyond
wing root; in males lying across face of eye, due to the large size of eyes in this sex.

at tip of labrum.

Rlesonotum prominent, carinate medially (Plate XXXIII, 142).
slender,slightly curved,

transversely crenulated, alittleBreathing horns long andenlarged toward tips; proximo-

cephalad of base of each horn a small rounded tubercle bearing two setae.

Lateral angles

about three short setae. Mesonotum convex, transversely wrinkled, witha few very short setae. Wing sheaths ending about opposite posterior margin of second
abdominal segment. Leg sheaths parallel, about subequal in length or those of fore legs aof thorax with

httle longer, ending ju.st before posterior margin of fourth abdominal segment.
Dorsalabdominal segments (Plate XXXIII, 143) each with two narrow basal rings and a muchbroader posterior ring; ventral segments (Plate XXXIII, 144) with the two basal rings
confluent; dorsal segments with each narrow basal ring having a narrow, slightly arcuatedband of spicules, and posterior ring having a narrow basal band of spicules which is moreor less arcuated, and a straight caudal band, immediately in front of which are four groups

two setae each, or, in some specimens, with the intermediate groups reduced to a single
segments with posterior ring having a broad transverse band of spicules,
with three setae in alinement at each end of this band; closer to base of segment, twostrong setae on either side; anterior ring darkened.
Pleura with large but apparently
nonfunctional spiracles on segments 2 to 7, a strong ante-spiracular seta, and three postspiracular setae, as illustrated (Plate XXXIII, 144).
Male cauda (Plate XXXII, 139 andl'^), terminating in two acute tips which are directed dorsad; at base of these lobes onofseta; ventral

The Crane-Flies of New York— PartII

835

two blunt, approximated, parallel lobes directed cephalad; at the base of longand dorsad, bearing on its
side and near the apex a few setae.
Dorsum of segment S with four prominent lobes
surrounding a hollow, these lobes ending in acute tips, the posterior pair directed more dorsad,
the anterior pair directed more laterad, at the base of the latter a pair of spiracles. Female
Cauda similar but with acidothecae prolonged, bearing on the side prominent lateral lobes(Plate XXXII, 141), corrt-feponding to the setiferous lobe of male as described above;quadrangle of tubercles on dorsum of segment S (Plate XXXII, 138) about as in male, alldorsal face,

apical lobes, a shorter, pointed, setiferous lobe, directed laterad

the tubercles ending in chitinized points.

— Ithaca, New York, May 1917.
— Ithaca, New York, June 1917.

Nepio7io(ype.

Neanotype.
Paratypes.

11,— Pupae with neanotype.

3,-

Tribe Hexatomini

The tribe Hexatomini, as here understood, comprises a very extensive
group of mediinn-sized crane-flies, made up of the members of the formergroups Limnophilini and Hexatomini. The more generalized membersof the group are herbivorous, but the two highest subtribes, Limnophilaria
and Hexatomaria, are carnivorous in their feeding habits.
The larvae of the lower divisions have the head capsule massive andcompact, the mentum chitinized and with the anterior margin toothed,and the hypopharynx usually well preserved. The mandibles are notformed into long, curved hooks, and the maxillae are of a generalizedorganization.
The higher subtribes, Limnophilaria and Hexatomaria,
have the head capsule long and slender and very much dissected, withthe constituent plates very narrow and separate.
The mentum is very
reduced or lacking, in some of the Limnophilaria consisting of an articulatedtransverse bar which is grooved with parallel fine striae.
In the group
Ulomorphae and in the Hexatomaria the chitinized mental region isnearly,if

not quite, lacking, allowing for great distention of this region

The mandibles are powerful, curved hooks, bearing a few
about midlength. In the group Ulomorphae the mandibles
are hinged at about midlength, the basal part being deeply grooved onof the head.

teeth

at

the inner face to receive the blade in a position of rest.

The maxillae

have the outer lobe prolonged into an elongate flattened blade which
extends out of the thoracic orifice when the head is retracted.
The pupae of the Ularia and the Epiphragmaria have a pair of large
spiracles on the dorsum of the eighth abdominal segment, indicating aclose phylogenetic relationshipwith the Limnobiini.

Dactylolabis has

Charles Paul Alexander836

tho lateral abdominal spiracles protuberant,
S(\e;ment very lar^e and conspicuous.

with those of the second

Pscnidolimnophila, the Ulomorphae,

have the pronotal ])reathing horns very long and slender,
In the Hexatomaria the lateralabdominal spiracles are large and functional.

andPilariawith the tips split into divergent flaps.

The innnatun; stages of tho Polymeraria, including the single tropical
American genus Polymera Wiedemann, are unknown. Their discovery
might result in the inclusion of this subtribe with the Pediciini rather
than with the Hexatomini.
The following keys separate the subtribes of the tribe Hexatomini:
Larvae1.

Mentum completely divided, each half with seven teeth on anterior margin; hypopharynx

a semicircular chitinized ring with numerous teeth around anterior margin.
Pseudolimno'philaria (p. 848)
2
Mentum when present not divided; hypopharynx not shaped as above
3
2. Abdominal segments 2 to 7 with a basal transverse creeping-welt
4
Abdominal segments 2 to 7 without such a welt
3. Mentum three-toothed; antenna elongate, the apical segment hemispherical.
Epiphragmaria (p. 843)
Mentum nine-toothed; antenna short, subglobular, the apex with two short papillae.
Ularia (p. 838)
4. Body depressed; head capsule massive, compact; mentum heavily chitinized, sevenDaclylolabaria (p. 852)
toothed; maxilla not projecting
Body terete; head capsule of slender bars, not compact; mentum feeble, at most a narrow,
5transverse bar which is delicately striate; maxilla projecting from thoracic orifice
Limnophilaria (group Ldmnophilae) (p. 858)
5. Mental bar present
6
Mental bar lacking
6. Dorsal plates of head capsule firmly united; epipharynx and maxilla densely hairy;
Limnophilaria (group Ulomorphae) (p. 869)
mandible hinged
Dorsal plates of head capsule widely separated, at most merely contiguous behind;
Hexatomaria (p. 876)
epipharynx and maxilla not hairy; mandible not hinged
^

Pupae
1.

2.

3.

Two large spiracles on dorsum of eighth abdominal segment
No large spiracles as above

2
3
Pronotal breathing horns subchitinized, directed strongly ventrad; abdominal segments
Epiphragmaria (p. 843)
without shagreened transverse bands; pupates in wood
Pronotal breathing horns flattened, directed laterad; shagreened transverse bands on
Ularia (p. 838)
abdominal segments; pupates in earth
Abdominal segments with large protuberant spiracles, those on second segment very
Daclylolabaria (p. 852)

large

Abdominal segments without conspicuous protuberant spiracles
4. Pronotal breathing horns elongate, split into two flaps at tip
Pronotal breathing horns not split into two such flaps
5. Abdominal segments with five or six rows of setiferous tubercles.

45
6

Pseudolimnophilaria (p. 848)

Abdominal segments without such rows of tubercles.
Limnophilaria (group Ulomorphae) (p. 869)

— PartThe Crane-Flies of New York
6.

837

II

Head and thorax without spines or tubercles; lateral abdominal spiracles small.
Limnophilaria (group Limnophilae) (p. 858)

Head and thorax oftsn with tubercles on antennal scape, labrum, or mesonotal scutellum;
Hexatomaria

lateral abdominal spiracles large, functional(p. 876)

The most important literature on the tribe Hexatomini is as follows:
Ula macroptera
Ula macroptera
Ula macroptera
Ula macroptera
Ula bolilophila
Ula elegans
Ula elegans

Epiphragma picta
Epiphragma picta.
Epiphragma fascipennis
Epiphragma fascipermis
PsexidolimnophiJa luteipennis

Pseudolivvwphila luteipennis
Dactylolabis wodzickii
Dactylolahis denticulata

Limnophila ferruginea
Limnophila hyalipennis
Limnophila lineola
Limnophila lineola
Limnophila nemoralis
Limnophila ochracea
Limnophila pallida
Limnophila dispar
Limtiophila pictipennis

Limnophila pictipennis
Limnophila punctata
Limnophila puTictata
Limnophila punctata
Limnophila punctata
Limnophila punctata
Limnophila bryobia
Limnophila sinistra
Pilaria fuscipennis
Pilaria fuscipennis
Pilaria fuscipennis
Pilaria discicollis
Pilaria discicollis
Pilaria tenuipes
Pilaria tenuipes
Eriocera spinosa

Larva

Stannius, 1829: 205.

Larva, pupa, general.
GeneralGeneralGeneralLarva, pupa, general..
Larva, pupa.

Perris,1849:337-34L

Alexander, 1915 a: 2.
Pierre, 1919-20:76.
Loew, 1869:4-5.
Alexander, 1915 a.
Malloch, 1915-17b:226. (Copy.).

Bremi-Wolf 1846.

Lari^a,Beling, 1873b: 589-590.
Larva, pupa
Larva, pupa, general... Needham, 1903:281-285.
Malloch, 1915-17 b: 224-225.
Larva, pupa
(Copy.)
Hart, 1898 [1895]: 202-204.
Larva, pupa, general.
Malloch, 19 1.5-17 b: 222-223.
Larva, pupa, general.
Nowicki, 1867:340-343.
Larva, pupa, general.
Larva, pupa, general.
Mik, 1894.
Larva, pupa, general.
De Meijere, 1916:204-206.
Beling, 1886: 198-199.
Larva, pupa
Beling, 1879:54.
Pupa
Beling, 1886: 199-200.
Larva
Larva, pupa
Beling, 1886:200-201.
Beling, 1886:202.
GeneralBeling, 1873 a: 556-558.
Larva, pupa
Larva, general
Perris, 1849: 331.
Larva, pupa, general.
Beling, 1879:51-52.
Larva
Brauer, 1883:55.
Larva
SchetTer, in Rossi, 1848: 10.
Larva
Osten Sacken, 1869:201.
Larva, pupa, general.
Beling, 1886: 195-197.
Larva
Gerbig, 1913: 158-161.
General
Cameron, 1917:63.
General]\Iik, 1881 205-206.
Larva, pupa
Hudson, 1920: 33-34.
Larva, pupa
Beling, 1886: 197-198.
Larva
Gerbig, 1913: 164-166.
General
Cameron, 1917 63.
Larva
Gerbig, 1913: 163-1(54.
General
Cameron, 1917: 63.
PupaHart, 1898 [1895] 204-205.
Pupa
Malloch, 1915-17b 223-224.
Larva, pupa, general.
Alexander and Lloyd, 1914:27.

.

.

.

.

.

.

.

.

:

:

:

:

.

30.

Eriocera spinosa
Eriocera cinerea

GeneralEriocera longicornis

Pupa, general

Larva

Alexander, 1915 c 149.
Alexander and Lloyd, 1914 2123 (as longicornis).
Alexander and Lloyd, 1914 23:

:

:

27.

Charles Paul Alexander

838
Eriocera longicornis
Eriocera fultonensis

GeneralLarva, pupa, general.

Hexatoma nigra
Hexatoma mcgacera

GeneralLarva, pupa, general.
Larva, pupa, general.

.

Alexander, 1915 c: 149-152.
Alexander and Lloyd, 191-1: BOSS.

Penthoplera albilarsis

Von Roser, 18S4.
.

.

Alexander, 1915c: 141-148.
Alexander, 1915c: 152-157.

Subtribe Ularia

Tho division Ularia includes only the genus Ula.

It represents a very

primitive group of crane-flies, presumably the most generalized of theentire tribe.

The head capsule of the larva is oval and very massive,with the prefrons large and distinct.

Tht) head capsule and its arrange-

ment of setae is not conspicuously unlike that of the eucephalous familiesof crane-flies, and this group of Tipulidae is presumably not very different
from the early tipulid ancestors. The mentum is heavily chitinized, notcompletely divided behind, consisting of two plates, one behind the other.

The outermost plate terminates in three teeth, while the second plate
The hypopharynx is notchitinized.
The antennae are very small; the basal segment is nearly
The mandibles
globular, bearing at its tip two blunt, conical papillae.

furnishes three additional teeth on each side.

are slender; the ventral cutting edge has about five narrow teeth; thereisa distinct brush of hairs at the prosthecal region.

Tho maxillae areTheof a generalized type, with the palpus large, flattened, and disklike.

abdomen is provided with six creeping- welts on the ventral surface of the
The spiracular disk is moderate in size, squarely truncated,surrounded by five subequal lobes which are heavily marked with black
segments.

on their inner faces.
The pupa has the cephalicsmall setae.

The

crest small

and provided only with very

pronotal breathing horns are very long, tapering to

The abdominal segments have a basal transverse band
The dorsal spiracles on the eighth abdominalof a shagreened appearance.
the subacute tips.

segment are large and distinct.

The closest relative of the division is apparently the genus Epiphragma
But this entire group of genera (thosein the subtribe Epiphragmaria.

included in the subtribes Ularia, Epiphragmaria, and Pseudolimnopliilaria)
isnot far removed from the tribe Limnobiini.

The Crane-Flies of New York— Part II

839

Genus Ula Haliday (Gr. soft)
1833
1864

Ula Hal. Ent. Mag., vol. 1, p. 153.
Macroptera Lioy. Atti dell' Institut Veneto, ser. 3, vol. 9, p. 224.

— Body covered with a short pubescence.

Larva.-

Basal annuli of abdominal segments

2 to 7 with a transverse creeping-welt on ventral surface.
cated, surrounded

Spiracular disk squarely trun-

Head
Labrum large,

by five subequal lobes which are fringed with very short hairs.

capsule massive, prefrontal sclerite very large, tapering to a point behind.

Mandible slender; ventral cutting edge with five teeth;
Maxilla simple; palpus large. Antenna very small;Mentum not deeply divided
basal segment subglobular, with two apical sensory papillae.
conspicuous, with tufts of hairs.

a tuft of hairs on prosthecal region.

behind; outer pLate with three apical teeth; inner plate adding three additional teeth to eachside.

Hypopharynx not chitinized.

Pupa.-— Cephalic crest small, setae tiny.

subacute, flattened apices.

ment.

Pronotal breathing horns long, tapering to

Wing sheaths ending opposite base of third abdominal seg-

Leg sheaths ending at about midlength of sixth abdominal segment.

Abdominal

segments on tergites and on apical sternites with transverse bands of microscopic points,
producing a shagreened appearance; dorsal spiracles on segment 8 conspicuous.

The genus Ula includes only six described species, of which three are
European, two are North American, and one is Javanese.
The adult flies of the American species are commonest in spring andsummer. They frequent cool, shaded gorges and ravines, or darklatewoods in mountainous regions, and may be swept from beds of low vegeand other species.

tation, such as ferns, yew,

The larvae of all the known species live in various species of fungi,
but go to earth for pupation.
In Europe the common genotype,Ula macroptera (Macq.), has been
Stannius (1829:205) foundthe larvae in a species of Agaricus. Ferris (1849:337-341) furnishes abrief account of the species, from which the following notes are taken:
The fungus in which the larvae were found was Hydnum erinaceum Bull.,growing on the trunks of living oak trees. The larvae are gregarious
and frequent galleries in the fungus, along which they progress by meansof their mandibles, by their ambulatory feet, and by the short hairs thatcover the body. They were found in the month of November. A month
later they went into the earth, where they transformed as pupae.
Thepupae bear a strong resemblance to those of Limnophila, but differ in
In February and March thethe shape of the pronotal breathing horns.
pupae come to the surface of the earth and the adults emerge. The
discussed rather frequently in the literature.

Charles Paul Alexander

840

caudal end of the larva is described as having but four lobes; no mention

made of the median dorsal lobe found in the American species, and it
presumably lacking or very reduced in size. Pierre (1919-20:70) has
reared this species from larvae living in Russula nigricans Fries.
Another European species, U. bolitophila Loew, was bred from larvae
living in fungi on beech trees in Austria (Loew, 1869:4-5).
In America, the immature stages of Ula elegans have been discussed
by the writer (Alexander, 1915a), the notes given below being in part
supplementary to his earlier account.
isis

Ula elegans 0. S.
1869

Ula elegans 0. S.

IMon. Dipt. N. Amer., part 4, p. 276-277.

Ula elegans is a rather common species in mountainous regions thi'uout
the northeastern United States.

The adult flies swarm in early spring

and again in the fall. At Ithaca, New York, on May 14, 1912, the writer
found them swarming at half past four in the afternoon. There wereabout fifteen to twenty individuals in a swarm within a foot or two of
the ground. In some cases the swarms were reduced to two or three
individuals, or in a few instances to a single specimen.

In copulation

the flies rest on the upper surface of near-by plants (Symplocarpus, in the
instance cited), withallthe legs on the support.

Copulation

firm, and the insects fly for short distance^^ still united.

israther

The only crane-fly

associated with Ula at that time was Limnophila ultima 0. S.

On September 15, 1912, the writer found a fleshy species of fungus(Fomes) growing on a much-decayed stump close to the ground. This
fungus contained a number of crane-fly larvae, including about thirty-fivelarvae of Limnobia cinctipes, a lesser number of L. triocellata, and many
larvae of Ula elegans. The larvae of these species frequented the upper
mushroom and had reduced the surface to a semi-liquid
At the end of a week the whole fungus was reduced to a very
decayed condition. The fungus, which was taken at Gloversville, New
York, was then transferred to Ithaca, and was placed in large glass jars,
with sand in the bottom to take up the liquids produced by decay and
The first adults emerged on October
to provide a place for pupation.
14.
Emergence continued until the 27th, when the remaining pupae
were killed and placed in alcohol. At that time they were very dark-layers of thestate.

The Crane-Flies of New York— Part II

colored and evidently nearly ready to emerge to the adult state.

841

At

times the larval movements are very active and eel-like, but at othertimes they are very slow and sluggish. At each movement forward,
the terminal segment partly telescopes into the subterminal and is thrownviolently backward.

skin

is

left

After transforming to the adult condition, the pupal

adhering to the sand, with the posterior half, or a smaller

portion, attached, often standing quite perpendicular to the surface

andvery conspicuous.
Larva.

— Length, 8.5-11.9 mm.
Diameter, 1.4-1.8 mm.

Color dull white; head capsule very dark brownish black.

Form moderately slender (Plate XXXIV, 145), but the body not greatly elongated as in
the Eriopterini and some Limnobiini.

Integument covered with a fine, short pubescence.
Abdominal segments 2 to 7 each divided by a constriction into a narrow basal annulus anda broad posterior annulus; basal ring at about midlength and at the end with narrow transverse rows of tiny setae; abdominal segments 2 to 7 on ventral surface of basal ring with
con-picuous raised transverse creeping-welts which are covered with microscopic points;
Region around anus
eighth abdominal segment suddenly constricted before spiracular disk.
protuberant, this evidently being an aid to propulsion. Spiracular disk (Plate XXXIV,
153) surrounded by five blunt lobes, of which the dorsal one is median in position andblunter than the others; iimer face of each lobe with a conspicuous brownish black mark;a fringe -of delicate blackish hairs surrounding disk, these hairs a little longer and more
prominent at ends of lobes. Spiracles almost circular, widely separated, the distance between
them being about three times diameter of one spiracle. Disk between spiracles with an
indistinct arcuated line.

Head capsule (Plate XXXIV, 146) massive and compact, rather narrow, posterior incisions
not extending deeply into capsule.

Prefrons very large, tapering gradually to a sharp point

behind; numerous setae on prefrons and on other sclerites of capsule.
147)

large,

Labrum (Plate XLXXIV,

conspicuous; epipharynx and lateral margins strengthened by narrow bandsof chitin; lateral an<i anterior margins of labrum with a dense fringe of long hairs; an irregulartubercle on dorsal surface on either side of labrum; epipharynx with a narrow transverseband of setae; juncture of clypeus and labrum with four setiferous punctures, two on eitherside, the posterior pair a little the closer together; clypeal region with a dark transverse basal

and terminal band; base of clypeus with three setiferous punctures on either side. Mentum(Plate XXXIV, 148) almost completely undivided, the usual median split obliterated except
behind; outermost mental plate terminating in three apical teeth, the median one of which isa little shorter and broader than the other two; behind (dorsad of) outer plate, another similar
Hypopharynx not chitinized.
plate which adds three more teeth to each side of mentum.
Anteima (Plate XXXIV, 149) very small, short, cylindrical or subglobular, about as broadas long; at the tip two hyaline papillae shaped like immature mushrooms, the outer one
about twice the size of the inner one. Mandible (Plate XXXIV, 150 and 151) slender, ending
in a blunt, slightly curved, apical point; on ventral cutting edge a row of about five flattenedCharles Paul Alexander842
teeth; themore apical ones larger, the basal one very broad and truncated; two or threedorsal teeth, gradually lessening in size from tip of mandible toward base; two conspicuous
tufts of long setae at prosthecal region; base of mandible on dorsal side with a large, somewhat curved, plate overlying head sclerites. Maxilla (Plate XXXIV, 152) with cardo longand narrow, transverse, with three conspicuous setiferous punctures bearing long setae;
outer lobe short and stout, a httle narrowed to tip; palpus rather large, flattened, disklike,the truncated apex with several tiny hyaline pegs; inner lobe with a sensory bristle

andan inner fringe of rather short hairs.
Pupa.

— Length, 6.4-7 mm.
Width, d.-s., 1.2-1.3 mm.
Depth, d.-v., 1.2-1.6 mm.

Mouth parts, wing sheaths, and leg sheaths rather dark brown;thoracic

abdomen light yellowish brown; pronotal breathing horn.s dark brown,

dorsum andpassing into lightyellow on apical third or quarter; mesonotal prescutum retaining its light color even in oldpupae and those preserved in alcohol.
Cephalic crest (Plate XXXV, 155) represented only by a small bilobed protuberance
behind and between antennal bases, each lobe tipped with a very tiny seta. Labrum short,Labial lobes appearing as a large, roughly quadrate plate, a little
bluntly rounded at tip.
narrowed behind and with all angles rounded. Sheaths of maxillary palpi very long andAntennae not conspicuous, rather
slender, rather stout at base, tapering to blunt tip.
widely separated at bases, ending just before or opposite origin of wing pad.
Pronotal breathing horns (Plate XXXV, 154 and 155) long and conspicuous, cylindrical,
gradually tapering to subacute and flattened apices; horns directed laterad and cephalad,widely divergent. Thoracic dorsum without lobes or spines. Wing sheaths extending to
Leg sheaths extending to about midlength
just beyond end of second abdominal segment.
of sixth abdominal segment; tarsal sheaths ending almost on a common level, the fore legs
being a very little the longest, the middle legs a little shorter. Abdominal tergites with transverse rows of scattered punctures; on tergites 2 to 6 a transverse band of subchitinized points,
producing a shagreened appearance; these bands located on basal rings of segments excepting
the last two, which are on extreme ends of segments 5 and 6; on sternites the bands appearing
only on segments 7 and 8, the other sternites being largely concealed by the unusually long
Male cauda
leg sheaths; band on segment 8 very broad, but narrowly interrupted medially.
with ventral lobes bluntly rounded and inclosing pleural appendages of adult (in fullycolored pupae these show thru the pupal skin as from twelve to fifteen strong, chitinized
points on the inner posterior face) dorsal lobes (Plate XXXV, 157) very short and blunt, and;approximated medially; on eighth tergite a broad rectangle of five lobes, including
an anterior and a posterior lobe on either side and a much broader anterior median lobe;
closely

just proximad of anterior lateral lobes a large and distinct spiracle, in the cast skin with thelarge tracheal trunks

stillattached.

Female cauda (Plate XXXV, 156) stout, tergal valves

a little longer than sternal valves and a little upcurved.

— Ithaca, New York, October— With type

Nepionotype.
Neanotype.

14, 1912.

larva".

Paratypes.— Type locality, October 12 to 19, 1912.

The Crane-Flies of New York— Part II

843

Subtribe Epiphragmaria

So far as is known

to the writer, the division Epiphragmaria includes

The immature stages are of a very primitive

only the genus Epiphragma.

The

organization.

larvae have the headwith the posterior incisions very shallow.
of

hair.

lateraltufts

only asingle lateral

The mentumiscapsule broad and massive,

The labrumisheavily chitinized,

broad, withentire,tooth on either side of the median tooth.

with

The

not chitinized but the entire surface is provided withThe
points, somewhat as in the Dicranoptycharia.
spinousrows of tiny

hj'popharynx

isantennae have the terminal papillae hemispherical. The mandibles are
not greatly elongated; each has an apical tooth and two rows of lateral
The maxillae are very simple in structure. The abdomen isteeth.
provided with six ventral

creeping-welts.

The spiracular disk is large

and simple, with the lobes short and obtuse.
The pupa has the breathing horns heavily chitinized, tapering to acute,The spiracles on the dorsum of the eighth abdominal
slender points.
segment are large and conspicuous.
The relationships of this group are obviously with the Rhamphidaria of
the Limnobiini and the Ularia of the Hexatomini, the three groups probably

being closer together phylogenetically than their arrangement in tribes

From a study of the adults alone, there seems to beconnection between the genera Rhamphidia, Epiphragma, and Ula,
which were placed in three widely separated tribes by Osten Sacken. A
would indicate.

littlestudy of the immature stages, the structure of the larval head, the abdomcreeping-welts, and the dorsal spiracles of the eighth abdominal

inal

segment of the pupa, indicate a relationship that cannot be denied.

Genus Epiphragma Osten Sacken(Gr.

upon + partition)

Proc. Acad. Nat. Sci. Phila., p. 238.

1859

Epiphragma 0. S.

Larva:— Body nearly smooth; basal annuli of abdominal segments 2 to 7 with a naked

transverse creeping-welt on ventral surface.

the dorso-median lobe often reduced.

Head capsule massive.

Spiracular disk large, with four or five lobes,

Spiracles large, circular.

Anal gills four, retractile.

Labrum broad, with lateral tufts of hairs and with two setae on

Mandible with a blunt apical tooth and two rows of two lateral teeth.
Antenna
stiff hairs.
two-segmented; basal segment elongate, apical papilla hemispherical. Ilypopharynx nota slender median tooth, and a broad,chitinized.
Mentum entire, with only three teeth
anterior margin.

Maxilla small, simple; palpus large; inner lobe with a dense tuft of—

fat tooth on eitherside.

Charles Paul Alexander

844— Cephalic crest erect, rectangular, the outer lateral angles produced into strong,

Pupa.-

chitinized spines.

Pronotal breathing horns large at base, the elongate tips nearly chitinized,

tapering into subacute points.

segment.

Wing sheaths ending

opposite base of third abdominalI^eg sheaths terminating just before end of fifth abdominal segment.

Abdominalband of spicules at caudal margin; pleurites with a transverse
oval area of similar spines; terminal sternites with a similar transverse band of spicules,
tergites vn\\\ a transverse

most powerfully developed on segment 8; dorsal spiracles on segment 8 conspicuous.

Epiphragma is a rather small genus (twenty-five species) of usuallylarge and handsome crane-flies, finding its center of distribution in the

New World.

The adult flies of the local species
Epiphragma fascipennis, the best known of these
are usually common.
species, is often found swarming in shady places, the swarms usuallytropical regions of theconsisting offrom twelve to fifteen individuals.

The immature stages of all the species thus far known are spent in decaying wood.

In Europe, Beling (1873b: 589-590) records finding larvae andpupae of E. pida (Fabr.) in decaying ash (Fraxinus) and beech (Fagus)
He gives the pupal duration of this species as about two
in the spring.
Needham (1903:281-285) -gives an excellent account of E.
weeks.
fascipennis, which he found in partly decayed stems of willow and buttonbush.
The same species has been found living in the stems of black
ash and elm. E. solatrix has been found in various deciduous trees ina dead or decaying condition. Bruch {in litt.) records an Argentinian
Epiphragma from decaying wood.

Epiphragma solatrix
1859(0. S.)

Limnophila (Epiphragma) solatrix 0. S.

Proc. Acad. Nat. Sci. Phila., p. 238.

Epiphragma solatrix is a beautiful crane-fly, a little more Austral in
H. S.
cUstribution than the more widely distributed E. fascipennis.
Barber found larvae and young pupae very numerous in a drift log ofsycamore (Platanus) at Plummers Island, Maryland, on May 19, 1913,
and it is the study of this material that is included in this paper. Shannon
found larvae in rotten wood at Rosslyn, Virginia, on November 23 andLater he reared both E.
25, 1912, and a pupa on January 3, 1913.
solatrix and E. fascipennis from larvae taken from the same log on thesame day. The species was again found in a decaying maple log on the
Potomac River opposite Plummers Island, Maryland, where it was associ-itsThe Crane-Flies of New York— Part II

845

ated with the larvae of the syrphid fly, Temnostoma bomhylans (Fabr.)
and the supposed larva of Protoplasa ([). 770).
Larva.

— Length, 18 mm.
Diameter, l.G mm.

Coloration pale whitish; anterior segments of body brighter-colored, more reddish.

Pronotum longer than other
Integument covered with a short appressed pubescence. Abdominal
segments 2 to 7 with conspicuous white creeping- welts on ventral surface of basal ring; there

Form stout and plump, not narrowed at ends of body.

thoracic segments.

Welts not interrupted medially, and entirely naked.

Spiracular disk (Plate XXXVI, 163)

unmarked, surrounded by five short lobes, the ventral pair very widely separated;dorsal lobe broad and obtuse; entire disk fringed with very short, delicate hairs which areinterrupted only between paired lobes; inner face of lobes suffused with pale brown at tips.
Spiracles large, circular, reddish with a very broad yellow margin, separated by a distancelarge,

nearly equal to two times the diameter of one spiracle.

Anal gills four, retractile.

Head capsule (Plate XXXVI, 158) broad and massive, the constituent plates firmly united
except on ventral parts. Labrum broadly transverse; a median epipharyngeal part whose
surface is finely pitted, bearing on ventral side near margin two stout setae; lateral angles oflabrum directed proximad and densely tufted with hairs; extreme lateral margins of labrumwith a curved hyaline seta; juncture of labrum and clypeus with two large setiferous punctureson either sid?; a few setiferous punctures on head capsule.

Mentum (Plate XXXVI, 159)

heavily chitinized, the outer face terminating in a single median tooth, behind which is a

Prementum (Plate
mentum, a moderately broad transverse plate who£etridentate plate with three flattened teeth, the lateral ones very broad.

XXXVI, 160)

lying just behindanterior margin is deeply and almost squarely notched, the lateral lobes thus formed being

rounded or feebly indented at their tips. Hypopharynx (Plate XXXVI, 161) lying above
and connected with prementum, consisting of a large flattened lobe, whose outer margin
is evenly rounded, the surface densely set with longitudinal rows of small, subacute papillae.
Antenna short, two-segmented; basal segment cylindrical, with auditory plate almost basal
Mandible moderate inin position; apical segment small, subglobular or hemispherical.
size,

broad at base with a blunt apical tooth and a few blunt lateral teeth, two on dorsal

cutting edge and two on ventral cutting edge, the more basal of these latter tending to beevanescent; two stiff setae at heel of mandible; a triangular lobe on dorsal face which
projects onto lateral parts of labrum.

iMaxilla (Plate XXXVI, 162) primitive in structure;

cardo elongate, transverse, with four setiferous punctures bearing powerful setae; outer lobe
glabrous, with palpus at its tip; palpus slightly elongate, the auditory plate just before
midlength; a seta on outer lobe below palpus; inner lobe of maxilla with a dense terminal

brush of stiff yellow hairs and a few sensory papillae.

Pupa.

— Length, 12-12.5 mm.
Width, d.-s., 2-2.1 mm.
Depth, d.-v., 2-2.1 mm.

Head, thorax, and appendages brown, becoming darker with age; pronotal breathinghorns light red; abdomen whitish, the terminal rows of spicules on the segments chitinized.

Charles Paul Alexander846(In older specimens the wing pattern shows on the sheath and is not distinctly fasciate asin E. fascipe7inis.)

Cephalic crest (Plate XXXVII, 166) erect, rectangular, the outer lateral angles directedventrad and laterad as powerful chitinized spines, which in ntiost cases curve outward; aLabrum subtruncatedor indistinctly bilobed at apex.

Labial lobes triangular, projecting caudad as two cones.

Maxillary palpi bent backward,

stout seta beneath each spine.

Cheeks projecting
on scapal segment with a flattened
antennae extending rather far beyond wing

lying along margin of cheek, stout, tapering suddenly to blunt tips.

as flattened ledges.

Antennae moderate

tubercle on either sideof

cephalic

increst;

length,root.

Pronotal breathing horns (Plate XXXVII, 164) with base enlarged, the elongate, subchitinized tips gradually narrowed,

and the extreme apices subacute; breathing horns at tips

tirected ventrad and proximad; a truncated setiferous tubercle before each breathing horn.

Mesonotum very deep.

A short, but high, median crest behind breathing horns.

angles of thorax produced into a slender lobe bearing a stout seta at tip.

ending opposite base of third abdominal segment.

LateralWing sheaths

Leg sheaths terminating just before endof fifth abdominal segment; middle tarsal sheaths usually a little shorter than the others; insome specimens, especially females, fore legs the shortest, hind legs the longest. Abdominal
XXXVII, 167) with two narrow basal rings and a broader posterior ring.

segments (Plate

Tergites on segments 2 to 7 near caudal margin of posterior ring with a narrow transverse

crossband densely beset with acute spicules; on posterior segments these bands becoming
gradually weaker, on segment 7 being very weak; two setae near lateral margin of posterior
ring at about midlength.

Pleurites with a transverse oval area with about twenty sharp spines

near caudal margins of segments; a stout seta lying cephalad of this area.

Spiracles distinct,

on ventral cephalic part of pleural posterior ring; a seta near dorsal margin of
second basal ring. Sternites with a tran.sverse band of spicules on segments 5 to 8, theseoval, lying

being weakest on segment 5 at ends of tarsal sheaths and very strong and powerful on segment 8, ^^fhere the band is interrupted medially; band often rather narrow but sometimes
much broader. A seta near midlength of posterior ring on either side. Male cauda (PlateXXXVIII, 170) with dorsal lobes very stout, globose at base but soon narrowed into slender
cylindrical appendages, each bearing three setae at tip;viewed from above, these lobes areseen to be separated by a wide U-shaped notch; ventral lobes blunt and with a flatteneddepressed disk at base of incision; eighth segment with a wide rectangle of lobes,

twoand two smaller anterior lobes with alarge

posterior lobes that are weakly
spiracle

proximad of each.

setiferous

Female cauda (Plate XXXVIII, 171) with tergal valves a little

longer than sternal valves, upcurved, and with a stout seta before tip.

———

Plumraers Island, Maryland, May 19, 1913.
Nepionotype.
With type.
Neanotype.
Paratypes.
Larvae and pupae with type.

Epiphragma fascipennis (Say)
1S23
18.59

1869

Limnobia fascipennis Say. Journ. Acad. Nat. Sci. Phila., vol. 3, p. 19.
Limnophila (Epiphragma) pavonina O. S. Proc. Acad. Nat. Sci. Phila., p. 239.
Epiphragma fascipennis O. S. Mon. Dipt. N. Amer., part 4, p. 194.

The Crane-Flies of New York— Part II

847

Epiphragma Jascipennis is a very common crane-fly thruout eastern
North America. As already stated, the immature stages are spent indecaying or partly sound wood, a wide variety of deciduous trees andshrubs being chosen, such as willow, elm, ash, buttonbush, and others.
Needham gives the pupal duration as about twelve days, larvae andpupae found on May 18 emerging as adults on the 30th. A fully grownlarva that the writer found in a decayed log beneath moss at Ithaca,
New York, on May 8, 1917, pupated early in the morning of the 10th.
The specimen died on the 18th, when about to emerge, and this would
give a much shorter pupal period than is generally recorded for the genus.

was noted at the same time that the larva superficially resembles theit wasassociated but from which it is easily distinguished by its massive headItlarva of the leptid fly Chrysopila thoracica (Fabr.), with which

capsule.

Needham found abundant pupae in a decaying log of black ash (Fraxinus)
near Freeville, New York, on May 6, 1915. The pupae occurred in burrows in the semi-decayed wood.
material was found at

Adults emerged on the 11th.

Additional

Mud Creek, near Freeville, on May 15, 1915, inelm (Ulmus).

The accountof the habits of the larvae as observed in Illinois

byNeedham (1903:281-285) is here quoted in part:
The larvae bore in the dead and fallen stems of buttonbush and willow, where these lie
on the mud at the borders of shallow ponds. I found them always in stems that were still
partially sound, tunneling beneath the bark or even into the deeper parts and into thesounder wood. These stems are frequently submerged in spring and autumn, and evenLa summer, when the pond has gone dry, ihey are always saturated with moisture
The most interesting thing about the larva, aside from its wood-boring habits, is its singular
adaptation to amphibian life. It must needs live part of the time wholly submerged beneaththe waters of the pond, and part of the time out on land; it has, therefore, both open spiracles
and tracheal gills; and, moreover, its tracheal gills are so placed that they may be withdrawn into the body in a dry time, where they escape the ills of too rapid evaporation.

In his description of the immature stages, Needham points out aprobable error of Beling in describing a sexual dimorphism in the larvaeof a species of this genus— Beling stating that the larvae producing

females have three caudal lobes while those producing males have

five.

Malloch (1915-17 b: 224-225) cites Needham's descriptions of this species.

—

Larva.
(No larvae are available to the writer for a comparison with this stage of
Epiphragma solatrix, but from Needham's characterization, and manuscript notes on specimens taker at Ithaca, New York, by the writer, the following differences seem to hold):Charles Paul Alexander

848

Spiracular disk surrounded by but four lobes, the dorso-median lobe being very reduced;
inner faces of these lobes, as well as disk itself, entirely unmarked with darker.

— Similar to pupa of Epiphragma

Pupa.'

Cephalic crest (Platedirected ventrad

orfsolatrix,

XXXVII, 165) low,slightly

but differing as follows

the lateral horns shorter,

downward; setae on

lateralface ofnot elevated, andspines often projecting

above (cephalad of) it. Pronotal breathing horns shorter and stouter than in solatrix.
Spines on eighth abdominal sternite widely separated or interrupted on median line. Male
Cauda (Plate XXXVIII, 169) with dorsal lobes much stouter than in solatrix.

— Bool's
Ithaca, New York, May
1917.
— Pupa, Mud Creek,
New York, May
1915.

Neanotype.
Paratype.

hillside,

8,

Freeville,

15,

The type pupa has one of the pronotal breathing horns deformed andbent caudad so as to be appressed against the body. Needham, who
has a very much larger series available for study, notes this same peculiarity

when he states that "a crumpled horn on one side is of rather common
occurrence" (Needham, 1903:284).
Subtribe PseudolimnophilariaAsat present constituted,

the division Pseudolimnophilaria includes

the single genus Pseudolimnophila, but other groups

may be added towhen the immature stages of other species of the old genus Limnophila
are made known.
The larva is of a distinctly primitive type, the headitcapsule being compact and massive, thepletely divided,

mentum chitinized and com-each half provided with seven or eight teeth.

The

hypopharynx is a heavily chitinized semicircle with numerous teeth aroundthe anterior margin. The antennae bear two elongate apical papillae.
The mandibles each have two blunt apical teeth and two rows of lateral
teeth.

The pupa is similar to that of the Limnophilaria, but the abdominal
segments have transverse rows of setiferous tubercles.
The larva shows many points of resemblance to the tribe Limnobiini,and the two groups are probably derivable from an immediate common
ancestor.

Genus Pseudolimnophila Alexander (Gr. false + Lwinophila)
1919

Pseudolimnophila Alex.

Cornell Univ. Agr. E.\p. Sta., Mem. 25, p. 917.

Larva.'— Body covered with delicate appressed hairs and numerous erect setae.

Spiracular

disk surrounded by four lobes, the ventral pair very long and narrow, with long fringes of
hair.

Gills

four,

long

and

slender.

Headcapsule massive,

the sclerites large,

fused.

The Crane-Flies of New York— PartII

849

I

Mandible short and blunt, of the generalized liranobiine type, the two apical teeth blunt.
Maxilla not greatly projecting. Antenna slender, with two elongate apical papillae. Hypo-

Mentum completely

pharynx a chitinized band, with numerous teeth along anterior margin.
divided, each half with about seven teeth.
Pupa.

— Cephaliccrest

tippedwith long setae.

Pronotalbreathing horns elongate,

Abdominal segments with transverse rows of setiferous
Lateral spiracles
tubercles, there being five such rows on the tergites and six on the sternites.
tips deeply split into divergent flaps.

protuberant.

The name PseudoUmnophila was proposed for a certain group of the
The adult

old genus Limnophila, inckiding L. luteipennis and its alHcs.
flies

closely resemble species of Limnophila, but the immature stages arevery different and of a distinctly more generalized type.
The larvae are herbivores, with the mentum heavily chitinized, completely divided into two halves, the margin conspicuously toothed, thehypopharynx heavily chitinized and with numerous teeth, and the mandiThe points of resembles short and blunt and with many obtuse teeth.
blance to the Limnobia type are numerous, and the head capsule shows

few features in common with Limnophila, in the restricted sense. The
pupae have the abdomen with numerous transverse rows of setiferous
The genotype is Limnophila luteipennis
tubercles bearing long setae.
Other species referable to this genus
0. S., of eastern North America.
are L. inornata, L. contempta, L. nigripleura, and L. noveboracensis, of the

Nearcticfauna;

L.

lucorum (Meig.), ofEurope; and L. frugi Bergr.,

L. claduroneura Speis., L. riatalensis Alex., L. spectabilis Alex., and several

other species, of the Ethiopian fauna.

Hart (1898 [1895]: 202-204) describes the habits of P. luteipennis insome detail. Larvae about half grown were found on March 17, 1895,
Hart ascertained
these attaining their growth and pupating on April 13.
by dissection that the species is a herbivore, feeding on fragments ofdead vegetation, numerous diatoms of many species, and minute algae.
As in many species of the Limnophilaria and a few other groups of
crane-flies, the spiracular lobes are fringed with long, delicate hairs which
spread out on the surface film of the water into broad fans. The larva
is unable to release itself from the hold of the water except by looping
the head and the anterior end of the body around the posterior end, anddrawing the latter thru this loop, repeating
have gone below the level of the film.
6

this action until the hairs.

Charles Paul Alexander

850

It is probable that other groups of species of the old genus Limnophila
will be found to deviate from the characters of tliis group as now restricted.

Species such as L. hyalipennis (Zett.), L. nemoralis (Meig.), and others in

Europe, and L. hrevifurca, L. emmelina,L.

mundoides,

L.

toxoneura,L. ultima, and similar anomalous forms in America, should be investigatedcritically when opportunity offers.

Pseudolunnophila luteipennis (0.
1859

Limnophila luteipennis O. S.

Pseudolimnophila luteipennis

S.)
Proc. Acad. Nat. Sci. Phila., p. 23G.

is

one of the most abundant species of

the tribe, and is very widely distributed thruout eastern North America.

The adult flies are common in swamps and along the marshy edges of
streams and ponds. The immature stages are characteristic swamp
inhabitants.
They have been described in detail by Hart (1898 [1895]:
Malloch (1915-17 b: 222-223) adds several supplementary notes
and figures of Hart's material.
202-204).

Larva.

— Length, 15-18 mm.
Diameter, 1.5-1.7 mm.

Coloration light brownish yellow, slightly darker toward posterior end of body.

Body covered with a delicate appressed brownish pubescence and abundant stiff bristles
and erect hairs on each segment, arranged in about five transverse rows; these bristles more
numerous, coarser, and blacker, on posterior segments, forming a large tuft on sides ofpenultimate segment of body. Spiracular disk (Plate XXXIX, 176) surrounded by four
lobes, the posterior pair very long and slender, finger-like, the lateral pair much shorter but
slender; the iimer face of these lobes with brownish black or dark brown transverse lines which
cover almost the entire face; an arcuate line extending from ends of ventral marks across disk
between spiracles; two small triangular marks located between ventral lobes; lobes fringed
with long, delicate hairs, those at ends very elongate, those toward base of lobes shortened

but continuous around disk.

Anal gills four, very long and slender, exceeding ventral lobesof spiracular disk.

Head capsule short and stout, exhibiting a very generalized condition.

Prefrons very

and but slightly exceeding lateral plates. Labrum
broadly transverse, with a large tuft of hairs on either side. Mentum (Plate XXXIX, 172)

broad, only slightly narrowed behindcompletely divided, each half with seven (or rarely eight) teeth along anterior margin, of

which the third (or foiu"th) from the inside is the largest; a flattened lobe just laterad of eachhalf of mentum; mental plates continuous behind with strong lateral plates of capsule; ina normal position the two halves of mentum overlying each other to a greater or less degree.
Hypopharynx (Plate XXXIX, 173) with anterior margin broadly rounded, with numerous(frombluntly rounded teeth. Antenna slender, with two long apical
Mandible (Plate XXXIX, 175) short and broad, of the generalized limnobiine

ten to fifteen)

papillae.

The Crane-Flies of New York— Part851

IItype; tips blunt, ending in two large approximated teeth, with a row of smaller lateral teeth

extending basad of each; ventral cutting edge with about four such lateral teeth, dorsal edge

with one or two much larger teeth; two strong setae near base of scrobe, and two others neara large brush of hairs at prosthecal region. IMaxilla not projecting, with

tip of mandible;

dense brushes of short, stiff hairs and with two short sensory tubercles which are tipped withvery elongate setae; palpi stout and chitinized at base, the apex narrowed and pale.

Pupa.

— Length, 10-13 mm.
Width, d.-s., about 1..5 mm.

Thorax reddish brown to black in color, depending on age of specimen; abdomen dirty
whitish with narrow transverse brownish lines, both above and below; pronotal breathinghorns dark brown or blackish.

Form subcylindrical, abdomen slightly depressed.

Cephalic crest of two slender tubercles

behind, tipped with long sctas; another pair of setae between antennal bases.

breathing horns elongate, transversely ringed, the tips widely and deeply

Prothorax with a prominent median carina.

divergent flaps.

breathing horn, each with about four stout setae.
tubercle bearing thrc-e setae.

Wings reaching end

Pronotal

split intotwo

A tubercle in front of each

Mesonotum above wing axil with aof second

abdominal segment.

Legs

reaching end of third abdominal segment, the tarsi ending about on a level or those of fore
legs a little longer than those of other two pairs.

Abdominal segments divided into two annuli, the basal one still further subdivided into
annulets; on tergum three annulets, each bearing a transverse row of setiferous tubercles, thesetae very longand conspicuous,insome cases the tubercles multisetose,with two widely separated, slender, elongate tubercles.

third annulet

Posterior annulus with a basal

and a terminal transverse row of setiferous tubercles. On sternum, four transverse rows of
setiferous tubercles on basal ring and two on posterior ring. On pleura, protuberant spiracles.
Lateral angles of segment 8 jutting out into stout lobes which are densely studded with
setiferous tubercles, on ventral face continued toward median line as a nearly straight row
Male cauda
of about eight setiferous tubercles, on dorsum a finger-like lobe on either sidet(Plate XL, 17.S) with sternal valves shorter than dorsal lobes, blunt at tips; tergal valves
Female cauda, (Plate XL, 179 and LSO) with tergal valvesacutely pointed, directed dorsad.
considerably longer than the more slender sternal valves, broad medially, tapering to thebroad, blunt tips which terminate in blackened points; two setae on outer margin.

———

Larch Meadows, Ithaca, New York, April 20, 1917.
Nepionoiype.
Female pupa, with type. No. 19-1917.
Neanotype.
Paralypes.
Larvae with type.
Pupae of both sexes from Bool's hillside, Ithaca, New
York, June 6, 1917. A female pupa, Orono, Maine, taken as a larva June 13, 1913,
emerged Jime 22. Female pupa, July 15, 1913, No. 75-1913.

Pseudolifunophila inornata (O. S.)
1869

Limnophila inornata 0. S.

Mon. Dipt. N. Amer., part 4, p. 219, 220.

Pseudolimnophila inornata is not so common as P. luteipennis but isfound in similar situations. The immature stages are spent in rich organic
mud. The only specimens that the writer has reared were taken inCharles Paul Alexander

852

Larch Meadows, near Ithaca, New York, on May 15, 1917, where they
were associated with larvae of Rhamphidia mainensis, PseiuioUmnophila
luteipennis, Tipula dejecta, and other swamp-inhabiting species.

— Very similar to that of P. luteipennis, but bodyLarva.-

isdarker and is conspicuously

blotched with whitish, especially on posterior parts of ventral segments.

Mouth parts

Antenna (Plate XXXIX, 174) with two very long andslender terminal papillae, one blunt at tip, the other much longer and tapering gradually
nearly the same in the two species.

to tip.

— Very similar to that of P. luteipennis, but somewhat smaller and darker-colored.

Pupa.'

Pronotal breathing horns (Plate XL, 177) darker brown, the divergent terminal flaps proportionately a little«more elongate, equal to almost one-quarter length of entire horn.

Dorsallobes at base of ovipositor more attenuated.

——

Nepionotype.
Larch Meadows, Ithaca, New York, May 15, 1917. No. 53-1917.
Neanotype.
Larch Meadows, Ithaca, New York, May 25, 1917.
No. 53-1917,

castpupal skin.

Subtribe Dactylolabaria

The division Dactylolabaria is proposed for the genus Dactylolabis,a small group of curious crane-fhes which are still not well understood.

The adult flies bear a striking resemblance to species of the genus Limbut the larvae and the pupae show characters that are not foundmore specialized divisions of the Hexatomini. The type of the

nopliila,in thegenus is Limnophila montana O. S. of the eastern United States.
Nearctic species included are L. damula O.

S.

Other

(western United States),

L. rhicnoptilaides Alex, (northwestern Canada), L. cuhitalis O. S. (eastern

United States), L. nitidithorax Alex, (western United States), and L.
hortensia Alex, (western Canada).

i—

The immature stages of two European species
D. wodzickii (Now.)
and D. denticulata Bergr.
have been made known by Nowicki (18G7:
340-343) and by Mik (1894), respectively. Their descriptions of thedetails of the larval head and the lateral spiracles of the abdomen of thepupa are incomplete, however. The larvae of none of the American
species have as yet been made known, but the pupae of D. cuhitalis 0. S.
were found by Needham and are described hereinafter.

—

Genus Dactylolabis Osten Sacken (Or. finger
1859

Dactylolabis O. S.

forceps)

Proc. Acad. Nat. Sci. Phila., p. 240.

—

Larva.
Body very depressed, the ventral surface, especially, being greatly flattened.
Head capsule compact; mentum not completely divided, its anterior margin with seven
teeth.

Mandible with but few lateral teeth.

The Crane-Flies of New York— Part II

853—

Pupa.'
Somewhat similar to pupae of Limnophilaria. Pronotal breathing horns shortand cylindrical. In some species, at least, second abdominal segment with a very large andprominent lateral spiracle.

Dactylolabis is a small genus including about thirteen known species
which are about -equally well distributed in Europe and North America.
Dactylolabis denticulata (Bergr.)
Limnophila denticulata Bergr.

1891

Mittheil. Natiu-f. Ges. Bern, p. 132.

The present knowledge of the life history of Dactylolabis denticulata^
due to the work of Mik (1894), v/hose paper is cited by other workers,
among them Griinberg (1910:54-55).

isLarvae and pupae were found on the dark brownish gray chalk cliffs
In certain places on the cliffs werebroad or narrow bands of black. The larvae and the pupae were foundalong these dark bands and showed a decided resemblance to their surroundings.
The surface of the rock was wet with dripping water, which
supported a flora of lowly plant organisms on which the larvae presumably
The adherence of the pupa to the last larval skin is of interest sincefed.
it presumably aids in the emergence of the adult.
A similar condition
is found in Cylindrotoma.
near Steiermark on August 3, 1891.

Larva.

— Length, 7 mm.
Width, 2 mm.

Form very depressed(Plate XLI, 183), the ventral surface flattened, almost leechlike.
Both dorsal and ventral surface provided with hairs, with longer and more numerous hairs
on sides of bodJ^ Body covered with small, black, structureless particles of earth or excrement, more numerous near margins of dorsal surface, these particles lacking a definite arrangement and being very firmly attached to the body hairs; when particles are removed, larva
is of a greenish gray color, subhyaline.
Abdominal segments divided into two rings bearing
long hairs which are more conspicuous on sides of body.
Head capsule (Plate XLI, 182)

small but compact, completely retractile within prothoracic segment.
large

Labrum relatively

and semitransparent, the margins with long hairs; on dorsum on either side a longdehcate seta.

Mentum chitinized; anterior margin with seven teeth, behind median tooth

Antenna two-segmented; basal segment cylindrical; second segment
Mandible three- toothed; a tuft of hairs at prosthecal region.
Spiracular disk surrounded by two fleshy welts which are deeply incised medially to appear

a brush ofhairs.

button-like, with apex pointed.

as four blunt lobes.

Spiracles diflficult to distinguish in a position of rest, elliptical, marginedwith blackish brown.

Pupa.

— Length, 8 mm.
Width, 1.75 mm.

Charles Paul Alexander

854
Coloration blackish;

Pupa(Platerock surface.

abdomen dark greenish gray.

XLI, 181) attached to old larval skin, which in turn remains attached to(Just before pupating, the larva sticks itself to the surface by means of its

Pronotal breathing horns cylindrical, a little swollen at ends. Sheaths of appendLeg sheaths attaining end*of fourth abdominal segment.

saliva.)ages dull and black in color.

Dactylolabis wodzickii (Now.)*

1867

Rhicnoptila wodzickii Now.

Verh. Zool.-Bot. Ges. Wien, vol. 17, p. 337-354.

The species Dactylolabis wodzickii was first found in the high Alpine
region of the Hungarian Tatras, at an altitude of from 6000 to 8000 feet,ina region frequented by the wall creeper (Tichodroma).

Here theadults and the immature stages were found together on the granite cliffs
where water dripped continuously, supporting a considerable algal flora.

Associated with the species were other flies, Liancahis virens (Scop.) andTricyphona schineri (Kol.).
The

Clinocera fontinalis Hal., as well as

degenerate wings of the insects prevent their flying, and they proljably
live and die close to the place where the eggs are deposited.

One fly wasTheobserved laying her eggs in the masses of algae in this situation.
larvae live in these masses and pupate in them.
Larva.

— I;ength, 12.5 mm.
Width, 2.4 mm.

Dorsum of body dark greenish brown, with dark longitudinal browTi stripes producing a
curious pattern of straightand convergent lines; ventral surface almost white, unmarked.

Body depressed, flattened, abdominal segments divided into a narrow basal and a muchbroader posterior annulus. Head capsule (Plate XLI, 185) compact, massive. Mandible
showing but a single inner tooth.

Posterior abdominal annuli with two long setae on lateral

Spiracular disk (Plate XLI, 186) surrounded by four lobes, t-he ventral pair longer

margins.

than the others; these lobes margined with darker chitinized lines and provided with fringes
Spiracles not described, and undoubtedly overlooked by theof moderately long hairs.
describer.

Pupa.

— Length, 12.5-13.5 mm.
Width, 2.6 mm.

Color brown.

Pronotal breathing horns short, cylindrical.

Wing sheaths ending opposite

Leg sheaths ending beyond middle of third segment.
Abdomen armed with rows of spines (Plate XLI, 184).
middle of second abdominal segment.

Dactylolabis cubitalis (0. S.)
1869

Limnophila cubitalis 0. S.

Mon. Dipt. N. Amer., part 4, p. 229.

Dactylolabis cubitalis is a local species, often occurring in great numbers.

In the gorge of Fall Creek near the Cornell University campus this species

The Crane-Flies of New York— PartIIcan be found in myriads by sweeping the rank herbage in May.

855
Tlie

writer has never located the immature stages, but the species was rearedby Dr. J. G. Needham at Ithaca, New York, on May 20,
is

1898.
There
no record as to where these larvae occurred, l)ut they are presumably

mud-inhaliiting forms, or possibly they frequent hal:»itats similar to thosedescril)ed for thetwo preceding species.

Pupa. — Length of cast skin, about 12 mm.
Pronotal breathinghoms (Plate XLI, 187) short, cylindrical, the tips scarcely enlarged.

IMesonotum behind with a prominent rounded tubercle on either side of median line, whichisset with

two spines, a larger outer spine and a small inner spine. Wing sheaths ending
Leg sheaths ending opposite apex of third

opposite apex of second abdominal segment.

abdominal segment, the tarsal sheaths terminating on acommon level.

Abdominal segments divided into two rings. Tergites with posterior ring having atransverse row of elongate tubercles, there being about eight on segment 2, about six onintermediate segments (3 to 5), and about four on segments 6 and 7; these tubercles fleshy,
but sparsely armed with chitinized spines. Basal ring unarmed. Pleura with a strong
tubercle on each ring; a blunt but prominent spiracle at extreme base of posterior ring; thisprominent on segment 2 (Plate XLI, 188), much exceeding lateral tubercles
and being about two-fifths length of pronotal breathing horns. Sternites with about six
tubercles on posterior ring.
Male cauda (Plate XLI, 189) with tergal lobes slender, runningspiracle veryout into acute chitinized points, which are directed caudad, slightly divergent; dorsum ofsegment 8 with posterior lobes powerful, about equal in size to, or a httle larger than, tergal
lobes of cauda just described; anterior lobes small, directed caudad and laterad;

two large
Female cauda with tergal valves moderately
elongate, stout, broad at base, narrowed toward tip, where they run out into long,. chitinized
points directed caudad and slightly dorsad.
lateral lobes

and two smaller ventral lobes.

——

Neanotype.
Cast pupal skin, Ithaca, New York, May 20, 1898.
Paratype.
Pupae, cast skins, with type.

Dactylolabis montana (0. S.)
1859

Limnophila montana 0. S.

The species

Dactylolabis

Proc. Acad. Nat. Sci. Phila., p. 240."-montanathruout the northern United States.

iscommon and widely distributed

It isa characteristic inhabitant ofrocky cliffs, where it rests in crevices on the almost vertical walls. The
immature stages have long remained unknown, but during the springof 1920 W. L. McAtee found a pupal skin and the teneral adult near

Washington, D. C. Mr. McAtee writes that the pupa was found in themoss that covers the almost vertical north faca of a cliff on Plummers
Island; this moss is mostly shaded, and grows on a thin layer of black soil
which, at that time of the year at least,issaturated with water.

The

Charles Paul Alexander856

pupal skin was not in condition for study, but now that the larval habitat

known it is hoped that more material
become available.

isof theimmature stages

will

Subtribe Limnophilaria

The division Limnophilaria comprises a large and heterogeneous groupof species which in their larval and pupal characters grade rather insensibly

into the next subtribe, the Hexatomaria, altho the adults of thetwosubtribes are very distinct.

The larvae of the various species in the group Limnophilae are rather
The head capsule is long and narrow, with thelateral plates very slender.
The labrum is ample, with, several sensory
papillae and setae.
The antennae (which are two-segmented in thesimilar to one another.

subgenera Phylidorea and Lasiomastix, and probably also in Dicranophragma and others) bear at their tips, in addition to the usual elongate
setae,

an oval or elongate-oval papilla whichisdelicately sculptured.

The mental region is feebly chitinized, and consists of two short longitudinalbars articulated at their cephalic ends with a transverse bar which functions
as the mental plate; this bar, or plate, is usually delicately grooved withparallel striae, atype of articulation that allows for great distention of
The mandibles are powerful chitinized hooks

the gular region in feeding.
])earing

two or more acute or flattened teeth at about midlength.

Theouter lo])cs of the maxillae project far cephalad as pale flattened blades.

The spiracular disk, in the primitive condition, is squarely truncatedand is surrounded by five subequal lobes which are fringed with rather
short hairs.
The dorso-median lobe is lost in most species, but the twopairs of lobes are more or less preserved in the other species known to thewriter, altho the lateral pair is sometimes reduced to a mere fringe or

The terminal fringes of the ventral lobes are often greatly
The anal gills are four in number, retractile,and rarely conspicuous. As is usual in many mud- or sand-inhabiting
tuft of hairs.

elongated, and fanlike.

species of Tipulidae,

the larvae are capable of greatly distending the

Tliis segment is often provided withnumerous transverse rows of fine points, and its inflation assists in thesubterminal abdominal segment.
larva's progression thru the soil.

The larvat; of the group are carnivorous, and almost without exception
are exceedingly agile and snakelike in their motions.

The Crane-Flies of New York— Part II857

Tho pupae of the group Limnophilae have the pronotal breathing horns
The abdomen is often
greatly depressed, with the lateral margins flattened and carinate andusually small, or at least not greatly elongated.

In the species of the subgenus Phylidorea thesegments of the abdomen are armed with rows of long, slender spines.
The group Ulomorphae seems to be closely related to the other members
the segments incised.

of the division, ])ut the immature stages present some curious conditions

which it is difficult to correlate with the same structures in the group
Limnophilae. The writer has removed a group of species from the oldgenus Limnophila, and has placed these in the genus Pilaria Sintenis.
In the larvae of the group Ulomorphae, the epipharynx and the maxillae
are densely hairy.
The maxillary lobes are slender. The mandibles
are hinged at about midlcngth, the basal segment being concave on its

when in a position of rest.
The mandibular blade has one, or sometimes two, elongate acute teeth
at the base.
The mental region is not chitinized. The spiracular disk
is surrounded by four lobes, of which the lateral pair are often very
inner face to receive the mandibular blade

reduced; the inner faces of the lateral lobes are capable of close approxi-

mation, so that when they are closely applied the spiracles are contiguous.
The pupae have the pronotal breathing horns very elongate, with theirtips split into two flattened divergent lobes, as in Pseudolimnophila anda few tipuline forms.

The following keys separate the genera and the subgenera of the subtribe
Limnophilaria:
LarvaeMental region not chitinized; maxillae and epipharynx fringed with conspicuous, long,
golden-yellow hairs; mandibles hinged; head capsule with dorsal plate spatulate at
its tip.
(Group Ulomorphae, p. 869J
2
Mental region a narrow, transverse, chitinized bar, finely striate; maxillae andepipharynx without conspicuous hairs; mandibles not hinged; head capsule not as
described above.
(Group Limnophilae, p. 858)
3
2. Length under 12 mm. basal tooth of mandibular blade nearly half the length of blade.
Ulomorpha O. S. (p. 869)
Length over 14 mm. basal tooth of mandibular blade about one-third or less the length
1.

;

;

of blade

3.

4.

Pilaria Smt. (p. 872)
Spiracular disk almost squarely truncated, surrounded by five subequal lobes producing
an eriopterine appearance
Dicranophragma O. S. (p. 861)
Spiracular disk obliquely truncated, surrounded by two pairs of lobes of which the lateralpair are the shorter
4
Mandibles with two or three acute teeth at about midlength; epipharj-nx with a circulararea bearing two biarticulate papillae
Phylidorea Bigot (p. 866)
Mandibles with three or four flattened and truncated teeth along blade; epipharynx
without papillae as above
Lasiomaslix O. S. (p. 863)

Charles Paul Alexander

858

Pupae
Pronotal breathing horns very long, cylindrical, the tips split into divergent flaps.
(droup Ulomorphae, p. 800)2
Pronotal breathing horns not elongate-cylindrical.
(Group Limnophilae, p. 858)
3
Abdominal segments with tubercles or spines only near posterior margins of segments.
Ulomorpha O. S. (p. 869)
Abdominal segments with three or four pairs of blunt, naked tubercles.
Pilaria Sint. (p. 872)
Abdominal segments depressed, lateral margins flattened, carinate, segments deeply

1.

2.

3.

4

incisedAbdominal segments not depressed nor incised; pronotal breathing horns bicolored.
Dicranophragma O. S. (p. 861)_
4. Abdominal segments with rows of acute slender spines; pronotal breathing horns very
short, semicircular in outline, bluntly rounded at their tips. Phylidoren Bigot (p. 866)
Abdominal segments without such slender spines; pronotal breathing horns short, broad,.

Lasiomastix O. S. (p. 863)

slightly compressed, tips a little pointedGroup Limnophilae

Genus Limnophila Macquart (Gr. sivamp + friend)
1834
1854
1861

Limnophila Macq.

Suit,

Phylidorea Bigot.

Ann. Soc. Ent. France, p. 4.56.

Limnomija Rond.

Dipt. Ital. Prodr., vol. 4, p. 11.

Larva.-— Form slender.

a Buff., vol. 1, Hist. Nat. Ins., Dipt., p. 95.

Spiracular disk surrounded

by

five,

or

more commonly

four,lobes, the ventral pair the longer and fringed with long, delicate hairs.

Anal gills four, notMandible notconspicuous. Head capsule flattened, very dissected, with plates narrow.
Maxillahinged, curved, chitinized, on cutting edge with a few acute or flattened teeth.
with outer lobe produced into a flattened projecting blade.
its

tip

with a short or elongate-oval papilla whichtransverse chitinized crossbar whichisisAntenna two-segmented, atdelicately sculptured.

Mentum afinely grooved.

— Cephalic crest setiferous. Pronotal breathing horns usually small, flattened,

Pupa.-

Mesonotum convex or flattened above.
armed with tubercles or spines.
tips not split.

Abdominal segments in some cases

The genus Limnophila comprises a very extensive group of crane-flies(more than one hundred and fifty species), which are found in most partsof the world but are apparently more numerous in the temperate regions.
The adult flies seem to be closely related, but the immature stages are sovaried in structure as to make it appear that the group must be a heterogeneous one and the similarity of the adults a result of convergent evolution.
The adult flies may be found resting on rank vegetation. Several of
the species {Limnophila ultima, for example) swarm in small groups offrom fifty to sixty individuals, copulation taking place in the air, as is
discussed more fully on page 711.

The Crane-Flies of New York— Part II

859

The immature stages of several species have been made known, butThestill very considerable gaps in the present knowledge.
Thelarvae are among the most carnivorous of all crane-fly larvae.
there arefeeble chitinization of them: ntal region allows for great distention, andoftentimes the prey of thei^e larvae consists of forms tliat are nearly aslarge as the captors themselves.

The data regarding extra-American

species

may be summarized asfollows:

Limnophila hryohia Mik (1881:205-206) was reared from moss, taken in

New Zealand, in the summer (December to
The moss was dried and then placed in a temIn March of 1879 an adult fly
perature of from 12 to 15 Reaumur.
appeared. This long interval of more than four years was supposed by
the Auckland Islands, nearFebruar}^) of 1874-75.

Mik to have been passed In^ the insect as a larva, the dry moss furnishing
th(5

food.

Possibly the eggs were carried over for a long period before

hatching.

Limnophila sinistra Hutton, of New Zealand, has been discussed in somedetail by Hudson (1920:33-34), v/ho states that this species is ,common inmost dens3 forests thruout the country. The larva inhabits fallen treetrunks in an advanced state of decay, forming burrows between the soft,
decayed part and the harder part of the wood. It is about 25 millimetersin length, cylindrical, tapering toward the head, which is very small, andfui'nis'ied with two minute jaws and a pair of very short antennae.
There
The extremity of the last segment isare eleven visible body segments.
truncate and deeply excavated, the concavity being protected by five converging spines, which can be spread out or drawn inward at the will of
the insect. Pedal warts occur on the undersides of all the segments except
the three irnxUiediately following the head and the terminal segment. The
pupa is about 12 millimeters in length, and rather stout. The breathinghorns are about one-third the length of the wing sheaths, and are modThere are two dorsal rows of hooks
erately stout and strongly recurved.
on each exposed abdominal segment and one ventral row near the termiThe valves of the ovipositor are strongly recurved andnal extremity.
The head and the thorax are dark brownish black, highlyver}^ stout.
polished; the abdomen is grayish ocherous, darker in the middle; the ovipositor

and the terminal segmentsofthe posterior tarsi arc reddish.

The pupa rests in a burrow made by the larva near the surface of the log.

Charles Paul Alexander

860

Edwards has decided that this insect and a few related species should be
The affinities of the fly, to judge from the details
supplied by Hudson, arc with the Epiphragmaria rather than with the
Limnophilaria, but until more details are forthcoming it cannot be defiplaced in a new genus.

nitely referred to that division.

Limnophila {Phylidorea) ferruginea (Meig.), of Europe, is closely allied
to L. adusta (page 867), and is of especial interest as being the type ofthe subgenus Phylidorea. It was reared by De Meijere (1916:204-206)
from larvae found living between saturated decaying leaves along thebanks of watercourses. Further mention of the species is made under
the discussion of the subgenus Phylidorea (page 866).
Limnophila hyalipennis (Zett.), of Europe, was found by Bcling (1886:
198-199) in piny woods, where the larvae live in old ant hills or in pilesof earth thrown up by wagon wheels.
The apparent lack of pupal breathinghorns in this species is discussed on page 755. The European species
L. lineola (Meig.) (Beling, 1879:54 and 1886:199-200), L. nemoralis(Meig.) (Behng, 1886:200-201), and L. ochracea (Meig.) (Behng, 1886
202) are all found in damp earth along the margins of woodkmd streams
or in damp spots in the woods. Larvae of L. dispar (Meig.) were foundby Perris (1849:331) living in the hard, withered stalks of an umbellifer,
Angelica sylvestris Linn., where they hollowed out long passages in thepith.
(This reference is cited by Mik, 1881:204, and by Osten Sacken,
1869:201-202.) Limnophila pallida Bel. was reared by Beling (1873 a:557) from larvae living in a decaying ash trunk.
The reference to Limnophila platijptera Macq. given by Heeger (1854)
is considered })y Osten Sacken (1869:4, 202) to refer to a mycetophilid,I

probably Bolitophila.

The American
following pages.

speciesthatare

now knownareconsidered inthe

In addition, Limnophila unica 0. S. has been bred fromlarvae found in decaying wood.
It isvery probable that some of the species listed above, as well as

many of the limnophiline forms that are still unknown as regards theirimmature stages,

will

be found to belong to some one or other of theremaining divisions of the tribe, rather than to the Limnophilaria as here
restricted.

This group requires more careful study than does any otherdivision of crane-flies.

IThe Crane-Flies of New York— Part II

861(Subgenus Dicranophragma Osten Sacken)
1859

Dicranophragma 0. S.

Proc. Acad. Nat. Sci. Phila., p. 240.

Larva. — Form short and stout.

Spiracular disk surrounded by five short lobes, theventral pair the longest, the dorsal lobe very blunt; disk surrounded by a fringe of delicate

Head capsule of the hexatomine type. Mandible
hook with a large acute tooth just beyond midlength.
Mentum chitinized, consisting of a narrow transverse bar

hairs which are longest at tips of lobes.

powerful, prolonged into a strong
Ma-xilla very longand slender.

which is ribbed with fine parallel grooves.

Pupa.

— Cephalic crest prominent, each half with three set^. Pronotal breathing horns

or slightly narrowed to a blunt tip.
Mesonotum convex, unarmed.
Wing sheaths short, broad, ending before tip of second abdominal segment. Leg sheaths
short,

cylindrical,

moderate in length, ending just beyond base of fourth abdominal segment, hind tarsi a little
longer than the others.

Abdominal segments divided into two rings; posterior ring witha subterminal transverse armature, on dorsum consisting of numerous setiferous tubercles,on pleura consisting of two or three groups of setiferous tubercles; basal ring with about
eight slender tubercles on dorsal surface.
Spiracles present, but evidently nonfunctional.

Dorsum of segment 8 with four slender tubercles.

Dicranophragma is a well-defined group of the genus Limnophila, with
about six known species whose center of distribution seems to be in the
Oriental region.

The tj^pe of the subgenus, Limnophila (Dicranophragma)
is the only described New World

fuscovaria, discussed in detail below,

The immatureform.

stagesare spent inrich

organic

mud in

cool,

Nat.

Phila.,

shaded woods.

Limnophila {Dicranophragma) fuscovaria O. S.
18-59

Limnophila {Dicranophragma) fuscovaria O. S.

Proc.

Acad.

Sci.

p. 240.

The adult flies of Limnophila fuscovaria may be easily distinguished bytheir broad, heavily spotted wings and the strong supernumerar}^ crossvein

They may be swept from rank vegetation in cool, shaded
The larva has the body stouter and less elongate than is usualin this group of species, but it possesses the same snakelike movements
as its near relatives.
The pupal duration is not longer than eight daysin cell R2.

woods.

(June 8 to 16, in 1917).
Larva.

— Length, 6.8-7.2 mm.
Diameter, 0.7-O.S

mm

Coloration light golden-yellow; maxillary lobes yellow; spiracular disk lined with dark brown.

Body rather short and stout, not of the exceedingly elongate type of the Eriopterini, gradually narrowed to anteriorend (Plate XLII, 190) prothoracic segment truncated anteriorly;;

Charles Paul Alexander

862
when headisretracted,

tips

of

maxillae projecting from prothoracic

orifice.

Seventhabdominal segment with a transverse fringe of stiff hairs on the sides, these directed lateradandslightlycaudad.

Body

noticeably constricted just before last segment.

Spiracular

XLII, 195) surrounded by five lobes, and in superficial appearance decidedly
eriopterine rather than hexatomine; ventral lobes the longest but still much shorter thanis usual in this tribe, slightly divergent, inner face with a double, V-shaped, dark brown
disk(Platemark, inner arm of V narrow, outer arm more expanded, especially at proximal ends, one
large inner and two smaller outer sensory bristles at apex of V; lateral lobes narrow, with aV-shaped mark similar to that of ventral lobes but narrower and darker; dorsal lobe broad,the inner face indistinctly lined with parallel, dusky, longitudinal stripes;

allfive lobes

surrounded by a fringe of long, delicate hairs lying just outside brown marks, these hairs
Spiracles circular, rather widely sepalongest at tips of lobes and continuous around disk.
rated; ring broad, light yellow, narrowly margined outwardly with black.

Anal gills four,

short and stout.

Head capsule (Plate XLII, 191) very dissected, consisting of six elongate slender plates,
Labrum (Plate XLII,

three on either side, articulating at a point laterad of mental plate.

192) large; anterior margin indistinctly trilobed, the median lobe hairy; on the ventral, or

epipharyngeal, surface, on either side, a slender cylindrical papilla, subhyaline, directedcephalad; sides of epipharynx with a tuft of long setae.

Mentum chitinized, consisting of

two lateral rods and an anterior transverse bar which is finely ribbed with parallel lines.
Immediately dorsad of mentum two roughened papillae (prementum) directed cephalad;esophagus retrorsely roughened. Antenna (Plate XLII, 193) stout, in caustic-potash preparations almost hyaline, tipped with an elongate cylindrical lobe distad of which is a small
Mandible
setiferous tubercle with two long, delicate setae exceeding antennae in length.
(Plate XLII, 194) powerful, very deep at base, produced into an acute, strongly curved tip;
beyond midlength a very large, acute tooth which is about one-third length of apicaljust

point, in its angle a small, hyaline, flattened, leaflike blade with a truncated apex; basadof large tooth

two or three gradually smaller teeth.

Maxilla with outer lobes stout, broadat base, gradually narrowed to tip, which is cut off by a constriction into a semi-oval structure; palpi

on inner dorsal face of maxillary lobe at about midlength, short, semi-globular,

with abundant sensory papillae; when head is completely retracted, the long, pale, sUghtly
divergent tips of maxillary iobes projecting fromorifice.

Pupa.-—-Length of cast skin, about 6 mm.

General coloration dark brown; pronotal breathing horns almost black, with apical fourthconspicuously light yellow; abdominal incisures pale.

Cephalic crest large and prominent, with three long setae on outer face.
at apex.

Labrum blunt

Labial lobes with a blunt point on either side, these directed proximad and caudad.

Maxillary palpi broad at base, suddenly narrowed to the long, cylindrical tip, which isdarkened. Antennae rather short. Pronotal breathing horns (Plate XLIII, 196) stout,
short, almost straight,

broad basally, tapering gradually to the blunt

tips.

Wing sheaths

very short and broad, ending before tip of second abdominal segment. Leg sheaths short,extending to just beyond base of fourth abdominal segment; all the tarsi ending about on alevel, orouter sheaths a little_the longer.

The Crane-Flies of New York— Part II

863

Abdominal segments (Plate XLIII, 198) each divided into two rings by a false constriction,
Basal ring on dorsum with a transverse
row of about eight slender, blunt tubercles before caudal margin; on pleura a similar but
larger tubercle opposite basal ring, this tipped with a long, delicate seta.
Sternum unarmed.
Posterior ring on dorsum with a subterminal row of blunt tubercles, simple or bifid, often
approximated or crowded, bearing one or two long setae; near base of tergum, opposite
spiracle, a setiferous tubercle; on pleura, opposite base of posterior ring and nearer dorsum
basal ring about two-thirds length of posterior ring.

than sternum, a vestigial spiracle with a double setiferous tubercle ventrad of it; near caudal
margin of pleura two or three compound multisetose tubercles (Plate XLIII, 197) or aggregations of simple tubercles, which are largest on seventh segment.
On sternum, two slender
tubercles on either side, near base of posterior ring.

Female cauda (Plate XLIII, 201)

elongate, with the tergal valves a little longer than the sternal valves; each of tergal valves

with a small tubercle at base.
lobes blunt, the dorsal lobes a

Male cauda (Plate XLIII, 199 and 200) with the ventral
longer and ending in an acute point; a tiny tubercle

littleon either side at base of dorsal lobes.

Dorsum of segment 8 with a trapezoid of four con-spicuous lobes ending in truncated tubercles; anterior pair of lobes smaller and a little more

widely separated than posterior pair.

——

Nepionotype.
Ithaca, New York.
Neanotype.
Ithaca, New York, May 31, 1917.
No. 57-1917.
Paraiypes.— F\ipa.l skins, June 11, 1917 (No. 99-1917), June 16, 1917 (No. 1.32-1917).
Abundant larvae, with nepionotype.

(Subgenus Lasiomastix Osten Sacken)
1859
Larva.

Lasiomastix 0. S.

— BodyProc. Acad. Nat. Sci. Phila., p. 233.

slender, with pencils of stiff setae on lateral margins of thorax.

Spiracular

disk with four lobes, the ventral pair a little the longer and bearing a fringe of long, delicate

Anal gills short and stout. Head capsule ofMandible powerful, cutting edge with a single row of three or four
Maxilla very long and slender. Antenna elongate, at tip withflattened, bladelike teeth.
a small elongate-oval papilla whose surface is delicately sculptured. IMentum chitinized,a strong transverse bar which is ribbed with fine parallel grooves.
Pupa: CephaUc crest very small, inconspicuous, bilobed. Pronotal breathing horns
short, broad, somewhat compressed, a little pointed at apex.
Mesonotum unarmed.
Abdominal segments depressed, armed with numerous tubercles or small spines, these being
most abundant on posterior annuli of segments.
hairs,

these about twice length of lobes.

hexatomine type.

—The subgenus Lasiomastix includes but six known species, three ofare from eastern North America. Limnophila {Lasiomastix)

which

macroceraiscommon and widely distributed thruout the Eastern States.

and L. (L.) subtenuicornis (Alex.) are found only
Northeastern States and are more local in their distribution. The
immature stages of L. (L.) macrocera are spent in rich organic mud. They

L. (L.) tenuicornis 0. S.
in theare very similar to those of the subgenus Phylidorea.

Charles Paul Alexander

864

Limnophila (Lasiomastix) macrocera (Say)1823^

Limnobia macrocera Say.

Journ. Acad. Nat. Sci Phila., vol. 3, p. 20.

Limnophila macrocera is a common swamp-inhabiting crane-fly, thelarvae Hving in organic mud.
At Orono, Maine, the writer found larvaeof this species associated with larvae of Bittacomorpha clavipes, Pilaria
teyiuipes, P. recondita, Erioptera chlorophylla, and other crane-flies, asw(^ll as with leeches, snails, worms of many kinds, and other forms oflife.

The larva is similar to that of other related species.

When placedin water it is very active and has the habit of darting the anterior quarterof its body from one side to the other, suggesting the striking of a reptile.

The pupal duration is not more than eight days (June 24 to July 2, June
28 to July
Larva.

6).

— Length, 14.5-15 mm.
Diameter, 1.4 mm.

Coloration, pale yellowish white.

Body terete, narrowed toward both ends but more noticeably toward anterior end.
Integument covered witth a dense appressed pubescence. Chaetotaxy as follows: two stout
setae on both dorsal and ventral margins of prothoracic orifice; thorax with pencils of stiff
setae near anterior margins of segments, two large lateral tufts and a smaller ventral pair;abdominal segments with a pencilof similar

bristleson ventro-lateral

margins, one on

anterior half, the other midlength, of each ring; a seta at posterior lateral angles of sternites;

four setae in a transverse row near posterior margin of tergites, the lateral pair the larger;
lateral margins of

cauda near base of lateral lobes and above gills with pencils of blackishsetae; basal part of segments on both sternum and tergum with transverse parallel rows offine scabrous points, this area very

narrow on second abdominal segment, consisting only
becoming broader and the rows more numerous,of three or four rows, the areas gradually

there being on sixth
ba'^al tliird ofand seventh segments about 28 to 30 rows which occupy nearly thesegments.

'

Spiracular disk (Plate XLIV, 208) surrounded by four lobes; ventral pair the longest, lateralpair a little shorter; ventral lobes near tips with a brush of delicate, exceedingly long hairs

which curl into loops at tips, the longest of these hairs about twice length of lobes bearing
them; this fringe of hairs continuous around disk, longest at ends of lobes, gradually shortened

toward their bases but not interrupted; lateral lobes with terminal hairs a little shorter but
still

longer than lobes themselves; inner faces of lobes delicately margined with dark brown,

those of ventral lobes having the outer lateral margins the broadest.
located at base of lateral lobes.

Spiracles rather small,

Anal gills four, very stout and plump.

Head capsule with a framework of long, slender plates, as is usual in this group.

Labrum(Plate XLIV, 205) broadly transverse; anterior margin irregularly rounded; anterior medianarea truncated, with the lateral angles slightly projecting, cut off squarely, each with a small,and caudad of each, two sensory papillae, the innermost
more than twice length of short, oval, outer one. Epipharyngeal

hyaline, sensory papilla; lateradelongate-cylindrical,

The Crane-Flies of New Yorkregion roughened by tiny groups of parallel ridges.

— Part II

865

Mental region consisting of a heavily

chitinized transverse bar which is delicately grooved, enlarged at ends, and articulating witha small ventral bar.

Antenna (Plate XLIV, 206) long and slender; basal segment short andbroad; second segment elongate-cylindrical, bearing at its tip a small, hyaline, sensory papillawhich appears delicately crosshatched byfine

impressedlines,

this papilla elongate-oval

with the apex broadly rounded; a long hair near base of this papilla and about twice its length.

Mandible (Plate XLIV, 207) long and powerful, the tip produced into a slender point, thecutting edge at about midlength with a single row of three or four flattened, bladehke teeth

which are truncated at their tips, these teeth varying considerably in their shape and relative
proportions.

Maxilla with outer lobe elongate, the base strongly chitinized, this chitinizedpart continuing up margin of lobe almost to

midlength, the small palpus and a long seta.

the apical part nearly hyaline; at about
Esophagus enlarged, the walls thickened andtip,roughened.

(A slide of a larva that was supposed to belong to this species differs considerably from

The antenna is much shorter, with the apical tubercle muchand elongate-cylindrical; the mandible has the teeth fewer and more acute, quite
as shown by De Meijere for Limnophila [Phylidorea] ferruginea.
It is probable that thisthe material described above.

largerspecies likewise is a Phylidorea.)

Pupa.

— Length, 14-16 mm.
Width, d.-s., 1.8-2 mm.
Depth, d.-v., 1.9-2.2 mm.

Breathing horns pale whitish brown, alittledarker at base; remainder of body darkbrownish black, abdomen a little lighter-colored; incisures of segments, and pleural line, pale.
Cephalic crest very small, inconspicuous, bilobed, each principal lobe subdivided into

two smaller lobes terminating in a small seta; another seta on anterior face. Labrum
Labial lobes terminating in sharp points projecting
rather blunt and truncated at apex.
proximad. Maxillary palpi short and stout, tapering to blunt tips (Plate XLV, 210).
Antenna of male elongate, reaching to beyond midlength of wing sheath; antenna of female
short, ending at about one-fourth length of wing sheath.
Segments of antenna cylindrical,
unarmed.
Pronotal breathing horns (Plate XLV, 209) short, broad, somewhat compressed, a little
pointed at tip. Thorax broad and deep, the pronotum flattened, carinate medially.
Mesonotum high, convex, transversely wrinkled. Wing sheaths ending some distance before
Leg sheaths short, ending before tip of third abdominal
tip of second abdominal segment.
segment, the hind legs a little longer than the other legs, which end about on a common
level.

Abdomen depressed, lateral margin carinate.

Abdominal segments distinct, incised, eachsegment divided into two annuli, basal one about half length of posterior one. Basal ringon dorsum with a transverse single, or somewhat double, row of small subacute spines, which
Pleura unarmed. Sternum with an
are more numerous and larger on mid-dorsal area.

obHque flattened lobe or wing on each side, directed caudad and proximad; between theselobes a more or less broken transverse row of from six to twelve slender tubercles.
ring

on dorsum with numerous small scattered tubercles.

Posterior

Pleura with a small circularspiracle near base and close to dorsal margin; a slender seta near spiracle and ventrad of it;Charles Paul Alexander

866

two other setae caudad of spiracle; at caudal margin a soli tarj* seta close to ventral margin.
Sternum with scattered slender tubercles, on segment 3 these appearing as a longitudinal rowof about ten alongside the hind legs; between this lateral longitudinal row and the scattereddiscal tubercles, a bare space, at anterior end of which is a large, stout, setiferous tubercle;
a subterminal transverse row of about fifteen acute black spines; on terntiinal segments thelateral longitudinal rows interrupted, or, en the seventh segment, lacking, on seventh segment
the subterminal armature of both dorsum and sternum more powerful, especially the intermediate spines of sternum, which are very large; numerous setae scattered along rows.
Segments 3 to 8 on ventral surface of posterior ring with a circular disklike area, median(In some specimens, especially females,in position and at about two-thirds length of ring.
the discal tubercles on the posterior ring are much smaller and lie in longitudinal rows.)]\Iale Cauda (Plate

XLV, 211) with tergal valves slender, elongate, slightly divergent, taper-ing to the acute tips, which bear a small subterminal seta; sternal valves short, blunt, witha flattened oval lobe between valves of sternum; eighth segment on dorsum with a trapezoidof four very long, slender, pointed lobes, which are provided with delicate hairs, the posterior
pair of lobes longer, stouter, and lying closer together; just above anterior pair of lobes a
blunt setiferous tubercle; pleural region of segment 8 with two spines; a small seta cephaladof these spines; sternal region of segment 8 with

two widely separated setiferous tubercles.

Female cauda (Plate XLV, 212) with tergal valves elongate, narrowed at tips, and armed
with sharp, black spines; a few small setae before tips; sternal valves short, blunt, narrowed
at tips.

— Orono, Maine, July
— With type
July
— Larvae, with type

Nepionolype.
Neanofijpe.
Paratypes.
1913.

larva,

19, 1913.
17, 1913.

larva, June 13 to July 27, 1913.

Pupae, June 9 to July 22,(Subgenus Phylidorea Bigot)
1854

Phylidorea Bigot.

Ann. Soc. Ent. France, p. 456.

The immature stages of the type species of PhyHdorea, Limnophila
by De Meijere (1916:(Phylidorea) ferruginea (Meig.), were described

In Holland, larvae were found in April on the banks of a ditch,where they were living between decaying leaves. The larvae are cylinThe antennae
drical, and are from brownish yellow to brown in color.
204-206).

are described as three-segmented,

the apical papilla being considered

The labrum bears on the epipharyngeal surface a median
The outerprojection provided with two conspicuous biarticulate lobes.
The mandibles arelobes of the maxillae project as flattened blades.

as a segment.

powerful and strongly curved, and have a group of small lateral teeth
at about midlength.

The pupa is about 10 millimeters long and is blackish brown in color.
The pronotal breathing horns are almost semicircular and are yellowish

The Cjiane-Flies of New York— Part867

IIbrown in color. The abdomen is depressed, with sharp lateral margins.
The abdominal armature is described as consisting of small hairs insteadof slender spines as in the American species of this subgenus.
In the Nearctic fauna, the subgenus Phylidorea includes Limnophila
adusta, L. similis Alex., L. novae-angliae Alex., L. lutea Doane, L. terras-

novae Alex., L. costata Coq., L. fulvocostalis Coq., L. insularis Johns., andprobably other western species.

It

seems quite possible, hioreover, thatthe species allied to Limncphila lenta O. S. also belong here.

Limnophila (Phylidorea) adusta 0. S. (supposition)
Limnoyhila adusta 0. S.

1859

Proc. Acad. Nat. Sci. Phila., p. 235.

A larva of a crane-fly that is almost certainly a member of this genus
was taken at Ithaca, New York, on May 30, 1917, in company with a
Ulomorpha (No. 88-1917). An unknown pupa was taken in the sedge
It was not
association on the Bool hillside, Ithaca, on June 4, 1917.
reared and its identity can only be surmised, but there can be little doubtof its relationship with Phylidorea.
Larva.

— Length, 8.6-9 mm.
Diameter, 0.8 mm.

Coloration a deep saturated orange-yellow, the body with silky iridescent reflections.

Body apubescence.
of,littlenarrowed at both ends.

Integument with along,appressed,

goldenNumerous long setae and pencils of hairs on body, usually at about midlength

or on posterior half of, the segments; posterior margin of each segment elevated into

prominent transverse ridge ofhairs.

aSpiracular disk surrounded by four lobes, ventral

pair about twice length of lateral pair; ventral lobes with a rather narrow, dark brown line

on inner face; outer margin fringed with very long dark hairs which are somewhat paler
at their bases, this fringe of hairs longest at tips of lobes, where it reaches a length of about
eight orten times length of lobe; hairs shorter toward base of lobe; a stiff sensory br'stle

on outer face of lobe, rather far back from tip; lateral lobes similar, with the fringe of hairs
yellow and about twice length of lobes. Spiracles of medium size, directed toward each other.
Anal gills four, fleshy, pale in color, posterior pair longer than anterior pair.
Head capsule of hexatomine type, the dorsal plates narrowly interrupted on mid-dorsal
line behind clypeal-labral sclerite.
Labrum very large, anterior margin provided witheight or ten papillae

area whichisand setae.

slightly elevated

Epipharyngeal region having a

and bears two bisegmented

large,

median, circularcylindrical papillae.

IMentalregion as usual in the genus, consisting of three hinged bars forming three sides of a rectangle,

the anterior transverse bar delicately grooved.

Antenna two-segmented, second segment

narrower than first and bearing at its tip an elongate sculptured papilla and a long seta.

Mandible of hexatomine type, a powerful curved hook, at about midlength of which is anChaeles Paul Alexander

8G8

acute tooth; in axil of latter a small flattened blade which is slightly widened outwardly andhas the tip truncated.

Maxillary lobe broad at base, tapering to narrow tip.

— Length, 12 mm.

Pupa.

Depth, d.-v., 1.6 mm.
Similar to Limnophila macrocera in general shape and color, differing as follows:

Labrum broadly obtuse at tip.

Pronotal breathing horns (Plate XLIII, 203) very short,stout but flattened, constricted beyond midlength, and with a row of breathing ports aroundapex; pronotum and mesonotum carinate medially in front; wing pad showing venation rather

Mi deep, basal deflection of Cm beyond midlength of cell 1st Mi.

clearly, cell

Abdominal

segments with lateral margins very deeply incised, the carinate lateral margins very accentuArmature of abdomen consistated, appearing as thin, flattened wings (Plate XLIII, 204).
ing of abundant elongate, acicular spines, some of which are sinuously twisted; on basal ringthese spines appearing as a subterminal transverse row on both

dorsum and sternum; onposterior ring, besides the subterminal transverse row, spines are scattered over surface inmore or less distinct longitudinal rows; at lateral carina a group of about a dozen long spines
at caudal margin, as well as a powerful spine below level of spiracle on extreme margin ofcarina; ventrad and caudad of this, three long setae,

two close beneath spine and the third

Female cauda (Plate XLIII, 202) about as in Ldmnophila macrocera,
but st;rnal valves divergent at their tips; eighth segment with a trapezoid of dorsal lobes
about as in L. macrocera, but sternum with four stout spines bearing setae on their sides;
underneath spiracle.

a blunt median lobe near base of eighth sternite.

New York, on June 4,
was associated with a characteristic swamp-inhabiting crane-fly fauna(Described from a female pupa taken on Bool's hillside, Ithaca,
1917, whereit—

Bitlacomorpha clavipes, Pseudolimnophila luteipennis, Limnophila macrocera, Pilaria recondita,and Prionocera fuscipennis.)(Subgenus Limnophila Macquart)
18.34

Limnophila Macq.

1863

Poecilostola Schin.

Suit. ^ Buff., vol. 1, Hist. Nat. Ins., Dipt., p. 95.
Dipt. Austriaca, vol. 2, p. 551.

Limnophila is the typical subgenus of the group, the type having been
designated as Limnophila pictipennis by West wood in 1840.

No species

subgenus have yet been described from North America.
The larva of L. punctata (Meig.) was described by Behng (1886:195-of this197)

and by Gerbig (1913:158-161) as hving in wet earth, by Scheffer(1848: 10) and by Cameron (1917:63) as living in decaying wood.

Gerbigfound larvae in sandy soil near both standing and flowing water. The
larvae are very active, are rust-brown in color, and attain a length of 15
millimeters with a diameter of from 1.5 to 2 millimeters.
]M-ovided with several setae

andsctiferous projections.

The body is
The spiracular

disk (Plate XLVI, 214) is surrounded by four subequal lobes and an additional reduced dorso-median lobe; all of these lobes are fringed with long

The Crane-Flies of New Yorkhairs,— Part II

8G9

and each of the four paired lobes has a sensory bristle near the tip.

The four anal gills are not very prominent.
The mandibles areis of the hcxatomine type.
two small, broad, blunt, sawlike teeth just beyond

The head capsulesickle-shaped, withmidlength.

—

L. pictipenms
Brauer (1883:55) found the larvae of another species
between wet decaying leaves in bogs.
His figure of the head capsule (1883, pi. 1, fig. 10) shows a typical hcxatomine head (Plate XLVI, 213). Beling (1879:51-52) fomid this species(Meig.), the type of the genus—

sand of a small, dried-out brook bed.

in theCroup Ulomorphae

Genus Ulomorpha Osten Sacken (Gr. Ula + shape) \1869

Ulomorpha 0. S.

Larva.-— Form slender.

Mon. Dipt. N. Amer., part 4, p. 232.

Body covered with a rich golden-yellow pubescence.

Rpiracular

disk surrounded by four unequal lobes, the ventral pair the longest, lying subparallel, fringed

with exceedingly elongate hairs.

Spiracles small.

narrow, at tip expanded into a spatula.

with a single very long lateral tooth at its base.
projecting, bladelike.

Head capsule narrow, the dorsal plate

Mandible hinged, blade very long and slender,
Maxilla densely golden hairy, outer lobe

Antenna with a very long, tapering, apical

papilla.

Mentumnotchitinized.

Pupa.-— Cephalic crest prominent, each lobe with three setae.

Pronotal breathing horns

very long and slender, sinuous, cylindrical, at apex split into two flattened divergent lobes.

Mesonotum short, very convex, unarmed.
segment.

Leg sheaths

short,

Wing sheaths attaining end of second abdominal

ending before tip of third abdominal segment; hindtarsi

Abdominal segments divided into two rings, posterior ringwith a subterminal transverse armature of stout black spines and a few long setae; sternal
armature stronger than that of dorsum. Pleura armed with a few similar spines. Acidothecae of ovipositor very long and slender. Dorsum of segment 8 with a trapezoid of four
a little longer than the others.

lobes.

The genus Ulomorpha includes five known species, the genotype,
Ulomorpha piloseUa, of the eastern United States, and four western
North American species. The immature stages of the genotype are spentin rich organic mud in shaded situations.
Ulomorpha pilosella
1859(0. S.)

Limnophila pilosella O. S,

Proc. Acad. Nat. Sci. Phila., p. 242.

Ulomorpha pilosella is not uncommon in cool Canadian woods thruout
flies may be swept from

northeastern North America, and the adultCharles Paul Alexander

870

rank vegetation in such haunts. The flies bear a marked resemblance
to the species of the subgenus Lasiomastix of the genus Linmophila, but
the pupae, especially in the structure of the pronotal breathing horns,

The larvae are different from the relatcnl and somewhat similar larvae of the subgenus Dicranophragma of the genus Limnoi:>hila, of Penthoptera, and of similar hexatomine genera, in their pale
whitish yellow coloration instead of the deep saturated yellows and orangesof the genera mentioned.
All, however, have the quick, restless movements so characteristic of this group of crane-flies. Larvae found on
Bool's hillside, Ithaca, New York, on May 14 and 23, 1917, transformed
to adults on June 9.
are quite distinct.

Larva.

— Length, 8.5-9 mm.
Diameter, 0.5-O.G mm.

Coloration pale whitish yellow, the eighth abdominal segment suddenly whitish.

Form slender.
of

stiff,

Body covered with a dense golden-yellow pubescence. A transverse fringe

erect, short hairs at posterior margin of prothorax.

A number of pencils of setaeor solitary bristles on sides of segments, one on each annulus, longest near posterior margins
of segments.

219)

Behindthese setae, tufts of small hairs.

Spiracular disk

(PlateXLVI,surrounded by four lobes; ventral pair the longest, lying subparallel to each other,outer margin fringed with long, delicate hairs, those near tip coarse and easily broken, someof the hairs at tips exceedingly elongate; inner face of ventral lobes heavily suffused with darkbrown, this color more intense proximally; lateral lobes short, with an apical fringe of coarse
yellowish setae.

Spiracles very small, widely separated, located at base of lateral lobes.

Anal gills four, very slender, pale in color, the posterior pair a little the longer. On sternumof eighth abdominal segment, before gills, a transverse row of four long, coarse setae.

Head capsule (Plate XLVI, 215) very long and narrow, the dorsal plate slender, at end
expanded into a spatula; lateral plates a little shorter than dorsal plate. Labrum (PlateXLVI, 216) and epipharynx broadly transverse, projecting, the anterior margin narrower,
truncated, on either side near base with a brush of long hairs; disk of epipharyngeal region

with four setae, posterior pair a little the closer together; a few tiny papillae on ventral

two large setae near anterior margin and another
Mental region not readily distinguishable in, thematerial available, but at the most with little or no chitinization. Antenna (Plate XLVI,217) with basal segment cylindrical, a little narrowed medially, the truncated apex with about
two or three long setae and a very long, hyaline, sensory papilla which tapers gradually to
Mandible (Plate XLVI,
apex, this papilla about three times length of segment bearing it.

surface; clypeal region emarginate, withimmediately behind base of antenna.

218) hinged, the base slender but powerful, with the inner face deeply concave to receive

mandible in a position of rest; blade of mandible produced into a very slender hook whichisalmost straight, a little curved at extreme tip, at its base a very large, acute, flattened

blade which is more than half length of mandible itself; in its angle this blade has a second,microscopic, tooth; prostheca with about five long, stout, comblike teeth exceeding themandible in length, and an additional shorter, flattened blade marked with parallel grooved

JThe Crane-Flies of New Yorklines.

Maxilla with dense tufts of— Part II

871

long yellow hairs; outer lobe produced cephalad as ahyaline, flattened blade which projects from prothoracic orifice when head is retracted.

Pupa.

— Length,mm.

8..5-9Length of pronotal breathing horns, nearly 2 mm.
Width, d.-s., 1 mm.
Depth, d.-v., 1 mm.
/

Head, thorax, and appendages dark brown; pronotal breathing horns similar, but terminal

half gradually paler, the tip almost yellow; abdomen pale brown.

Cephalic crest consisting of two prominent lobes, each with three strong setae, the mostventral directed outward; just before primary crest and lying between antennal bases, a very
low, slightly bilobed crest which is not setiferous.
Labrum elongate, obtusely rounded atapex and separating labial lobes, the latter produced caudally into subacute points. Maxillary

palpi stout at base, narrowed to tip (Plate
least),

XLVH, 221).

XLVn, 220) high, feebly carinate medially.
andAntenna short (in female sex, at

extending but a short distance beyond knee joints of fore

slender, sinuous, transversely(Plate XLVIL 222 and 223).

mesonotum

PronotRim (Platewrinkled, at tip split into two flattened divergent lobes

Mesonotum very short and convex, with numerous black dots

which are most abundant anteriorly.
Lateral angles oflegs.

Breathing horns separated basally, very long

Two small setae on either side behind wing axilla.

blunt, but tip produced into a slender setiferous tubercle.

Wing sheaths ending opposite tip of second abdominal segment. Leg sheaths short, ending
just before tip of third abdominal segment, hind legs a little longer than others.

Abdomen with a narrow basal ring and a much broader posterior ring, the latter armed
before posterior margin with a transverse row of small black spines, strongest on pleura,

XL VI I, 224) the spines reduced in number, there
being from one to five (or in some cases none), and occurring only at or near ends of row;
weakest on dorsum; on dorsum (Plateends of row with two setae; on either side of median line a group of three closely approxi-

mated setae; usually segments 2 and 3 have the spines weak or lacking; segments 4 to 6
with two spines, and segment 7 with one spine, but in some specimens the number is slightly
Sternites (Plate XLVH, 225) with the intermediate segments (4 to 6) having
about twenty spines in an almost continuous row which as a rule is uninterrupted; ateach end of row about two strong setae; near base of posterior ring a narrow transverse
area with two setae at each end.
Pleura with a few powerful spines, small or lacking on
increased.

more numerous on posterior segments, there being usually two
on segments 5 and 6, and four on segment 7; on pleura at about midlength of posterior ring and nearer dorsal side, three black setae in transverse alinement,these somewhat longer on basal segments; opposite basal ring a stout seta.
Female cauda
basal segments, larger andon segment(Plate4,thre^;

XLVn, 223) elongate; tergal valves slender, slightly upcurved, near apex with asharp black spine whichside before apex.

isdirected dorsad, laterad, and caudad; two weak setae on either

Dorsum of segment 8 withfour lobes; posterior pair elongate, slender,

curved, and divergent; anterior pair blunt, small, andmore approximated; just ventrad ofthese lobes a stout seta; pleural region with two powerful spines, above theof which is a stout seta; a seta near ventral margin.

Ncpionotype.— Ithaca, New York, May 30, 1917. No. 88-1917.
With tvpe larva, reared June 9, 1917.
Paratypes.— Larvae,' May 30, 1917. Pupa, June 13, 1917.
Neanolype.

—more dorsal

Charles Paul Alexander

872

Genus Pilaria Sintenis (Lat., derived from the long antennal verticils)
1888
1919

Pilaria Sintenis.

Sitzber. Nat.-Ges. Dorpat., vol. 8, p. 398.

Eulimnophila Alex.

Cornell Univ. Agr. Exp. Sta., Mem. 25, p. 917.

Larva.'— Form moderately slender.

Spiracular disk surroundedby four unequal lobe.s,

Headlateral pair in some cases very reduced, elongate ventral lobes fringed with long hairs.

capsule of Ulomorpha type.

Maxilla densely hairy.

Mandible hinged, blade with one or two acute teeth at base.

Mentum not chitinized.

— Pronotal breathing horns elongate-cylindrical, tips

Pupa.'

split

into flattened

lobes.

Abdominal segments with three or four pairs of naked tubercles.

The genus Pilaria includes a group of species of the old genus Limnophila,
comprising Limnophila tenuipes and L. pilicornis and theirrelationships are plainly withallies.

Its

Ulomorpha rather than with Limnophila.

The genotype is Limnophila pilicornis (Zett.), of northern Europe.

Other

included species are L. tenuipes, L. recondita, L. imbeciUa 0. S., L. edwardi
Alex., L. quadrata, L. stanwoodae Alex.,and L. osborni Alex.,

of

North

America, and L. discicolUs (Meig.), L. fuscipennis (Meig.), L. subtincta(Zett.), and probably other species, of Europe.

The immature stages are spent in mud or moist earth. In Europe,by Gerbig (1913:
163-164) and by Cameron (1917:63). P. fuscipennis is described by
Behng (1886: 197-198) as Hving in mud near a ditch. Gerbig (1913 164Pilaria discicolUs (Plate XLVIII, 232) has been found:166) found the larvae (Plate XLVIII, 231) in a similar situation.

Brauer(1883:54) describes them as living between decaying leaves in swamps.
Cameron (1917:63) states that the larvae are found in decaying wood,
this record possibly being an error.
According to Beling, the pupal duration is not more than ten days.
In America, P. tenuipes has been discussed by Hart (1898 [1895]: 204-205) and by Malloch (1915-17b:223224), as stated under the discussion of the species.
The species of the genus Pilaria may be separated by the following
keys:
LarvaeVentral lobes of spiracular disk elongate, heavily marked with brownish black; coloration
pale yellow
P. tenuipes (Say) (p. 873)
Ventral lobes of spiracular disk short, pale; coloration deep yellow.
P. recondita (O. S.) (p. 874)

Pupae
1,

Pronotal breathing horns short, black; lobules of cephalic crest blunt and rounded.
P. quadrata (0. S.) (p. 875)
Pronotal breathing horns elongate, pale, brownish yellow or yellow; lobules of cephaliccrest elongate, finger-like

2

The Crane-Flies of New York2.

— Part II

873

Antennal sheaths of male elongate; pronotal breathing horns longer, pale yellow.
P. lenuipes (Say) (p. 873)

Antennal sheaths of male short; pronotal breathing horns shorter, yellowish brown.
P. recondita (O. S.) (p. 874)

Pilaria te7iuipes (Say)
1823
1869

Ldmnobia tenuipes Say. Journ. Acad. Nat. Sci. Phila., vol. 3, p. 21.
Limnophila tenuipes O. S. Mon. Dipt. N. Amer., part 4, p. 210-211.

Pilaria tenuipes is a widely distributed crane-fly thruout eastern North

The immature stages are commonly found in the mud ofswamps, or near streams and other bodies of water. This is the unknown
Limnophila described by Plart (1898 [1895] 204-205), and also consideredin much detail by Malloch (1915-17 b: 223-224), who found the pupae
along the banks of the Sangamon River in Illinois.
America.

:

Larva.

— Length, 16 mm.
Diameter, 1.4-1.5 mm.

Coloration of living larva, pale brownish yellow.

Body covered with a long, dark-colored,

appressed pubescence, more conspicuous on posterior segments.
near base and apex of segments.
distention,

coveredwith

Antepenultimate segment

numerous transverse rowsofof

Lateral pencils of setae

body capable of globular

microscopicroughened points.

Spiracular disk (Plate XLVIII, 230) moderately large, surrounded by four lobes; ventral
lobes long and slender, inner face with closely approximated, transverse, brownish black lines
which cause entire face to appear dark; near tips of lobes these black marks tapering out

into a long point; basal parts of dark marks subcontiguously hollowed out interiorly to forma large pale area below spiracles; ventral lobes fringed with long, pale hairs, some of themexceedingly elongate.

Lateral lobes very small, blunt, tending to be reduced, bearing shortfringes of dark hairs.

Anal gills slender, pale in color.

Head capsule of LTlomorpha type and not very di.fferent from that of the type genus;
narrowed gradually behind to near tip where it expands into
Epipharyngeal region of labrum and maxillae fringed with dense tufts

dorsal plate broad basally,a very large spatula.

and brushes of long yellow hairs.
Maxillary lobe relatively small but elongate, hyaline,
tapering to flattened apex.
Antenna (Plate XLVIII, 229) with basal segment elongate,
bearing at its tip an elongate apical papilla which is a Httle longer than the segment, bluntly
rounded at its tip, and delicately sculptured. Mandible (Plate XLVIII, 228) very long,
hinged, at its base an acute tooth equal in length to about one-third length of mandible; inthe type larva, the left mandible a little longer than the right mandible. Mental region
not chitinized.

Pupa.

— Length, 10-15 mm.
Width, d.-s., 1.5-1.6 mm.
Depth, d.-v., 1.6-1.8 mm.

Coloration dark brown; pronotal breathing horns light yellow, extreme bases brownish.

Cephalic crest small, composed of three slender, finger-like lobes which are tipped withstrong setae; on front, before crest, two setiferous lobes.

Labrum small, bluntly roundedCharles Paul Alexander874
at apex.

Labial lobes oval.

Maxillary palpi slender.

elongated, reaching to just beyond wing root.
elongate,cylindrical,

sinuous,

Antennaoffemale moaerately

Pronotal breathing horns (Plate XLIX, 233)

transversely wrinkled, apex scarcely enlarged but deeply

split on inner margin; a setiferous tubercle ventrad and laterad of breathing horns.

with a very high anterior mediancrest.

ThoraxA slender, setiferous tubercle above wing axil,

and two others on either side of median line. Sheaths of halter s long and slender. Leg
sheaths ending about on a level, or those of hind legs the shortest and those of fore legs alittlelonger.

Abdominal segments divided into two annuli. Tergites (Plate XLIX, 235) on posterior
ring with a caudal row of blunt, naked tubercles; at end of row two setae; on either side of
median area and just in front of row, a large setiferous tubercle; at base of ring two naked
tubercles, one on either side of median lin&; two slender setiferous tubercles near marginof ring.
Basal ring with four naked tubercles.
(The third pair of tubercles found in the
pupa of Pilaria quadrata is vestigial.) Pleura with four tubercles, two on each ring, thebasal one of each ring setiferous, the posterior one naked.
Sternites with six naked tubercles
on basal ring, arranged in three transverse pairs; on posterior ring at base two setiferous
tubercles, each with two bristles, directly behind last naked tubercle of basal ring; at caudal
margin two or three large tubercles near end of row and about four or five small naked tubercles
between. Female cauda (Plate XLIX, 234 and 236) very elongate, tergal valves slightly
upcurved, terminating in a sharp spine and with two setae on outer face. Eighth tergite
with four elongate lobes; posterior pair blunt, directed laterad; anterior pair elongate, slender,

A blunt lobe on pleural region.

with two setae laterad of each.

Pleura with three or four

powerful tubercles, with a seta located between the more dorsal pair.

Sternal region withfour small setae, two on either side of Taroad median area.

Ne.pionotype.-~ Orono, Maine, July 1, 1913.

Neanotype.

— With type

No. 50-1913.

larva.

Pilaria recondita (0. S.)
1869

Limnophila recondita 0. S.

Mon. Dipt. N. Amer., part 4, p. 212-213.

Pilaria recondita is a common crane-fly thruout the northeastern
United States. The immature stages are swamp inhabitants, and arevery frequently found in exactly the same situations as are those of P.
tenuipes.
P. recondita belongs to the same group as P. tenuipes, and thepupae of the two species are very difficult to distinguish.
Larva.

— Length, 15 mm.
Diameter, 1.2 mm.

Color a uniform light yellow.

Body covered with a long,appressed, yellow pubescence and with a few long setae.

Spiracular disk very small, in a position of rest almost closed, surrounded by four lobes;
ventral lobes moderately elongated, fringed with long, golden-yellow hairs which are longest
at tips of lobes; if bent backward these elongate hairs extending to beyond gills; inner faceof lobes almost

unmarked, with only a delicate brown line extending from tip toward baseThe Crane-Flies of New York— Part II

875

for a distance equal to about one-half length of lobes; lateral lobes small, subdorsal in position,

separated by a narrow notch, their inner faces opposed to each other, margin fringed withshort, golden-yellow hairs.

Headcapsule as inAnal gills four, moderately elongated.

P. tenuipes.

Antenna with sculptured

apical papilla tapering to

Mandible withblunt tip; besides this papilla, an even longer, hyaline, flattened blade.
apical biadelike part shorterand stouter, with two subequal stout triangular teeth at base

(Plate XLVIII, 227).

— Very similar to pupa of P. tenuipes, but smaller. Antennal sheaths of male

Pupa.short.

P. tenuipes but still much longer thanOn abdominal tergites, along caudal margin

Breathing horns a httle shorter than inin P. quadrala, of a pale yellowish brown color.
of posterior ring,from four to seven naked tubercles between the setiferous tubercles(inMale cauda. (Plate L, 237) with dorsal lobes stout, cylindrical,
narrowed at tips, divergent, direited caudad and ventrad; on outer face before tip a slender
Segments onseta; ventral lobes blunt, with a flattened ventral tubercle at base of notch.
dorsum with posterior lobes blunt, straight, directed caudad and shghtly laterad, but not so
P. tenuipes, four orfive).

strongly as in P. tenuipes.

———

Nepionotypc.
Orono, Maine, .July 3, 1913.
Neanolypc.
No. 112-1917.
Ithaca, New York, emerged June 11, 1917.
Paratypes.
Pupa, Orono, Maine, placed in rearing as a fully grown larva, June 26, 1913;
emerged as an adult male, July 3, 1913, showing a pupal duration of seven days. Larva,
Orono, Maine, July 5, 1913 (No. 74-913).

Pilar ia quadrala (0. S.)
1859

Limnophila quadraia 0. S.

Proc. Acad. Nat. Sci. Phila., p. 241.

quadrala is a widely distributed spring and early summer
The immature stages are very similar to those of P. lenuipes
and P. recondita. A pupa was found by Dr. Needham in the Indian
Spring, Ithaca, New York, where it was found floating among the water
cress.
From this pupa an adult female fly was reared. On June 3, 1917,
the writer found two fully matured male pupae in Chickaree Woods near
Pilaria

species.

Ithaca.
There had been a very heavy rainstorm on the preceding day,and the low spots in the woods had been converted into small ponds,

many of the insects that normally live in the mud or beneath the decaying
leaves being forced to the surface.

The pupae of P. quadrala, as wellas an abundance of Tipula larvae, were found clinging to small islandsof debris floating on these temporary woodland pools.
emerged on June 3.

The adult flies— Length of cast pupal skin, 9-12 mm.

Pupa.-

Coloration almost black, including pronotal breathing horns; abdomen more dusky gray.

Cephalic crest small, black, trilobed, each lobe with a seta at apex.
blunt at tip.

Labial lobes rounded.

Labrum narrow,

Sheaths of maxillary palpi elongate, tapering to theCharles Paul Alexander

876
slender points.

Antenna short in both sexes.

Pronotal breathing horns

moderately elongate, cylindrical, transversely wrinkled, at tips smooth,slightly enlarged.

(PlateL, 238)andflattened,

A tubercle with two long setae above wing axil. Two sstiferous punctures

on dorsum on either side of median line. Wing sheaths ending before tip of second abdominal
Leg sheaths ending before tip of third abdominal segment, the tarsal sheaths
ending about on a level or the hind legs shorter.
Abdominal .segments with tergites (Plate L, 2.39) 2 to 6 provided with eight naked,
discal tubercles, arranged in four transverse pairs, the third pair more approximated; lateradof third pair of naked tubercles, two small setiferous tubercles; on seventh segment one of thefour pairs of tubercles lacking; near caudal margin of segments a transverse row of weak,
setiferous tubercles; on either side of median line, in alinement with discal tubercles andsegment.

just anterior to the transverse setiferous row, a large tubercle provided with three

Pleural region carinate, each segmant

setae.

armed with four slightly curved tubercles: anterior

one solitary, setiferous; sacond ons solitary, nakad; third one with two or three setae; posterior
one bifid, naked. Sternites (Plate L, 240) with six naked, discal tubercles corresponding
Subterminal armature weak, ends of rows
to those of tergites but reduced in number.
tuberculate;

an isolated setiferous tubercle ventrad and laterad

Cauda (Plate L, 242) elongate;

dorsallobes

ofelongate-cylindrical,

ends of rows.
directedMalecaudad andslightly dorsr. 1, tapering to acute tips; three weak setae on outer ventral face; ventral lobes

blunt,

much shorter than dorsal lobes, with a blunt median lobule between them at theirtergum provided with a large, blunt, median tubercle, with two
which are directed caudad and with two tiny lobes on either side in
front; sternum v/ith a transverse row of four separated setiferous cubercles; caudal margin
with a transverse row of about eight or nine pale tubercles on either side, the outermost thebase; eighth segment onlarge posterior lobes

largest; a small

seta above second tubercle at ends of row.

Female cauda(PlateL,241) very long and slender, subacicular, sternal valves a little shorter than tergal valves;
tergal valves terminating in blunt cylindrical points.

—

Neanotype.
Male pupal skin, Ithaca, New York, June 3, 1917.
Paratypes.— Pupa, a male skin with type pupa; a female skin, Ithaca.

Subtribe Hexatomaria

The subtribe Hexatomaria comprises a well-defined division with but
four known genera, three of which are North American and are consideredin this paper.

The only other group of crane-flies with which the species

may be confused are certain of the Limnophilaria, especially the Ulomorphagroup of genera.
The larvae havo the labral sclerite of the head capsule large, separated
from the remainder of the capsule by a distinct suture. The epipharyngeal
region is restrict(Kl to the anterior median part of the sclerite, and is provided with two large tubercles on either side, which are tipped with twoor "".hree hyahne, cylindrical papillae. Between these papillae is a brushThe Crane-Flies of New York— PartII

877

The mental region
The dorsal plates of the head capsule are widely separated from each other by a median split.
The pupae are often armed with spines or tubercles on or about thehead and the thorax. The lateral abdominal spiracles are large andof hairs surrounding two pairs of setiferous tubercles.

isnot chitinized.

functional.

The generaof thesubtribc Hexatomaria

may be separated by thefollowing keys:
Larvae1. Coloration of body a deep saturated orange-yellow; spiracular disk with ventral lobes
unlined with darker, bearing at tips a few very long hairs.
Penthoptera Schin. (p. 891)
Coloration of body pale yellow, whitish, or greenish; spiracular disk with lobes linedwith dark brown or black2
Hexatoma Latr. (p. 877)
2. Size small, form slender (length 14-15 mm., diameter 1-1.3 mra.)
Size larger, form stouter (length over 15 mm., diameter over 1.6 mm.)
Eriocera Macq. (p. 881)
.

.

.

.

Pupae
1.

2.

Pronotal breathing horns short and stout, red at base and apex, the remaining partdark-colored, transversely wrinkled; horns bent strongly toward each other at tips.
Penthoptera Schin. (p. 891)
Pronotal breathing horns not as above
2
Size small (length under 10 mm.)Hexatoma Latr. (p. 877)
Size larger (length over 12 mm.)
Eriocera Macq. (p. 881)

Genus Hexatoma Latreille (Gr. six + I cut)
1809
1818
1818
Larva.

Hexatoma Latr.

Gen. Crust, et Ins., vol. 4, p. 260.
Nematocera Meig. Syst. Beschr. Zweifi. Ins., vol. 1, p. 209.
Anisomera Meig. Syst. Beschr. Zweifl. Ins., vol. 1, p. 210.

— Size small (lengthof Hexatoma megacera

about 14 mm.).

Spiracular disk sur-rounded by two pairs of lobes, ventral pair the longer, inner face marked with a narrowbrown line which is expanded at its inner end. Head capsule with lateral angles of labrum
elongate and densely clothed with hairs.

Pupa.

— Size small (length under 10 mm.). A large spinous tubercle on scape

No median projection on mesonotal scutellum.
Wing pads with cell Ri very small andof antenna.

Pronotal breathing horns short and straight.

M with but a single branch reaching wing margin.

The genus Hexatoma includes a small number of forms with a chiefly
Holarctic distribution, there being about eleven Palearctic, two Ethiopian,and one North American

species so far described.

The adult flies of

the North American species, Hexatoma megacera, are common on vegetation

along the banks of rather large streams.

They are discussed herein inunder the account of this species. The adult flies of some of thelarger European species have habits quite like those of the genus Eriocera,detailCharles Paul Alexander878
asdiscussed

describes in

elsewhere in

this

paper

(page

705).

Riedel(1909:29)

some detail the habits and swarming of H. hicolor (Meig.),

the males of which are very active in the forenoon during the hours ofbrightest sunlight, the females resting on the willow branches near by.

Similar habits are recordedby Riedel (1910:30) for H. saxonum(Lw.).

The immature stages of the European species are practically unknown,
the only original reference being that of Von Roser (1834), who states thatthe larvae of H. nigra Latr. live in the sand along the banks of streams.

Hexatoma megacera (0. S.)
1859

Anisomera mej2cera 0. S.

Proc. Acad. Nat. Sci. Phila., p. 21:2.

The adult flies of Hexatoma megacera are on the wing during the monthsofMay and June and may be swept from the rank vegetation along thestreams from which their larvae emerged.
lation, resting positions,details, areThe following notes on copu-

egg laying in nature and in captivity, and otherquoted from published field observations (Alexander, 1915 c:

143-145)—

May 14, 1911 This usually rare insect was common on a grassy plot of land along
Cascadilla Creek [Ithaca, New York].
The flies sit on the blades of grass, the long antennaeof the male directed straight ahead.
The males are very poor fliers and prefer to drop to thegroimd when disturbed and clumsily work their way off along the ground. When approachedfrom the side they are much more easily alarmed and fly away. When approached fromabove, they do not move until the stick, finger, or whatnot, is within a couple of inches,when they remove the fore feet from the support and, on nearer approach, fall to the ground.
When in copulation, the female tries to disengage by rapidly vibrating the wings in attempted
flight, repeating this often, from every one to five seconds until disengaged or exhausted.
The male can disconnect himself at will. In copulation the female is always uppermost
unless exhausted, when both sexes lie flat on a grass-blade.
The female has the head up,
the male the head downward; copulation always takes place on a vertical support, usuallya blade of grass, sometimes a plant stem. The sexes remain in copulation for quite a longtime and are perfectly motionless. All of the legs of both sexes are on the support unlessin a position where this is physically impossible, in which case as many as possible are used;
the hind legs of both sexes are held at right angles to the support, the forelegs in front. After
copulation the female generally drops to the ground, the male, after a few moments' rest, flies
away. Specimens in copulation were found in abundance from 2 to 4.30 p.m. when no more
could be discovered. From 4.30 to 7 p.m. solitary males were common, but no females could
be found on the grass-blades. At 4.30 p.m., a few females were found clinging to the trunksof the willow trees about two feet from the ground.
At 5..30 p.m., females were noted insmall groups over the water, evidently engaged in oviposition, as they frequently dipped
down to the surface. These latter were in company with a large swarm of dancing empidid
flies (Rhamphomyia).
Of the great numbers that were picked from grasses in the afternoon a considerable proportion were females and toward 5 p.m. they commenced egg-laying
on the sides of the containing vessel, large shell vials. By 7 p.m. the sides of the vials were
black in places with the large, dark -colored eggs. This data would seem t3 place the time
The females are very good fliers and often travel for longfor oviposition at about sunset.

The Crane-Flies of New York— Part879

IIdistances before alighting as is shown toward sunset when they fly for long stretches upThe males are rather poor fliers, due in part, possibly, to the weight of the long
stream.
antennae and, as stated before, this sex prefers to skulk rather than fly. When the males
fly, they do so heavily and seize the first support that they collide with and hang on, occasionally flying on immc^diately to another support.
When the male comes in contact with a stem,he very often ascends to the top by means of a part-flying, pan-climbing motion and, on reaching the summit, flies off to another place.
As a rule the flies, especially the females, alight ona single grassblade, but very often the males are observed on two blades, the legs of one side
on one blade and those of the opposite side on the other; when the body thus hangs between
the stalks, the tarsi diverge from one another, whereas on a single support, the legs converge.
Hexatoma was preyed upon by lirge numbers of a scatophagid fly that occurred in greatabundance in this vicinity and seemed to be subsisting almost entirely on these Iflies. At
least twenty of these pred'ice:}us flies were noted with Hexatoraae and this species seemed
to constitute the principle insect enemy of the crane-fly.
They would lurk on the grassblades and sally forth after their prey, carrying it back to some point to feed upon it.
It isprobable that the blood is taken since the body of the Hexatoma appeared almost uninjured
when examined. On an old beam where males had a habit of walking up the vertical face,a small spider's web was found, in which eleven specimens were entangled, two being still
alive; eight of these were males, the remaining three, females.
The males especially can walk up smooth surfaces, as glass, moving the legs alternately
and awkwardly. The first pair taken were in copulation but in placing them in the vial
they became disengaged and ran about in the container. After a short time they began to
copulate in the tube.
May 15, 1911
pair were taken in copulation at 10 a.m.; at 8 p.m. they were still incoitu, but this is exceptional as most of the pairs disengage very readily.
In the morning
the species is very active and although the males do not fly far, they fly readily and it isdifficult to pick them up by hand.
The females are excellent fliers especiallythe morning.
Several pairs were taken in copulation and each pair was isolated in a separate vial in order
to ascertain the number of eggs per female.
The clutch was determined by dissection.
When the captive insects began to oviposit, the eggs shot out from ihe body, at first slowly,
then more rapidly, one per second, later much slower again, the eggs being extruded one ata time. The total period of o'l/iposition required seven minutes; at the end of sixty seconds,
in the space between sixty and seventy seconds, eighteen eggs were laid, or 1.8 per second.
Toward the end of egg-laying, the eggs appeared much more slowly, one in two seconds. The
eggs are quite sticky or viscid and adhere to the glass. When the female is in danger ofdeath, as when she falls into the water, she begins, at once, to deposit the egg-complement.
In nature it seems probable that one egg is laid at each descent to the water. As soon as
the female touches the water, although she has not deposited an egg all day, she immediately
starts to deposit the oblong black eggs.
After the last egg is expelled the muscles of theovipositor still go through the motions of expulsion.
One specimen was placed in the waterand as usual began to deposit her eggs. She was decapitated, laid eleven eggs and tried tolay still more but failed.
The number of eggs laid varied from 316 to 372 with an averageof 347; the time required for ovnposition varied from seven minutes to seven minutes and forty
seconds.
In most cases the number of the egg-complement is probably between 300 and 400.
J

—A

m

The greater part of the larval existence is probably spent in the water,
and it is only when the larvae are fully grown and ready to pupate thatOn April 26, 1914, gravel from the bank of CascaCreek was carefully examined, but no signs of larvae or pupae were

they come to land.
dilla

On May 6, however, the same bank was examined
and about ten larvae and seventy-five pupae were found. Sometimes theimmature stages are very abundant. On May 12, 1917, near the place
to be discovered.

Charles Paul Alexander

880

just described, the writer found larvaeand newly transformed pupae ingreat numbers, the former pupating in the dry sand rather distant fromthe water's edge.

In one dry patch of sand on a rocky ledge, sixty

specimens were found in six square inches of soil. The insects are often
found in gravel or coarse sand that is thickly penetrated by grass roots
and rhizomes, rarely in pure gravel. The}- are most commonly foundThe immature
Hcxatoma are associated ^^dth larvae of Eriocera spinosa, E.

in soil that has l)een recently deposited after freshets.

stages of

Tipula bella, and Atherix probaljly variegata Walk., with pupae
and with many beetle^ such as Paederiis

cinerea,of Chrysops excitans Walk,,
littoran'us Grav.,

Dyschiriics

Gastrolohium bicolor (Grav.), Bledius sp., OmophronsphaericoUis

Say,

Tachistodes

discoideus (Dej.), Laccobius agilis

partiarius(Say),

sp.,

Anadaptus

Rand, and other species characteristicof the sandy margins of large streams.

The pupal duration of Hexatoma

and one-half days.
Larva. — Length, 14-15 mm.
Diameter, 1-1.3 mm.

is sixColor of body, pale brownish yellow.

Body covered with rather abundant appressed hairs, the subterminal distended part ofabdomen with transverse rows of very short spines or setae, there being from thirty-five toSpiracular disk (Plate LI, 246) surrounded by four lobes; ventral lobes
forty such rows.
the longer, with a fringe of a few long hairs at apex; on inner face a long, narrow, brown
mark, extending from the tip inward, the proximal end expanded; lateral lobes with numerous
long hairs which are gradually shorter toward base of lobes. Spiracles circular, situated atbase of lateral lobes; a dusky mark extending from spiracles dorsad.
Head capsule of usual hexatomine type, as described for Eriocera longicoriiis (page 888).
Entire general features and details of mouth parts very similar to those of Eriocera longicornis.
Labrum (Plate LI, 243) transversely oval, with frame strong and chitinized, anterior medianpart produced into a small lobe (Plate Lll, 250) bearing two lateral papillae and two setiferous
tubercles surroundedby short hairs; lateral margins of labrum produced into prominent

lobes directed proximad and cephalad, densely hairy and entirely protecting anterior marginof labrum.

Antenna (Plate LI, 244) with apical papillae shorter than segment, the largest
IMandible (Plate LI, 245) long and slen-

papilla transversely sculptured (Plate LII, 251).

der, the lateral teeth more accentuated than in Eriocera; largest tooth with a flattened trun-cated blade in its axil, this in some cases broken up into two or three small blades; basadof largest lateral tooth a flattened lobe which is barely indicated in the species of Eriocera
studied.

Maxilla with outer flattened blade conspicuous; palpus nearside, similar to the condition

Pupa.

its

base on innerobtaining in Eriocera; just laterad of palpus a powerful seta.

— Length, 9.2-9.6 mm.
Width, d.-s., 1.2 mm.
Depth, d.-v., 1.4 mm.

The Crane-Flies of New York — Part II881

Pupa (Plate LI, 247) very similar to that of Eriocera longicornis, differing only in its small
size,

greater development of seapal spine, lack of projection on mesonotal prcscutum,

anda few lesser characters.

Cephalic crest (Plate LI, 248 and 249) as viewed from beneath,very different in shape.

Fore pair of legs much shorter than the others, ending just beyond

posterior margin of second abdominal segment; hind pair of legs extending far beyond theothers, ending beyond midlength of third abdominal segment; in some .specimens the tarsal

segments much closer to posterior margin of third abdominal .segment, but usually a marked
difference in tips of tarsi of the various legs.

Cephalic crest as viewed from side, triangular,

Viewed from beneath, lobes con.spicuand separated by a deep
median split. Spine on scape of antennae very large, conspicuous. Tubercle on labrum
Antennal sheaths of male very long, those of female much shorter.
strongly developed.
Pronotal breathing horns short, straight. Mesonotum strongly wrinkled along medianWing pads dark, venation not showing clearly but, ifline; scutellar lobe not developed.
characteristic of Hexatoma alone.
made out, the very short cell R' and the reduced
Posterior leg sheaths extending beyond level of middle legs, these, in turn, being longer
ending in an acute point directed strongly forward.

ously triangular, pointed, lying parallel or slightly divergent

Mthan sheaths of fore legs.

Abdomen (Plate LI I, 2.52) with about thirty-four spicules on sternite 4, about thirty
ina straight, uninterrupted row on tergites 3 and4,and about twenty on

tergite 5.

Chaetotaxy about as in Eriocera longicornis, but the seta lying ventrad of spiracle on pleuritea
much farther ventrad and very weak. Male cauda with sternal lobes strongly rounded,enlarged,and bent suddenly dorsad.

Female cauda as thatof male; ovipositor

viewedfrom side with an obtuse notch; from beneath, sternite obtusely pointed and with a deep
median .split; from above, tergite almost flat across caudal margin, the lateral angles rounded,with a deep median split.

— Ithaca, New York, May 1914.
— TypeMay
Paratypes. — Several hundred larvae and pupae from type
Nepionnlype.

6,

K-eanotype.

locality.

Genus Eriocera Macquart
18.30
18.38

1838
1848
18.50

1857
1859
18.59

1912
1916
Larva.

2, 191.3.

locality.

(Gr. wool + horn)

Caloptera Gu^r.
Voyage de la Coquille, Zool., Ins., pi. 20, fig. 2.
Eriocera Macq.
Dipt. Exot., vol. 1, p. 74.
Evanioplera Gu^r.
Voyage de la Coquille, Zool., vol. 2, part 2, p. 287.
Plerocosmus Walk. List Dipt. Brit. Mus., vol. 1, p. 78.
Allarithmia Loew.
Bernstein und Bernsteinfauna, p. .38.
Oligomera Dolesch. Natuurk. Tijdschr. Nederl. Indie, vol. 14, p. 387.
Arrhenica 0. S.
Proc. Acad. Nat. Sci. Phila., p. 242.
Physecrania Bigot. Ann. Soc. Ent. France, ser. 3, vol. 7, p. 123, pi. 3, fig.
Androclosma Enderlein. Zool. Jahrb., vol. 32, part 1, p. 'M.
Thous. Ins. Japan, add. 2, p. 471.
Globericera Matsumura.

1.

— Spiracular disk surrounded by four lobes which are rarely (as in Eriocera cinerea,

subobsolete, inner face lined with brown or black, tips with fringes of moderately long hairs.

Head capsule long, narrow, the constituent plates very slender; dorsal plate completely
divided tho contiguous or approximated behind.

Labral sclerite large and conspicuous,

sensory tubercles and papillae crowded on median cephalic region.

7

Mandible long, acuteCharles Paul Alexander882

at tip, with two teeth at about midlength.

IVTaxillawith outer lobe greatly prolonged into

Antenna cylindrical or clavata, with three or four long papillae at tip.
Mentum not chitinized, in E. cinerea with a flattened rectangular plate on either side, thisarmed with numerous hooks and spines.
Cephalic crest of various shapes and sizes, very reduced in Erwccra spinosa.
Pupa.a flattened blade.

—

Antennal sheaths of males of several species (E. spinosa, E. lomjicornis, E. cinerea) very
Pronotal breathing horns of various shapes, acutely
long, extending beyond end of wing pad.
pointed in E. spvnosa, short and blunt in several species.

Head and thorax often wit

spines or tubercles on scape of antenna, on labrum, or (in E. spinosa) on face of eye;one on dorsum of second abdominal segment (in E.
Abdominal segments with a conspicuous transverse armature of spines neartubercle on scutellum (in E. longicornis)
spinosa).

iaposterior margin.

,

Lateral spiracles large, distinct.

Eriocera is an extensive genus (including approximately 150 species)of medium-sized to large

flies,

mostofwhich are tropical.

The genus

has not been found in Europe, but elsewhere it is represented by a host

The habits of the adult flies have already been noted (pageThe immature stages are spent in sand or gravel near runningof species.

704).

water,

more especially along large streams. A more complete accountby Alexander and Lloyd (1914) and by Alexanderof the genus is given(1915 c: 148-152).

The occurrence of the flies is somewhat local. During an entire summer
Maine in 1913, the wiiter did not find a single specimenof collecting inany species; and Dr. Dietz has stated that the only living individual
which he has found was a single male of Eriocera spinosa taken in the
Pocono Mountains in Pennsylvania. On the other hand, the flies areoften found in countless numbers, and several species may be foundassociated together. Thus, at Ithaca, New York, in the sandy gravel
along Cascadilla Creek, the immature stages of four species of Eriocera
and one of the closely related Hexatoma occur together in unlimited
numbers.
The species of Eriocera may be separated by the following keys:ofLarvae1. Lobes surrounding spiracular disk obsolete or nearly so; a flat, chitinized plate with serE. cinerea Alex. (p. S86)
rate margins on either side of mental region
Spiracular disk surrounded by four slender lobes; no plate as described above on mental

region2.

2

Very large (length 40-45 mm., diameter 4-5 mm.); spiracular disk with ventral lobes
narrowly lined with black, iimer ends of each forked, Y-shaped; literal lobes narrowhUned with black, inner ends of marks expanded
E. spinosa (O. S.) (p. SS.'j)\
Smaller (length under 30 mm., diameter under 2.5 mm.); spiracular disk not marked
as above

3

The Crane-Flies of 5j^ew York3.

— Part II

883

Ventral lobes of spiracular disk bearing one or two very long, dark setae in addition to
the shorter yellowish fringe; inner face of each lobe with a capillary black line which
is suddenly expanded at its inner end into a triangular brown mark, the two marks
inclosing an oval pale area between their proximal ends; lateral lobes with a capillary
black line
E. fultonensis Alex. (p. 8^0)Ventral lobes of spiracular disk with apical fringe consisting of numerous long, pale setae;
inner face of each lobe lined with pale brown, at about midlength gradually expanded
into an elongate-triangular mark, the two marks inclosing a linear pale area betwp'^ntheir proximal ends; lateral lobes with a brown line
E. longicornis (Walk.) (p. 888)

Pupae
1.

Size large (length 25 mm. or over); pronotal breathing horns tapering to acute tips;
cephalic crest small, reduced to four small tubercles; cell Mi on wing pad present;a strong spinous tubercle on either side of median line at base of second abdomuial
tergite; a tubercle on eye
E. s-pinosn (O. S.) (p. 883)
Size smaller (length under 18 mm.); pronotal breathing horns blunt at tips; cephaliccrest prominent; cell Mi on wing pad lacking; no tubercles on second abdominal tergite

or on eye

2.

3.

2
E. longicornis (Walk.) (p. 888)
No tubercle on mesonotal scutellum
3
Pleurites of abdominal segments with a transverse row of three setae ventrad and slightly
caudad of spiracle; antennae of male elongatedE. cinerea Alex. (p. 883)
Pleurites of abdominal segments with two stout setae dorsad and caudad of spirarh;
antennae short in both sexes
E. fultonensis Alex. (p. 890)

A tubercle on mesonotal scutellum

Eriocera spinosa (0. S.)
1859
1869

Arrhenica spinosa 0. S. Proc. Acad. Nat. Sci. Phila., p. 244.
Eriocera spinosa O. S.
Mou. Dipt. N. Amer., part 4, p. 252-253.

commonest of the large species of the genus
The larvae occur in great numbers beneathrocks in rapid water in the autumn, when they form a considerable proEriocera spinosa

in eastern

is

the

North America.

When about to pupate they gobanks and live for some time in the sand or gravel.
The habits of the larvae have been discussed by Alexander and Lloyd(1914:16-17) and by Alexander (1915c: 149).
The larvae were found on May 1, 1913, along the banks of Fall Creek,
Ithaca, New York, in considerable numbers.
They were associated withyoung and mature pupae of E. longicornis, which were emerging in great
numbers at the time. On May 27, both larvae and pupae of E. spinosa
were found to be very abundant, the larvae being more numerous in thewetter places, the pupae in the drier spots. They occurred at various
distances from the water's edge, from within a foot of the margin to as
far back as eight or ten feet.
The pupae are found in short, more or lessvertical, burrows, from one to three inches below the surface.
Not often
portion of the insect life in the streams.

to the neighboring

884Charles Paul Alexander

were larvae and pupae found in close proximity to each other.
of E. spinosa, as well as of

all

Pupae

other species of the tribe as known, arevery active when removed from their burrows, wriggling rapidly to and
fro,

27,

and are exceedingly tenacious of life: Larvae, as found on May
were mostly contracted; a fonv, however, were expanded and had thesubtcn-minal segment of the alxlomen swollen.

In this regard it may bementioned that almost all of the larvae of crane-flies that live in the sandor

mud along the banks of streams have this ability to inflate the endof the abdomen.

Larvae of Eriopterini, of Pediciini, and of Hexatomini
have been observed with this conspicuous enlargement. It is undoubtedlyused to propel the larva thru the soil by alternate expansion and contraction of the segment.

Larvae of E. spinosa were placed in breeding jars on May 13 and adultemerged on the 28th. It is prolmble that the pupal stage is notlonger than from ten to twelve days, at the most.
On May 30 a large
number of lai*vae and pupae were brought into the laboratory in a bucketflies

of gravel.

Some of the fully matured pupae transformed in the pailwhile being brought to the laboratory.

The larvae are carnivorous. Their powerful, sickle-shaped mandible?
are capable of inflicting a painful bite on tender parts of the hand.
S. G.
Rich placed larvae in dishes together with the nymphs of various dragon

The smaller of the nymphs were eaten by the Eriocera larvae, thus
confirming previous observations on the carnivorous habits of the species.

flies.

Larva.

— Length when

fully extended, 40-45 mm.

Diameter, 4-5 mm.
Color varying from very pale whitish to rather dark brown; in life, the skin showing
conspicuous bronzy reflections.
Spiracular disk (Plate LIV, 262) surrounded by four slender elongate lobes, one pair being
lateral,the other ventral, in position; inner face of lateral lobes with a capillary black hne,

this beginning as an enlarged black spot just ventrad of spiracle,

reaching tip of lobe; dorsal
outer edge of lobe with a dense fringe of long, conspicuous, reddish hairs, inner edge of row
beginning just laterad of spiracle where the hairs are very short, gradually becoming longer

where they are as long as the lobe itself; ventral lobe with a capillary black line on
proximal edge, this dividing at base of lobe, the lower branch running along ventral margin
to tip,and approaching its fellow of the opposite side on median line of body; adense fringe of conspicuous reddish hairs at tip of lobe and continued on outer dorsal sideof stigmal field

for a short distance toward base; a few dusky brown spots on stigmal field between spiracles;

two sniall hairs between spiracles. Spiracles rather small, widely separated. Underneathcaudal lobes and behind penultimate swollen segjnent, four anal gills, shon, stout, cyhndrical.

tThe Crane-Flies of New York— Part885

IIthe inner pair directed caudad. Head capsule (Plate
the lateral pair directed outward,
1.8-2 mm. (acrossits length, measuring 3.5-3.S mm. by
toproportion
in
broad
267)
LIV
third length of segone
than
n.ore
not
short,
antennaoftip
Papillae at
dorsal plates).
prominent conical tooth at midlength, such as
ment. Mandible (Plate LIV, 271) lacking a
species.
otheris found in E. cinerea andPupa.— Length: male, 20.5-27 mm.; female, 25-2S.5 mm.
Width, d.-s.: male, 3.4-3.9 mm.; female, 3.4-4 mm.
Depth! d.-v. male, 4-4.2 mm. female, 3.-5-4 mm.
;:

pale yellowish to dark brown or almost black,
In life pupae varying in color from veryof old pupae; body often showmg bronzy
thoraxandheadofthe deepest color being thatreflections.

•

.

r

level of antennae; viewed from
Cephalic crest very reduced, scarcely projecting beyond
angles produced into small pomted lobes
beneath somewhat quadrate, the anterior lateralside a second pair of lobes is seen, these bemg
bearin- a small seta at apex; viewed from
Spine of antennal scape very
setiferous.
subequal to anterior lobes in size, and likewiseof eye with a conspicuous
surfacecaudalInner
ventrad.
large somewhat curved, directed
Tubercles at ba.se oftubercle.
Tentorial region produced into a small mediantuberclechitinized; a small seta above eachstrongly
tips
their
together,labrum very large, close
Pronotal breathing horns
small seta on each cheek.
of these labral tubercles and anotherand rather pointed at tips, the organ arcuated so that

Ion- slender broad at ba.se, flattened

Mesonotal scutellar lobe (Plate LV, 280) prominent, ratherofWing pad light brown, venation showing very clearly, the presencestron-ly projecting.
antennae in male sex being found in this species alone
cell m[ in connection with elongate
oppositeLeg sheaths with tarsal sheaths ending on a level, about
in eastern North America.
end of third abdominal segment.
tubercle on either side of median line.
Second abdominal tergite with a conspicuous basalrows of spines, there being about
Abdominal segments (Plate LVI, 283) with subtermiiial
destitute of spines but with fourand7
6
tergites2
to5;
tergites
twenty to twenty-two on
conspicuous setiferous tubercle on ventrosubapical setiferous tubercles; tergites 2 to 7 with amargin, bearing aring; eighth tergite concave on posterior

apex is bent strongly ventrad.

cephalic angle of each posterior
pair of strong apical tubercles

on either side of -median line.

rather restricted, longitudinally wrinkled.

Pleural region of

abdomenSpiracles large, elliptical, transverse, placed about

Three small setiferous tubercles ventrad and caudad of
opposite midlength of segments.
on dorso-cephalic angle of each pleuron. S.ernites
tuberclespiracle, and another similar
segments 4 to 6 with from sixteen to twenty
on segment 3 with two spines on each outer angle;
setferous tubercle caudad and dorsad
isolatedanspines;tenspines- segment 7 with about
setiferous tubercle about midlength of posterior ring;of end's of row segments 4 to 7 with arow of strong spines which are intersegment 8 lacking soft pleural region, bearing an apicalsmall space on median line of venter, there being about
rupted only on dorsum and for atwenty of these spines in the circlet.

Male cauda (Plate LV, 281 and 282) with ninth sternite
a small lobe on ventral side at end ofrounded swollen, with a deep median furrow bearing
points separated by a V-shaped
conicalstrongtwo
intocaudadsplit- ninth tergite produced
dorsad, each one a little split near tip on
notch these points directed caudad and slightly
Female Cauda (Plate
at about midlength.
outer face and with a prominent lateral tooth^^Charles Paul AlexanderA^epiono^ype.— Ithaca, New YorkAe«7i«////;e.— With type larva

Para/^pes.- Numerous larvae

Mav 1 lOH

and pupae with types. May 1-15,

1913.

Eriocera cinerea Alex.
1912

Eriocera cmerea Alex.

Psyche, vol. 19, p. 169-170 pi
13

fig

9

^'''' The larvae were'"

foutd'TMiTTe "iJlT"" ""'T","' "^'r"•' '""''^^ "f Cascadilla Creek,
aiiica, JNew
It"!c-.
Now YorkTl!'"'°"f
Yo k. They
have beenfound at;,past few yea,., but always
in scanty numbers.

various dates durinir theLarvae foundilpri

Atneux, and otherinsects.
The larvae arc stouter than/««,«on« and are pale whitish yellow,

touale on24,

May 16.

tho.seofE,uite devoid of the greelsh

infs

Additional larvae and pupae
were found onMay

and a few pupae on May 30 1917

(IOm'zT ^T*^' 'r™ 1^; '""^--''-'rf-cribed by.Alexander and Lloyd

later in this paper.

Larva.— Length, 15-lfi
1mm.
Diameter, 2-2.2 mm.
Diameter
Color light yellow.

Head capsule long and narrow, measuring
w.;tK

-^^

about 1

prox,„,a, anterior a„g,e.
produced if.:!:;.''^

I

hv n ')7r, ,

j

,

,

abL ; riitrurt; r"'°

The Crane-Flies of New York— Part887

II

genus (Plate LII, 254); this'structure located on either side of capsule on ventral face, aflattened, subrectangular plate whose surface, except at base, is densely set with small spinesand large

pits;

inner

margin provided with large, acute spines, beginning at about one-

third length of sclerite, gradually enlarged toward tip, at inner angle acute; these teeth inter-

rupted before outer posterior angle, which terminates in a blunt, flattened lobe.
cylindrical, apex obliquely truncated, terminating in

Antennaa slender apical papilla which is longer

than the segment that bears it, broad at base, tapering gradually to tip; two or three long
apical setae; small auditory plates at about one-third length of segment.
IjII,

Mandible (Plate

255) a powerful, slender, curved hook, at about midlength with a strong pointed tooth

bearing in its axil a smaller tooth; as is usual in the genus, an egg-shaped chitinized pieceisolated in

one of the ventral tendons of mandible.

dible, outer lobe persisting as

Pupa.

Maxilla arising just ventrad of man

a very elongate, bladelike organ.

— Length, 13-15 mm.
Width, d.-s., 1.6 mm.
Depth, d.-v., 1.7 mm.

Head and appendages dark brown, thoracic dorsum a little paler; abdomen with posteriorrings of tergum and sternum dark brown, producing a banded appearance.

CephaHc crest (Plate Lfll, 257) very large and conspicuous, consisting of two rounded
lobes behind, each tipped with a strong seta; anterior part of crest directed ventrad and consisting of two lobes, the larger bearing a strong seta on outer face;viewed from front, theseby a very narrow, U-shaped, median notch; crest of female a Uttle
smaller. Two blunt tubercles on scapal segments of each antenna.
Junction of clypeus andlabrum with two tubercles, above and slightly laterad of each a strong seta. A strong seta
on cheek below eye. Labrum broad, very obtusely rounded at tip. Labial lobes rectangular, widely separated.
Maxillary palpi blunt at tips (Plate LIII, 258). Antenna of male elongate, exceeding wing and ending opposite base of last tarsal segment of hind leg.
Pronotalanterior lobes separated

breathing horns short, slender, cylindrical, scarcely longer than cephalic crest.

Two long

curved setae and a smaller straight seta laterad and ventrad of base of each breathing horn.
Lateral angle of thorax with two setae; two long setae above wing axil (Plate LIII, 256).

A strong seta on either side of mesonotum and a group of two small setae in front of eachof these.

Wing sheaths extending to base of third abdominal segment. Leg sheaths extend-

ing to base of fourth abdominal segment; tarsal sheaths ending about on a level, or thoseof fore legs considerably shorter.

Abdominal segments (Plate LIII, 259) divide 1 into two subequal rings. Chaetotaxy as
pleura with a seta on dorsal margin of basal ring; a transverse row of three setaeon posterior ring, lying ventrad and slightly caudad of spiracle (as in E. spinosa) tergum withbasal ring unarmed, posterior ring with a subterminal row of sharp black spines; two longsetae at each end of row and a few small setae at intervals along row; a solitary seta on basal
lateral part of posterior ring; armature weaker on posterior segments, on segment 7 being
reduced to four .separated groups of setae, the outer groups with a single spine; sternum withbasal ring unarmed, posterior ring with a subterminal transverse row of stout black spines
with two long setae at each end of row and an isolated seta laterad and caudad of end ofrow; at base of ring on either side a group of two setae, the lateral one the smaller. Male
Cauda (Plate LIII, 260) with the sharp dorsal lobes directed dorsad, rather acute at tips, twofollows:;

Charles Paul Alexander

888

above, these setae seen to be separated by a
deep U-shaped notch; eighth segment with a dorsal pentagon of five closely approximatedsetae on outer face before tips; viewed fromgroup of threelobes; just laterad of these a

setae, the posterior

one long and slender, theanterior one short and stout; pleural region produced into a long lobe tipped with a slenderseta;

on sternum two small setae on either side.

Female cauda (Plate LIII, 261) elongate,

tergal valves very long and slender, two delicate setae on either side before tip and a stouterone at tip.

—

Ithaca, New York, April 28, 1917.
Neanolype.— Fall Creek, Ithaca, May IS, 1917
Paratypes.
Larvae and pupae with types.

Nepionotype.

—

Eriocera longicornis (Walk.)Anisomera longicornis Walk.

1848
1869

Eriocera lofigicornis O. S.

Eriocera longicornis

List Dipt. Brit. Mus., vol.

1,

p. 82.

Mon. Dipt. N. Amer., part 4, p. 253-254.

probably the commonest species of the genus
The adult flies are sometimes very abundant,isin the eastern United States.

occurring in swarms in late afternoon and early evening in May, someof theswarms numbering thousands of individuals.

the day, theflies

At other times of

may be found resting quietly on bushes.

live in the sand near the water's edge.

The larvae
The pupal duration is seven days.

The detailed life history of this species is given on pages 704 to 708.
Larva.

— Length, 17-19 mm.
Diameter, 2-2.3 mm.

Color, greenish brown.

Body covered with a long, appressed, dark pubescence.

Penultimate segment of abdomencapable of great distention and destitute of pubescence; last segment of body conspicuously

narrowed.

Spiracular disk (Plate LIV, 265) surrounded by four slender lobes, the ventral

pair the longer, bearing at tip elongate hairs, some of which exceed the lobes in length; onlateral face at about midlength a small pencil of hairs; on ventral face one or two long setae;

inner face of ventral lobes lined with pale brown, beginning as a narrow brown mark at tip,
at about midlength gradually

expanded into an elongate triangular mark, the two lines

inclosing between their inner ends a pale linear mark; lateral lobes similar to ventral lobes,

fringed with long yellow hairs which are longer than the lobes; inner face of lateral lobes linedwith pale brown.

Spiracles rather large, separated by a distance equal to about one and one-

A few setae in transverse alinement on last segment

half diameter of one.

Anal gills pale.

behind lateral lobes.

Two pairs of short black setae behind gills.

Head capsule and mouth parts very similar to those of Hexatoma, dorsal plates of capsuleseparate from each other, not fused as in the Ulomorphae; inner margins of dorsal plates
straight and parallel.

Labral sclerite broadly transverse, narrowed at ends, lateral margins

produced into long lobes which are densely tufted with short, golden-yellow hairs. Median
lobe of epipharyngeal region projecting, provided with two large sensory papillae, one on
ether

side,

and a few othersetilerous papillae near tip, surroundedby numerous hairs.

The Crane-Flies of New York— Part889

II

Antenna elongate, a little narrower at base, at tip with three or four hyaline, seta-like papillae
which are of various diameters and shorter than the segment, the largest of these papillae
delicately sculptured with transverse lines.
Mandible acute, curved, at about midlengthwith a blunt, flattened tooth, this with a smaller similar tooth in its axil.

Maxillary blade

very long and slender, about half length of capsule.

Pupa.

— Length: male, 13.2-15.2 mm.; female, 14-15.4 mm.
Width, d.-s.: male, 2.1-2.2 mm.; female, 1.8-1.9 mm.
Depth, d.-v. male, 2.1-2.3 mm.; female, 2.2 mm.
:

Young pupae very pale; soft abdomen almost white; chitinized anterior part of body very
Older pupae much darker, the chitinized part becoming black with a bronzy
reflection; abdomen very dark brownish gray; breathing horns dark brown on apical half.
pale brown.

Cephalic crest (Plate LV, 275) very prominent, elongate, tapering to the subacute tips;
lobes with blunt tubercles behind, as well as four long setae on each lobe, three on dorsal

margin and a longer one on lateral face at about midlength; viewed from in front, lobes
separated by a broad, square or U-shaped notch; ventral part of crest produced forward

between antennal bases as a depressed lobe bearing a stout seta on either side.
on antennal scape very prominent.

with a small rounded knob cephalad of each.

diamond-shaped.

Maxillary

palpi

Tubercle

A slightly smaller tubercle on either side of clypeus,
very

Labrum truncated.

broad,

rectangular,

tips

Labial lobes roughly
truncated.

Antennal

sheaths of male greatly elongated, enlarged at base; viewed from beneath, the swollen bases
just above and proximad of inner margin of eye, with
Antenna of male exceeding wing pads, those of female
ending just beyond wing base. Pronotal breathing horns short and stout, straight, transversely wrinkled, directed cephalad, dorsad, and laterad; when viewed from beneath, comThoracic notum convex; me.sonotum transversely
pletely concealed by large cephalic crest.
wrinkled (Plate LV, 277); median lobe of mesonotal scutellum projecting dorsad and caudad as a blunt point (Plate LV, 272). Two or three setae above wing axil. Lateral angles
Wing sheaths attaining end of second abdominalof thorax subacute, with a weak seta.
segment. Leg sheaths ending before caudal margin of third abdominal segment; tarsi ofhind legs the longest, the two inner pairs ending about on a level (Plate LV, 273).
Abdominal segments (Plate LVL 285) divided into a basal and a posterior ring; tergitea
on posterior ring with a subterminal transverse row of spines, these varying from about thirtytwo on segment 3 to about fourteen on segment 7; these rows of spines interrupted on dorsomedian line; at each end of row, three long setae, and two additional groups of setae interspersed along row; two setae on either side at anterior -lateral angle of ring; tergites on basal
ring unarmed; sternites on posterior ring with a subterminal transverse row of from twentyfour to thirty-two spines, with two setae at each end of row; an isolated seta on caudolateral margin, close to pleura; a group of two approximated setae near base of posterior
Sternites on basal ring unarmed; pleuritos
ring, on either side, about at level of spiracles.

nearly contiguous on medianline,

scapal tubercle described above.

on basal ring with a solitary seta at about midlength, but slightly nearer dorsal margin;posterior ring with two setae dorso-caudad of spiracle, and a third seta ventrad of it.

Cauda (Plate

A'al

•

LVL 286) very blunt, much narrower than remainder of abdomen; ventral

lobes very blunt; dorsal lobes short, stout, ending in sharp points directed dorsad, on out'^rface a long

and a short seta; segment 8 with a dorsal trapezoid of four lobes, the posterior

S^Charles Paul Alexanderrpair the longer, each with two setae; anterior
pair of lobes a little more widely
separatedlaterad of latter pair of lobes, a tubercle bearing
three setae; a long, powerful seta on
pleuratwo setae on either side of median line of sternum. Femalecauda (Plate LV, 274) similar'
but ventral lobes more pointed, sUghtly exceedinglevel of dorsal lobes.

Nepionohjpe.— Ithaca, New York, April IS, 1917.
Neanotype.— Fall Creek, Ithaca, May 2, 1913.
Paratypes.— Several hundred larvae and pupae withtypes.

Eriocera fultonensis Alex.
1912

Eriocera fultonensis Alex.

Psyche, vol. 19, p. 168-169, pi. 13, fig. 7.

Eriocera fultonensis is a rather common but usually

the northeastern United States.

local species thruout

The larvae are found in the same situ-i-

tions as are described for the other species of
the genus, in sand or gravel
near the margins of usually large streams. On
May 30, 1913, larvae were
found in considerable numbers along the banksof Fall Creek, Ithaca
New York, where they occurred in company with numerous
larvae andpupae of E. spmosa, a few large tabanid larvae, asmall tabanid pupa
and the following beetle associates: Bembidion,
Schizogenius Tacliy-'
Gastrololjium, and a few others.
The pupal duration is seven days (froni
May 31 to June 6, 1913).

Larva.— Length, 18-26 mm.
Diameter, 2-2.3 mm.
Color, pale flesh yellow; anterior segments

Body long and slender.

of body a little darker
Spiracular disk (Plate LI V, 266) with ventral lobes

lateral lobes shorter; ventral lobes at theirtips with one or two

which are from two to three times length of lobes; inlong and slendervery elongate blackish hairs

addition to these the usual apical fringeof yellowish hairs not exceeding lobes; nearbase on outer side a s:,iall pencil of hairseachventral lobe with a very delicate capillary blackline which expands abruptly at
its'innerend into a brown area, these two areas inclosing
between their proximal ends a more or lessoval pale area (in some specimens the innerends completely
in others the brown lines are notencircLn- this pale area while
continuous over the disk); lat3ral lobes shorter,

fringed with yellow hairs which are longer

black hne.

similarly

than lobes; inner face of lobes with a capillar,-Spiracles rounded oval, widely separated.

Head capsule and mouth parts almost as in Hexato^na

nierjaccm and Eriocera hnguonns
as already described; epipharyngeal region(Plate LIV, 268) produced into a hemisphe-.calrounded lobe which is densely covered with fine hairs;
two large papillae on either side neartip, between them a terminal tuft
of long yellow hairs surrounding twolong slender setiferous papillae. Antenna with sensory papillae
at tip short, about one-third length of segment.

Pupa.— Length, 14-16 mm.
Width, d.-s., 1.8-2 mm.
Depth, d.-v., 2.5-2.7 mm.

The Crane-Flies of New York— Part II

891

Fully colored pupae dark brown; cephalic crest paler; pronotal breathing horns pale yellow,

darkening into brown at tips; wing pads light yellow, with dark venation showing clearly;
pleurites of abdomen of a darker

brown than stemites or tergites.
Body somewhat similar to that of E. longicornis, but general form much stouter.

Cephalic

LVI, 287) prominent, tuberculate, consisting of four lobes, the posterior lobes
somewhat the larger, on posterior and lateral faces with two stout setae; a stout seta on
crest (Plateventral face of anterior lobes.

Scapal spine lacking.

a small setiferous tubercle above each.

Clypeal tubercles large, blunt, with

Labrum bluntly rounded at apex.

elongate, diamond-shaped, tips rather acute.

Labial lobes

Pronotal breathing horns rather long and

Mesonotum (Plate LV, 279)
Wing pads usually showing venation clearly on palebackground; vein r connecting R\ with Ri+i distinctive of this species, lack of cell Mi
transversely wrinkled, longer than cephalic crest.

slender,more convex than in E. longicornis.

separating this pupa from that of E. spinosa and E. brachycera. Legs sheaths ending. about
on a level, the hind tarsi a little longer than the two inner pairs.
Arrangement of setae on abdomen (Plate LVL 288) about as in E. longicornis. Pleurawith two stout setae dorsad and caudad of each spiracle, and a weak seta ventrad of spiracle
and close to it; basal ring with a single pleural seta; spicules on caudal margin of posterior
ring small and numerous, on intermediate segments about forty in number; seventh sternite
with about four to six spines at each end of row, the broad median area devoid of spines.
Female cauda (Plate LV, 276) with tergal valves exceeding the long sternal valves,s^arce'y directed dorsad (this condition may be compared with that in E. longicornis). Malecaada with abdomen bluntly rounded at tip.

— Ithaca, New York, May 30, 1913.
— With type larva, June 1913.
Paratypes. — Larvae and pupae with types.
Nepionotype.

Ne n'tli/pc.

6,

Genus Penthoptera Schiner
ISGj

Penthoptera Schin.

(Gr. sorrow + wing)

Wien. Ent. Monatschr., vol. 7, p. 220.

— Spiracular disk surrounded by four blunt lobes, the ventral pair a

Larva.-

inner face not marked with darker, at tip with one or more long setae.
as in Eriocera.

little the longer,

Head capsule about

Coloration a deep saturated yellow.

— Cephalic crest with lobes rounded, setiferous.

Pupa.'

on head or thorax.
a flattsneJ heal,

No distinct tubercles or spines

Pronotal breathing horns short, stout, cylindrical, apex expanded into

stem caarsely wrinkled, base enlarged.

Abdominal armature weak.

Spiracles not well developed.

Penthoptera is a small genus which includes four European and three
American species, two of the latter occurring in tropical America. The
eastern North American Penthoptera alhitarsis, discussed below, has been
considered in some detail by the author in another paper (Alexander,
1915c: 152-157).

Charles Paul Alexander

892

Pentho'ptera alhitarsis 0. S.
1809

Pcnthoptera albitards O. S.

Mon. Dipt. N. Araer., part 4, p. 257-258.

The larvae of Penthoptera albitarsis are usually not uncommon in richorganic mud in shaded places thruout the range of the species.

Larvaemany sizes, some very small, others apparently almost fully grown,
may be found at a single time. This would probably indicate that theofspecies emerges at intervals thruout thesummer, rather than that it isdouble-brooded.
Larva.

— Length, 10-12 mm.
Diameter, 1-1.2 mm.

Colbr bright chestnut-yellow, anterior half of body richer- and deeper-colored; thoracic
segments suffused with brown; skin with a silky, iridescent reflection.

Body provided with numerous long, appressed hairs. A few setae on body, the following
being the most conspicuous: one on lateral do. sal margin oflateral lobes;

last

segment, near base ofa series of four groups of one or two in each row across dorsal surface of thethree thoracic segments at about midlength; a group of two or three long setae on sides nearcaudal margin of segments.

Subterminal enlargement of abdomen with about twenty-fivetransverse rows of fine points.

Bpiracular disk (Plate LVII, 292 and 293) with four blunt lobes; ventral lobes densely
fringed with long, pale hairs, those toward ends of lobes longer; one ormore elongate setae

near tip of each ventral lobe, these being longer than lobes themselves; lateral lobes witha similar fringe of rather short, yellow hairs; spiracular disk almost free from dark markings,

a pale brown line extending dorsad from each spiracle and an indistinct brownish line

along ventral margin of laterallobes.

Spiracles

capsule rather broad, dorsal plate with inner

circular.

anterior

Anal

angles

gills

pale.

Head

Labrum(Platefour,

rounded.

LVII, 289) almost as in Eriocera spinosa, the extreme cephalic epipharyngeal parts with theand seti'erous tubercles; lateral papillae bearing at their tips three or four
two pairs of setiferous tubercles, a basal larger pair
and a more apical smaller pair.
Mandible (Plate LVII, 291) a little more curved than isusual papillae

slender pegs; between these papillae

usual in this subtribe, inner margin with a double tooth at about midlength.

Pupa.

— Length, 10-10.5 mm.
Width, d.-s., 1.4-1.5 mm.
Depth, d.-v., 1.5-1. G mm.

Thorax dark brown, wing and leg sheaths paler; pronotal breathing horns dark brown,swollen bases and tips much paler, light orange; abdomen brownish yellow.

Cephalic crest consisting of two widely separated rounded lobes behind, each bearing two
setae; anteriorly the crest appearing as a large depressed lobe between antenna! bases, witha very large, stiff seta on either side.

A powerful seta on each side of region of clypeus. A

seta on genal region between eye and sheath of maxillary palpus.

or a little truncated at apex.

pentagonal in outline.
sheaths.

Labrum evenly rounded

Labial lobes widely separated, roughly rounded or indistinctly

Maxillary palpi broad, ending bluntly beneath or just before antennal

Antennae endiag just beyond wing root in female, considerably longer in male.

The Crane-Flies of New York— Part II893

Pronotal breathing horns (Plate LVII, 294) short, stout, cylindrical, apex expanded into aflattened circular head, stem coarsely

and transversely wrinkled, base enlarged; breathinghorns widely separated at their bases, but bent proximad so as to be almost contiguous at theirtips;

two strong setae laterad of base of breathing horn and an additional one in front of it.

Mesonotum transversely WTinkled, with a distinct carina anteriorly (Plate LVIII, 296). Twogroups of two setae on either side of median line, with an additional solitary seta; two longer
setae above wing axil.

AVing sheaths ending before tip of second abdominal segment.

Legsheat'is ending at from two-thirds length to opposite end of third abdominal segment; tarsal

sheaths ending about on a level, or, in some specimens, the hind tarsi a little longer than the
Others (Plate LVIII, 297).

Abdomen indistinctly divided into a narrow Vjasal ring and a broader posterior ring; basal
ring further very indistinctly subdivided into

two subequal annuli.

Abdominal armature

weak; on sternites a subterminal transverse row of delicate spines with two setiferous tubercles
at each end of row; on posterior ring two setae on either side at about midlength; tergites

with four groups of two or three setae near posterior margin and an additional group of twosetae on lati'ral margin of posterior ring near base; pleural region with a stiff seta on extreme
anterior part of basal ring.

Spiracles distinct; a group of

two setae caudad and slightly
Femaleventrad of each spiracle, with an additional solitary seta caudo-ventrad of these.

Cauda (Plate LVIII, 298) with tergal valves of ovipositor only a little longer than sternalvalves, at tip ending in a short, rather blunt point directed dorsad; on outer face before tip
a short, stiff seta; segment S on dorsum with a close trapezoid of four irregular lobes; two
setiferous tubercles on dorsal and lateral part of eighth segment, the more dorsal of thesewith two setae, the lateral one with a single seta; sternum with four stout setae, of v.'hich twoare lateraland two are median inposition.

Male cauda (Plate LVII, 295, and Plate

LVIII, 299) with sternal valves short and blunt; tergal valves slender, ending in an acutepoint directed dorsad; a few short setae on outer face before tip.

Nepinnniype.— Ithaca, N-w York, May 25, 1917.
Bool's hillside, Ithici, June 5, 1917.
Paralypes.
Abundant larvae and a few pupae with types.

——

Neanoti/pe.

Subtribe Polymeraria

Genus Polymera Wieck-mann
1821

Polymera Wicd.

Polymera
species,is(Gr.

many + part)Dipt. Exot., vol. 1, p. 40.

a tropical

Americang(^niisincluding fifteen

described

one of which, Polymera ycoryiae Alex., occiu's in the southeastern

United States. A single additional species, P. magnijica Meunier (1906:
The385), has been deso'il^ed from the Baltic amber (Lower Oligocene).
only species concerning the ecology of which we have any record is P.
geniculata Alex., which has
in Porto Rico.

hxm found living in crabholes beneath rocks

In this connection the long-horned deinoceritine mosquitoes

894Charles Paul Alexander

which live in similar habitats should be considered.

Howard, Dyar, andKnab (1915:213) say, in describing these mosquitoes:Those crab-hole inhabiting species possess peculiarly developed antennae in order,
as we
suppose, to enable them to detect the approach of iheir crustacean hostand fly out of the
h-.ics before being overwhelmed in the water in thebottom by the incursion of the crab
whose body must completely fill the entrance to the hole.
'

It is

curious and suggestive that the males of Polymera should likewise

possess elongatedand very complicated antennae.
Tribe Pediciini

The Pediciini constitutes a small tribe which seems to be divisible into
two well-marked subtribes, the more generalized Adelphomyaria indicatinga relationship with the Hexatomini.

The larvae of the Pediciini have the labrum broad and the epipharynx
The mentum is completely divided into two parts,
each halt with not more than four, usually three, teeth. The hypopharynx
is labriform.
The maxilla consists of two lobes, distinct and separate
usually feebly armed.

the Adelphomyaria, more or less approximated or fused in the PediThe mandible is powerful, ending in a strong apical point; thecutting edge has about four teeth; there is a simple tuft of setaeon theprosthecal region in the Dicranotae and in Pedicia.
The head capsulei:icaria.

very elongate, massive, and compact, with the posterior incisions veryi3

shallow.

In the Adelphomyaria the cauda is surrounded by four lobes
which are fringed with exceedingly elongate hairs; in the Pedicaria there

are two ventral caudal lobes, each tipped with a very few setae.
Theanal gills are four in number and are segmented, the terminal
segment "being more or less retractile.
In the Pedicaria prolegs aredeveloped

on the abdominal' segments of some of the genera.
All of the species of the tribe, so far as is known to the writer,arecarnivorous in their larval state, which is spent in mud or earth close towater.

The two subtribes of the Pediciini may be separated as follows:
LarvaeSpiracular lobes four in number, fringed with very long, delicate
„

.

,

,

,

-Spiracular lobes two

hairs.

Adelphomyaria (n.

895)
m number, ventral in position, each tipped with from six to eight setae.

Pedicaria (p. 899)

The Crane-Flies of New York— Part II
is as

The most important literature on the tn\x Pediciini
„

,.

.

~

Oenonl.

^^""^'^enej^

Pedjcia
Pedicia rwnsa::

TrirlXna immacnlata
Tricyphonn schiurri
Dicrnno,a bi.n.cuInU.
Drcranota tnmaculata
Dicranota hmncuMa
Dicranota bimaculaia
Rhaphidolabis tenuipes

^S!;,

S'^

Wesenbcrg-Lund. 1915: 335.

•

generalTri'-^/phont immaadata

Mows:

RchefTor, in Rossi, 1848:9.

..

^™ r^'^f - nvosa

895

Need'.ia.n, 1903:285-286; 1905:8.

Larva, general
Larva, pupa, generid,
Larva, pupa, genera
Larva, pupa, general.

'^ "/^;.

.

^^^'^^..^.^^^"'.

General(general''

10 Vfi- 195
iq'^-19r,
196.
l^eMeijc^-e 1916.
Behng 18.9.47.

•

i^arva,Copy.)

Wesenbe?g-Lund, 1915: 342-343.
1915-17 b: 219-220.
(Codv )

Needham. 1908 a: 212-214.

Larva, general

Subtribe Adelphomyaria

The division Adelphomyaria, as known, includes
Adelphomyia Bergroth, a curious genus

but the single genus

of small crane-flies which, in thesuggest the hexatomme subgeneral appearance of the adults, strongly
been associated with
Limnophilaria. The immature stages have not
tribe

of a

by rearing, and there is, of course, the possibility
the insect herein describedmistaken reference. The immature stages ofgroup it belongs,
recognized, however, and, no matter to what

the adult

flies

are easilyitdeserves subtribal rank under the Pediciini.

Genus Adelphomyia Bergroth (Gr.

brother + fly)

Mittheil. Naturf. Ges. Bern, 1890, p. 134.

1891 Adelphomyia Bergr.
of stiff setae, producing a spiny appearance.
Larva (supposition).- Body with pencils
which are fringed with exceedingly elongate
lobes
short
four
by
Spiracular disk surrounded
Head capless than the diameter o one.
Spiracles large, separated by a distancehairs.
Mandible acutely pointed.
behind.
exceptunitedfirmlyplatessule long and massive, all the
Antenna two-segmented, the terminal segment
Maxilla of two elongate separated lobes.
Mentum completely divided, eachlabriform.
Hypopharynx
with three small papillae.
{minula, supposition) or thelargestpair on each side thehalf with four teeth, the middle
supposition).
{amerkana,largestsecond from the inside thelobe with three setiferous tubercles.
Pupa (supposition^.- Cephalic crest small, eachwith light yellow. Mesonotum
tippedbroadlylength,inmoderate
Pronotal breathing horns
segment. Leg sheaths
abdominal
third
Wing sheaths extending beyond base ofunarmedarmature weak, especially
Abdominal
segment.
abdominal
fifthofextending to beyond baseon posterior segments.

four European
Adelphomyia is a small genus of crane-flies, including but
species from Africa. The
and three North American species, and a doubtful

Charles Paul Alexander896

insects resemble tiny species of the s^nus Limnophila.

The adult flies

are not uncommon on rank herbage, especially ferns, in woods and usually

As already stated, the immature stages have notbeen reared, but larvae found by the writer in Maine are referred withnear running water.

considerable confidence to Adelphomyia americana and A. cayuga, whilelarvae and pupae of another species taken at Ithaca, New York, seem to

The larvae show a curious combination of tribal
The general appearance and the structure of the spiracular

be those of A. minuta.
characters.

disk are altogether those of one of the Hexatomini; but the head capsuleand the details of the mouth parts indicate a relationship with the Pediciinithat cannot be denied.

Adelphomyia minuta Alex, (supposition)
1911

Adelphomyia minuta Alex.

Can. Ent., vol. 43, p. 287-288.

Adelphomyia minuta is a characteristic late spring species, common in
boggy woods and on vegetation along rapid streams. Larvae and pupae
which are referred to this species were sifted from organic mud taken on
Bool's hillside, Ithaca, New York, thruout May and early June, 1917.
The pupae referred to this species strongly resemble those of Dicranophragma but in reality are very different. The species is discussedherewith in the hope that it may be definitely recognized in the future.
Larva.

— Length, 4.5-5 mm.; caudal

fringe, 2 mm. additional.

Diameter, 0.4 mm.
Coloration, saturated yellow with a faint orange-brown tinge.

Form narrow, body tapering gradually to both ends, spiracular disk narrowed.

Body

clothed with a delicate appressed pubescence and numerous tufts of conspicuous stiff hairs

which produce a spiny or bristly appearance; the more conspicuous of these tufts located on
pleural region, there being three such rows on abdominal segments—

one on basal ring, thesecond and largest at base of posterior ring, and the third just before posterior margin ofsegment and more ventral in position; only the large intermediate tuft present on thoracic
segments, the small brush on anterior annulus of abdominal segments lacking.
Spiracular
disk (Plate LIX, 305) with four lobes; ventral pair not more than three times length of lateralpair; inner face of lobes margined with brown; at tip of ventral lobes a fringe of exceedingly
elongate hairs, which are from one-third to nearly one-half length of entire body and about
fifteen times length of lobes bearing

them; a stiiT sensory bristle located in black margin

at tip of lobe; lateral lobes with fringe of hairs relatively

much shorter, tho still long.

Spiracles large, close together, separated by a distance less than diameter of one, the middle

piece black, the ring pale yellow.

Anal gills four, slender, hyaline, each subdivided by con-strictions into four lobes which are gradually narrowed from the base outward, the last being
cylindrical.

The Crane-Flies of New York— Part897

II

Head capsule very compact, lateral plates united with broad dorsal plate except for a short
Labrum (Plate LIX, 300) broadly transverse; cephalic margin truncate or

distance behind.

very slightly concave; epipharyngeal region with about five transverse interrupted rows of

Mentum (Plate LIX, 302) of two entirely separated plates, each half with four teeth,

setae.

those at the ends the smallest, the outermost tooth tending to be reduced, the two middle
teeth of each side notably larger than the others;

outside of mentum a thin plate,

proximal margins nearly contiguous at median line of body.

its

inner

Hypopharynx labriform, pro-beyond level of both labrum and mentum; outer lateral angles densely hairy,
median posterior area with about eightsmall, hyaline spines. Antenna (Plate LIX, 301), in
caustic-potash mounts, hyaline; basal segment elongate-cylindrical, bearing at its tip about
two papillae as follows: a long, slightly curved, cylindrical papilla which is finely sculptured,and immediately pro.ximad of this a slightly smaller second segment of the antenna, bearing
near its tip three tiny papillae, an inner flattened subspatulate blade, and two longer cylindrical papillae; near base of this second an tennal segment a long seta.
Mandible (Plate LIX,
303) rather long and slender, apical point narrow; about four small lateral teeth near ventral
cutting edge, the most basad of these acute; a large dorsal tooth on cutting edge and twovery small acute teeth near base; on dorsal face of mandible two long setae, and two
somewhat shorter setae near heel of mandible; about five or six stout setae at prosthecal
region.
Maxilla (Plate LIX, 304) consisting of two elongated lobes which are separate fromeach other, the outermost the longer, the pale rounded palpus terminal in position; inner
lobe shorter, with three elongate setae, of which one is apical and the longest is sub-basal injecting

position.

Pupa.

— Length, 4.5—4.7 mm.
Depth, d.-v., 0.8 mm.

Color light yellowish brown, the thoracic dorsum paler; pronotal breathing horns darkbrown, apical quarter abruptly light yellow; abdomen brown, lateral and posterior partsof each segment darker.

Cephalic crest small, each lobe with three small setiferous punctures on anterior face;ventrad of crest between antennal bases a prominent median lobe.
truncated, indistinctly bifid.

Labrum broad; apex

Labial lobes large, with a deep LT-shaped median notch behind.

Maxillary palpi rather short and stout, ending before joint of fore legs (Plate LX, 307).
Pronotal breathing horns moderately elongate, curved slightly laterad, strongly divergent,
cyHndrical, of uniform diameter thruout their length.

Mesonotum not very gibbous (PlateThorax ^vith a high anterior median carina. Wing sheaths extending be3'ondbase of third abdominal segment, the venation indistinct. Leg sheaths rather long, endingat about one-third length of fifth abdominal segment; hind legs much longer than the

LX,

306).

others,middle legs a very little longer than fore legs.

Abdominal segments indistinctly subdivided into two rings, a narrow anterior ring and amuch broader posterior ring; abdominal armature very weak, lacking on segment 7; basal
annulus on tergites and sternites with a number of small pits on sides, these sometimes sparssor lacking; posterior ring with a transverse row of numerous long, stout setae or delicate
spines before margin.
Spiracles weak, at base of posterior ring.
Male cauda (Plate LX,
309 and 310) with sternal lobes elongate, contiguous except at extreme tips; tergal lobes
terminating in very slender and acute curved points, directed caudad and dorsad; tergite

Charles Paul Alexander898

8 swollen, with four blunt lobes, the posterior pair large, with their posterior faces setiferous.
Female eauda (Plate LX,308) with sternal valves inueh shorter than tergal valves and rather
blunt at tips; tergal valves broad at base, narrowed at tips, which terminate in acute blackspines, situated on lateral margin before apex, the spines directed dorsad, laterad, and caudad.

The larvae were common near Ithaca, New York, from May 10 toJune 5, 1917. The pupae are described from one male and two females
washed from mud from Bool's hillside, at Ithaca, on June 11, 1917. A
little later in June adult flies of this species were common at this location.
Adelphomyia americana Alex, (supposition)
1912

Adelphomyia americana Alex.

Pomona Journ. Ent., vol. 4, p. 829-831.

Larvae that were rather common in the rich organic mud from theStandpipe Woods, Orono, Maine, from July 1 to 14, 1913, are referredwith some doubt to Adelphomyia americana. They are unquestionably
congeneric with the species last described {A. minuta, supposition) andwith the form discussed in the following pages as A. cayuga.

A shorttime after the larvae of these three species were obtained, the adult flies

appeared in considerable numbers in the same situations and there seems
Associated with the larvae
to be but little doubt as to the reference.
of this species in the organic mud were a few larvae of Penthoptera,
Rhaphidolabis, and other species of crane-flies.

The larva of the present species averages larger than that of A. minuta,
when fully grown measuring 5.5 millimeters in length. In coloration
The ventral lobes of the spiracular disk have the dark
it is light yellow.
markings on their inner face much more extensive, the apical half being
The mouth parts are similar to those of A. minuta
suffused with brown.
as described, but the outermost of the two large teeth of each half of thementum is reduced in size so that only a single tooth is of conspicuous size.
(Described from larvae taken at Orono, Maine.

No. 66-1913.)

Adelphomyia cayuga Alex, (supposition)
1912

Adelphomyia cayuga Alex.

Pomona Journ. Ent., vol. 4, p. 831.

The supposed larva of Adelphomyia cayuga occurred with specimensof the preceding {A. americana, supposition) at Orono, Maine, from July

This is a smaller species than the preceding, measuring
1 to 13, 1913.
but 4 millimeters in length, and is much paler, the color being almost

The Crane-Flies of New York
white.

— Part899

II

The condition of the mental teeth is ahnost as in A. americana,but the outermost of the two large intermediate; teeth is a little larger.
The larvae were rather frequent in the mud beneath saturated moss.
(Described from larvae taken at Orono, Maine. Nos. 57- and 67-1913.)
Subtribe Pedicaria

The Pedicaria comprise a well-defined division of the tribe Pediciini,the more generalized Pediciae, withincluding two groups of genera—

the genera Pedicia, Tricyphona, Ornithodes. and Hhaphidolabina, andthe specialized Dicranotae, with the genera Dicranota, llhaphidolabis,and probably Polyangaeus.

The genera of the Pedicaria may be divided in the main as follows
Larvae1.

Abdomen without prolegs, but with raised welts on segments 4 to 7,

these covered.2
with a microscopic scurfiness. (Group Pediciae)
Abdomen with conspicuous cylindrical prolegs on segments 3 to 7, those with circlets
Group Ihcranotae (p. 906)
of conspicuous chitinized booklets around their ends
2. Abdominal segments 4 to 7 with raised welts on both dorsal and ventr il surfaces; sid-sof
Rhaphidolabina Alex. (p. 901)
hypopharynx not parallel, narrowed toward baseAbdominal segments 4 to 7 with welts on ventral surface only; sides of hypopharynx
3
subparallel
3. Size very large, when fully grown 40 mm. in length; mental plates without a small lateral
Pedicia Latr. (p. 899)
tooth
Size smaller, when fully grown under 30 mm. in length; mental plates with a small lateral
Tricyphona Zett. (p. 903)
tooth

The writer is unable to separate the genera of the group Dicranotae
with the material available, and is inclined to suspect the congenerousness
The characters given by Mallochof Rhaphidolabis with Dicranota.
(1915-17 b: 217) to separate the two genera do not hold at all in a series.
It is impossible also to

key the pupae with the present knowledge ofthe group.

Group Pediciae

Genus Pedicia Latreille (Gr. afield)
1809
1916

Pedicia Latr. Hist. Nat. Crust, et Ins., vol. 4, p. 255.
Daimiotipula Matsumura. Thous. Ins. Japan, add. 2, p. 463.

Pedicia is a small genus including but six described species, one occurringin Europe,

one in Japan, and four in North America.

Of the last-named,Charles Paul Alexander

900

two are eastern and two are western in their distribution.

They include

the largest and most beautiful species of the Limnobiinae.

The European Pedicia rivosa (Linn.) was found by Behng (1879:45-46)
living in brooks and springs, or in wet spots among saturated leaves andother debris, sometimes associated with the larvae of Tipida lutescens

The pupae live inFabr.

cylindrical verticalburrows, clothed in theand are able to move up and down in these passages
Pupation lasts from one to two weeks.
Needham (1903:285-286, and 1905:8) was the first to describe andfigure the larva of the commonest eastern species, Pedicia alhivitta Walk,
last larval skin,

Pedicia alhivitta Walk.
1848

Pedicia alhivitta Walk.

List Dipt. Brit. Mus., vol.

1, p.

37.

a beautiful fly, common and widely distributed
thruout the northeastern United States and Canada. The adults areon the wing in midsummer, and a few individuals may usually be foundThe much rarer and more local P. contermina Walk, is a vernal
in June.
species, on the wing in May and early June.
The larvae of P. alhivitta live in cold springs and beneath saturated
moss at the edge of streams. The writer has never succeeded in rearing
Pedicia alhivitta

isthis species to the adult condition.
Larva.

— Length, 40-44 mm.
Diameter, 5-5.5 mm.

Color dark grayish brown above, paler at sutures and on posterior half of body; paler
beneath, more grayish.

Body covered with a short, appressed, dusky pubescence.

Thoracic segments with a

Abdominal segments with a few delicate lateral
setae on posterior ring, at about midlength of segments.
Ventral creeping-welts on abdominal
segments 4 to 7 completely divided on median line, the welts covered with a microscopic
scurfiness.
Spiracles (Plate LXL 311) circular, separated by a distance about equal topencil of small setae on pleural region.

diameter of one, situated on a slightly protuberi'.nt elevation.

Spiracular lobes two, ventral

each with about six setae at tip.

Anal gills (Plate LXI, 315)in position, short, slender,

short,

stout at base, before tip a constriction cutting off the elongate conical terminal

segment, which is partly telescopic within the next basal segment.

Head capsule (Plate LXI, 312) massive, elongate, as in this division.

Labrum broadlytransverse, lateral parts a little enlarged and projecting anteriorly into blunt lobes, with along seta near inner margin; median region of labrum with two widely separated setae, justlaterad of each of which is a small papilla.

band of small spines.

Epipharynx roughened into a narrow transverse

Mentum completely .divided, each half continuous with ventral plate

The Crane-Flies of New York— Part901

IIof same side; anterior margin of each half with three slender, flattened teeth, the middle oneof which is slightly the shortest.

Hypopharynx conspicuous, labriform; anterior margin witha deep notch to form distinct lobes at lateral angles, and with several rows of small chitinizedtubercles.

Antenna(Plate

LXI, 313)

small;

basal segment elongated, slightly

curved, acircular auditory plate near base, at tip with numerous papillae, two of which are very long,nearly as long as segment itself; in addition to these, three or four tiny cylindrical papillae.

Mandible powerful; ventral cutting edge with a row of about four teeth which are successively enlarged from tip to base; basal tooth very broad and Cat, with outer margin truncate

or slightly concav(^; teeth on dorsal cutting edge indistinct; a pencil of moderately delicate

setae on scrobal region of mandible,

and another at prosthecal region.

Maxilla elongate;outer lobe larger than inner lobe, chitinized, apex with a very flat circular palpus (Plate

LXI, 314) which is provided with a few disklike papillae around margin and a few scatteredsensory papillae over pale apex; inner lobe with a long, powerful seta on ventral face andsmaller setae near tip.

Nepionotype.

— Ithaca, New York, June

1,

1917.

P araty pes. ^LsLTVue from type locality.
Rhaphidolabina Alexander (Gr. diminutive of Rhaphidolabi^

Genii.s

DIG

R'laphiMabina Alex.

Proc. Acad. Nat. Sci. Pliila., p. 540-541.

— Body covered with an abundant, appressed pubescence and tufts of erect hairs

Larva.-

which are more numerous on anterior end of body.
surfaces of abdominal segments 4 to

7.

Creeping-welts on dorsal and ventral

Spiracular lobes two, moderately elongated, eachSpiracles large.
Anal gills four, long and diaphanous. Head
Mandible powerful. Antenna short, with two elongate papillae.
Hypopharynx labriform, anterior margin concave. Mentum completely divided, each halfwith three large teeth and a much smaller lateral tooth.

with about six hairs at tip.

capsule long and massive.

— Labrum truncate. Pronotal breathing horns short-cylindrical or slightly flattened.

Pupa.-

Abdominal segments with circular areas of spicules on pleurites.
a broad transverse band of tiny spines.

Intermediate tergites with~

The genus Rhaphidolabina includes only R. flaveola, a curious pallid
fly of the northeastern United States, serving as a connecting link between

Tricyphona on the one hand and the Dicranotae on the other. The
The immature
stages are spent in rich organic earth in the same situations as are frequented by the adult flies.
adult flies are common on rank vegetation in cold woods.

Rhaphidolabina flaveola (O.
1869

The

S.)

Rhaphidolabis flaveola 0. S.

Hon. Dipt. N. Amer., part 4, p. 288.

Maine
At Orono, Maine, larvae were numerous in the richwriter has found the larvae of Rhaphidolabina flaveola inand in New York.

Charles Paul Alexander902
organic

mud of the Staiulpipo Woods.

One large larva was placed ina watch crystal with a fully grown larva of Adelphomyia cayuga (supposiIt at once seized th(; latter in its mandibles at about the thirdtion).
abdominal segment, and carried it helplessly all around the dish,occasionally shaking it, quite as a terrier does a rat.
Larva.

— Length, 9.S mm.
Diameter, 0.8 mm.

Color, brown to orange-yellow on anterior segments of body, becoming darker on abdominal

segments due to the increase of pubescence.
Body covered with an abundant, appressed pubescence. Form terete; body moderately
Abdominal segments 2 to 8 divided
elongated, gradually narrowed toward both ends.
into a narrow basal ring and a much broader posterior ring; segments 4 to 7 with conspicuous
dorsaland ventral welts, which are larger and more conspicuous on posterior segments;
and bisected by a deep longitudinal medianthese welts occupying basal ring of segments,

impression, their surface covered with microscopic points.

length with a transverse row of

stiff,

Thoracic segments before mid-dark brown hairs grouped in tufts or pencils; thesehairs occurring on abdominal segments also, but less prominent here and occupying posterior

region of segments; pencils

more numerous on ventral and pleural regions, much scantier

on dorsal surface except on pronotum.

On sides of last abdominal segment, between spiracles

and spiracular lobes, several long setae arranged in a transverse row, lacking on mid-dorsal
and ventral regions. Spiracular disk (Plate LXII, 321) reduced to two ventral lobes, as inthis division, these lobes slightly united basally, not very elongate, blunt at their tips,

bear about six dark setae.

which

The two spiracles large and conspicuous, entirely exposed on

dorsum of last segment, separated by a distance a little less than diameter of one; middle
Anal gills four, long and slender, nearlypiece of spiracles large, black; ring brownish yellow.
hyaline, divided into lobes by shght constrictions; posterior pair of lobes a little longer thananterior pair.

Head capsule elongate, flattened, massive, as in this tribe. Labrum conspicuous, exceedmentum and hypopharynx, the anterior margin fringed with long hairs. Mentum

ing(Plate LXII, 316) completely divided, each half with three subequal prominent teeth andan additional much smaller lateral tooth; middle tooth of each side a little broader. Hypopharynx (Plate LXII, 317) labriform, anterior margin deeply concave, roughened. Antenna(Plate LXII, 3L8) short, the segment short-cylindrical, a little enlarged toward truncated
apex; at tip several papillae, of which two are exceedingly elongate, very slender, about twice
length of basal segment.

Mandible (Plate LXII, 319) powerful, ending in an acute point;

ventral cutting edge with about four teeth, the basal one of which is the largest, subtruncate,

the next outer tooth a large, flattened, acute blade, two or three smaller flattened teeth justbefore tip; dorsal cutting edge with about two small teeth.

Maxilla(Plate

LXII, 3-0)

short and very stout, the outer lobe much longer than the slightly smaller inner lobe.

— (The following notes are taken from the cast pupal skins of the bred specimens.

Pupa.'

Labrum truncate.

Labial lobes ovate.

beyond base, tips blunt (Plate LXII, 322).
ally

to tip.

Pronotalbreathing

Maxillary palpi short and stout, broadest jupt

Antenna moderate in length, tapering gradu

horn (Plate LXII, 323 and 324) short, cylindrical or

The Crane-Flies of New York— Part II

circular breathing pores along margin
slightly fiattoned, with a row ofWing sheaths short. Leg sheaths much longer.
slightly enlarged.

903

of distal end, whichisabundant,
circular areas which are armed with
Pleural region of abdominal segments withspicul<-s
seven
to
five
from
being
there
spicules,ofrows
short, straight or slightly arcuated,scalehke tubercles,
magnification these appearing as flattened,ineach row; under highsegments

dorsum of intermediate abdominal
fringed with the spicules described above;
sharply curved spine^;is rather densely set with tiny,
which
band
transverse
with a broadlying posterior to
bands
dorsalweaker;muchor
on posterior segments these rows lacking
with dorsal plate rather smal each
325)
LXII,
(Platecauda
Maleareas.
level of laterala small seta;
outer lobe minutely tuberculate and withhalf indistinctly bilobed at tip;either side with a small tubercle bearingon
tergiteeighth
tips;
at
blunt
large,
ventral lobes,

two setae.
Nrmonotvve— Ithaca, New York, May

14, 1917.

,.

at,„ii14,

ivSC,^ Ithaca, May 2G, 1917. Cast pupal skin; larva placed m rearing May,•

Pupae, taken as larvae May 27, 1917, pLaeed
No. 79 - 1917.

in''p;™S;r^Saef'wiTMype larva.
rearing, emerged June 7, 1917.

Genus Tricyphona Zetterstedt
1837
1856
185G
I860
1881

Larva

Triaiphona Zett.

Amahins Hal.

(Gr.

three

+ hend)Isis von Oken, p. 65.

Ins. Brit., Dipt., vol. 3, p. 15.

l«.i.
Bophrosia Rond. Dipt. Ital. Prodr., vol. 1, p.
391.
Crunobia Kol. Wien. Ent. Monatschr., vol 4, p.
191.
Nasiterna Wall. Ent. Tidskr., vol. 2, p. 179,

basal annul!
- Body moderately elongate, with ventral transverse creeping-welts on
Ana g.Usof abdominal segments 4 to 7.

Spiracular lobes two, moderately elongate

divided into two to four lobes by from

one to three constrictions.

four,

Head capsule massive

in an acute point and with about
Eyespots distinct. Mandible powerful, ending
lobes, the slender inner lobe
Maxilla stout, consisting of two distinct
four lateral teeth.
basal segment cylindricalsmall;
Antennalobe.
outerclosely approximated to the larger

elongate.

at tip, withtwo exceedingly long, hyaline papillae

T rnma^^data).
Mentum completely divided,(possibly lacking inroughened.
Hypopharynx labriform. anterior margin concave,large teeth.
anterior margin of each half with three,,
angles rounded.
-Pronotal breathing horns short and stout, blunt,

Pupa

on tergites andsegments with transverse bands of spicules

sternites,

au^
Abdominal•iand circular areas

on pleurites.

including more thanTricyphona is the largest genus of the Pediciini,
are found in thespeciesAlmost all of thesefifty described species.
eastern Umted
theIn
Antipodal.
Holarctic region, but two or three are
and appardistributed
widelymostStates Tricvvhona incmistans is the(as I.
otherscertamandcommonest species. This species
ently theinhabitants, but other species
paludicola Alex.) are characteristic swamp

Charles Paul Alexander904

occur along running streams (T. vcrnalis [O. S.]) or near cliffs (T auripennis.

[O. S.]).

The immature stages of the known species are usually spent in moist
In Europe, T. immaculata (Meig.) was reared by Beling (1879:47)
from larvae taken beneath decaying vegetable mold in the bed of adried-up woodland stream. Other specimens were found in old horse
manure in beech woods. De Meijere (1916:195-196) found the same
species in decaying leaves and other vegetable matter in wet spots near
the banks of streams.
T. schineri (Kol.) was found on August 19 in awet spot in beech woods, where the larvae were associated with pupaeof Pedicia rivosa in' damp earth beneath 'debris, adults emerging on
September 6 and 12 (Beling, 1879:47). The only American species
whose immature stages have been found is T. inconstans, described
earth.

herein.

Tricyphona immaculata (Meig.), the genotype, as described byDefrom T. inconstans as herein
The antenna bears a very short terminal

Meijere in the paper cited above,
described in the following points:

differs

two very long ones in the local species. The small
on the mentum is not mentioned nor figured as appearing
The anal gills are short and with but a single conT. immaculata.

{)apilla instead of thelateral toothin

The pupa hasstriction.

the pronotal breathing horns small, kidney-

shaped, the outer margin rounded.

The fore legs are a little shorter thanthe middle legs, and these in turn are somewhat shorter than the posteriorlegs.

The skin of the abdomen is very delicate, with crossrows of verysmall spicules, arranged in numerous irregular transverse rows, at theposterior margins of the segments,

about seven such crossrows being

present.

Tricyphona inconstans (O.

S.)

Amalopis inconslans 0. S.

1859

Proc. Acad. Nat. Sci. Phila., p. 247.

Tricyphona inconstans is an abundant species thruout eastern North
It has been recorded also from Europe, but the latter records
are almost certainly erroneous.
The writer found larvae of this species
America.
in richmud at Larch Meadows, near Ithaca, New York, on May 15,

1917, in association with larvae of Rhamphidia mainensis, Pseudohmnophila

May 27, 1917 (No.

luteipennis,

and P. inornata.

52-1917).

An account of the association in which these larvae occurred

Adults emerged on

The Crane-Flies of New York
is

givon

in

connection— PartII

905

with the discussion of Rhamphidia mainensis(page 831).
Larva.

— Length, 17-17.5 mm.
Diameter, 1.3 mm.

Color pale yellowish white; anterior parts of body sometimes a more saturated yellow.

Form moderately

stout,

body alittleof body almostand scanty scattered hairs.

narrowed toward «nds; surface

glabrous, the vestiture being a microscopic pale pubescence

Ventral surface of abdominal segments 4 to 7 with a prominent transverse welt, whichishollowed out medially so as to appear as paired prolegs, these being unarmed with hooks
or points.

Spiracular lobes short, divergent, tapering gradually to the blunt tips, which areprovided with seven or eight setae.
tion, small,Spiracles (Plate LXIII, 330) on a transverse oval eleva-

rounded, separated by a distance greater than diameter of one; middle piece ofspiracles large,

black, rings narrow.

Anal

gills

four,

very long and slender, constricted

into about four lobes which are successively narrowed from the base outward, the terminal
division very slender.

Head capsule of the usual elongate, massive type of this tribe.

Labrum large, projecting

beyond hypopharynx, anterior margin with long hairs. Mentum (Plate LXIII, 326) large,
completely divided, each half with three subequal narrow teeth and an additional reduced
lateral tooth; on proximal margin of each half of mentum, near base of innermost tooth,a few tiny notches dovetailing into those of opposite half. Hypopharynx (Plate LXIII, 327)
projecting far beyond mentum, labriform; anterior margin deeply concave and provided withsmall rounded papillae; lateral angles smooth, rounded, subchitinized. Antenna (Plate
LXIII, 328) small; basal segment cylindrical, bearing at its tip two elongate papillae which
are about one-half longer than basal segment alone; besides these an oval papilla. Mandible(Plate LXIII, 329) of the usual pediciine type, powerful, the apex running out in. a long,
curved point; ventral cutting edge very flat, cut into about five teeth, the two basal of which
are very large; the most basal of these teeth squarely truncated, the left mandible with anadditional small tooth on lower inner angle of this blade; the next outer tooth rather acute;outermost teeth small and flattened; dorsal cutting edge, as usual with this division, withtwo very small teeth located far out on apical point; a tuft of a few long setae on back ofmandible near heel. Maxilla (Plate LXIII, 329) powerful, as in this group of genera,
but not exerted from the prothoracic orifice when head is retracted; outer lobe very
stout, feebly chitinized basally, hyaline at apex,

which bears the flattened, diskhke palpus;

inner lobe slenderer, shorter, with a few setae and small papillae.

Pupa.

— (The following notes are taken from the cast skins of the reared specimens.)

Labrum with apex broadly

triangular.

bluntly rounded (Plate LXIII, 331).

Labial lobes large, ovate, tips narrowed andPronotal breathing horns (Plate LXIII, 332 and .333)

very short, stout, roughly cylindrical, the apices truncated; in lateral outline, pentagonal,
with a row of breathing pores around margin of truncate apex.
versely roughenedbyshort,

Thoracic dorsum transirregular grooves.

Abdominal sternites with broad transverse bands of spicules on segments 5 to 7, the last
bands on tergites 4 and 5, these bands subequalin size; pleural region with a large, roughly circular area of short rows of microscopic spicules,of these three bands the weakest; similar

Charles Paul Alexander906

these areas near base of segments.

Female cauda (Plate LXIII, 334) with tergal valveselongate, tapering gradually to the rather blunt tips; sternal valves very small, blunt at tips;

at base, on dorsal side of eighth segment, two setiferous tubercles.

Nepiotwlypc— Larch Meadows, Ithaca, New York, May 15, 1917.
Neanotijpc.

— Ithaca, May

No. 52-1917.

27, 1917, a cast pupal skin.

Group Dicranotae

Genus Dicranota Zctterstcdt (Gr. a fork)
1838

Dicranota Zett.

Ins.

Lapponica, Dipt., p. 851, no. 164.

Dicranota is a small genus of crane-flies (about fifteen described species)

Four species arc found in eastern

occurring thruout the Holarctic region.

North AnKMica.

None of the American species have been reared.

In

Europe, the life history and morj)hology of Dicranota bimaculata (Schum.)has been discussed in detail by Miall (1893) and by Wesenberg-Lund(1915:342-343). Larvae were found by Miall in numbers in the muddy

banks of small streams and ponds, where they bury themselves in the
mud and gravel. They creep about with ease and rapidity between thesand and gi'avcl, and are able to swim well by a looping movement. Theirfood consists lai'gcly of small worms, Tubifex rivulorum Lam., whichabound in these haunts.

The pupal stage is passed in damp earth.

Thelarva is stated to be about 18 millimeters in length, but this seems to bea maximum figure.
fine,

The body is dirty white in color and is covered withappressed hairs.

Abdominal segments 3 to 7 bear paired retractile

pseudopods, which are circled with three rows of chitinizod hooks gradually
decreasing in size from the tips inward.
are distinctly segmented.

The head

The anal gills, four in number,capsuleiselongate and massive,

The mentum is completely divided, its anterior margin
having the usual six teeth. The mandible is of the usual pediciine t3'pe,as in the tribe.

with acute teeth on the ventral cutting edge antl a brush of hairs near
(Plate LXIV.)
The pupa is small, only about 10 millimeters in length, and has the
pronotal breathing horns expanded and flattened at the tips. The dorsal
surface of the abdomen is provided with roughened plates armed witli

the prosthecal region.

rather strong and dense spines, there being one such plate on the third
segment, two on the fourth to sixth segments, and one on the seventhsegment.

The Crane-Flies of New York— Part II

907

Genus Rhaphidolabis Osten Sackon {Gr. needle + forceps)
1869
1911

Rhnphidolahh 0. S. Mon. Dint. N. Amer., part 4, p. 2Sl:-2'7.
Claduroides Brun.
Rec. Indian Mus., vol. (J, p. 288.

Rhaphidolabis is a small genus, including about fifteen described species
The larvae strongly resemble

ranging thruout the North Temperate Zone.

those of Dicranota, and the two genera are undoubtedly very closely
related.

Rhaphidolabis tenuipes O.
1869

S.

Mon. Dipt. N. Amer., part 4, p. 287.

Rhaphidolabis tenuipes O. S.

Needham (1908 a: 212-214) found adult flies of the species Rhaphidolabis
numbers in tent traps set over Beaver Meadow Brook

tenuipes in greatin the Adirondacks in July and August,tainly belong here were found

1907.

Larvae that almost cer-among the rounded stones in the creek

bottom.
Larva.

— Length excluding caudal lobes, 8-9 mm.
Length of caudal lobes, 1 mm.

Abdominal segments 3 to 7 with prominent fleshy prolongations on ventral surface, thesebeing retractile, unpaired, and widely separated on mid-ventral line, and bearing at tip acirclet of outcurved booklets,with series of smaller booklets beyond.

the two ventral lobes long and obtuse at tips.

Spiracular disk with

(Further details are given by Needham.)

Rhaphidolabis cayuga Alex, (supposition)
1916

Rhaphidolabis cayuga Alex.

Proc. Acad. Nat. Sci. Phila., p. 543-544.

Larvae which the writer refers to this species were found in Needham's
New York, on March 30, 1916. Later in the season, in
April and May, adult fli(;s of the species arc very numerous in this glen,
and these account for the specific reference.
Glen, Ithaca,— Agreeing very closely with descriptions of both Dicranota and Rhaphidolabis

Larva.-

tenuipes.

Spiracles large, lying in a distinct longitudinal groove, by the closing of which they

Antenna long and slender, tapering to apex, which
numerous short papillae. Mandible with third lateral tooth of ventral cutting
row very long and acute, much exceeding second tooth; basal tooth shaped like a prun-are capable of being entirely hidden.

bear;

ing knife, with the cutting edge sinuate; at prosthecal region of mandible, a small tuft ofabout seven or eight long setae.
apical quarter.

IMa.xilla

having the two lobes firmly united except on

908Charles Paul Alexander
Tribe Eriopterini

The tribe Eriopterini comprises a vast assemblage of usually small cranewhose geographical range is coextensive with that of the family.
The lai'vae, so far as known, are herbivorous. They show a remarkableuniformity in the structure of the head capsule. In the generalizcnl
members such as Molophilus and some Erioptcra, the ventral bars of theh(^ad capsule are toothcnl at their anterior ends and form mental plates
which are apparently homologous with those of the Pediciini. In Chionea
an appanuitly similar condition exists, to judge from Brauer's figures.

flies

In the majority of species, however, the ventral bars of the capsule are

not enlarged nor toothed anteriorly and do not function as the mental
The hypopharynx is preserved as a hemispherical cushion whichplates.
isdensely provided with setae.

The mandibles are small and are blunt
numberat their tips; the teeth of the cutting edge are usually three inand blunt, but in some species (Ormosia, Gonomyia) they are longer
and more prominent. A distinct prosthecal lobe or hook is usuallydeveloped, and near the base of the mandible is a slitlike opening bearing
a fringe of long, yellow setae. The maxillae are rather large, hairy lobes.
The labrum and epipharynx is long and narrow, and densely hairy. The

antennae are remarkably uniform thruout the group, consisting of a stout
cylindrical basal segment with a slightly smaller apical papilla of an
In the Elephantomyaria the condition is somewhat
elongate-oval shape.
similar, the mandibles being very small, and the esophageal region being
conspicuously grooved with parallel lines and ridges. The spiracular
disk is obliquely truncated and indistinctly lobed in Chionea and Teucholabis; surrounded by four lobes in Elephantomyia; squarely truncatedand surrounded by four (in Rhabdomastix) or usually five lobes. In
the undetermined Eriopterine No. 1, the five lobes are spatulate flattened
blades with the margins hooked. Anal gills are usually present andvariously developed in the different genera.
The pupa is rather slender,
in the Elephantomyaria with the rostral sheath very long and the palpi
strongly recurved. The head usually bears a small setiferous crest whichis rarely lacking.
The pronotal breathing horns vabiy considerably inform and relative size, being flattened into fans in some Gonomyia, small
and trumpet-shaped in other Gonomyia and in Gnophomyia, elongate
in most species.

The mesonotum is usually armed at the crest with spines,
The leg sheaths are very short in Gno-

hooks, or setiferous tubercles.

The Crane-Flies of New York— Part II

909

phomyia but are longer in other genera, and the middle tarsi are usuallyshorter than the other legs.

The abdominal segments are subdivided

two rings, the posterior ring with a transverse row of spines andinto

The lateral spiracles are small, protuberant,of theand, in some cases at least, apparently functional. The dorsum

setae before the margin.

eighth abdominal segment is provided with four or five lobes,
in some cases (as in some Ormosia) spinous at the tips.

The writer has subdivided

which arethe tribe Eriopterini into two divisions:

Toxorhina
the Elephantomyaria, with Elephantomyia and presumably
Nearcticothertheand Ceratocheilus, and the Eriopteraria, including
requirewill
relatives
genera as known. It is probable that Cladura and its
known.
a division when their immature stages are better
The genera of the tribe Eriopterini may be separated by the following
keys
Larvae2

1.

Spiracular disk surrounded by four lobes
Spiracular disk not as aoove
j
iV
i-'
lives
Ventral lobes with a single powerful seta; coloration a saturated golden yellow;
S. (p. 95^)
ElephantomyiaO.
under bark
Ventral lobes fringed with short setae; coloration pale yellow; lives in earth.
Rhabdomastix Sk. (p. 942)
'

,'

'

'

'

"

'

i

2.

without lobes.

Spiracular disk obliquely truncated, appearing indistinctly trilobed or
Spiracular disk squarely truncated, surrounded by five lobes
lives in earth.
4. Mandibles with eight 'teeth; menial plates with seven teeth;
3.

.

.4

5

Chionea Dalman (p. 9o())

Mandibles with less than eight teeth; mental plates not toothed; lives under bark

Teucholabis O. S. (p. 945)

5.

Spiracular disk ending in five flattened black plates which are finely toothedmargins

Spiracular disk not as above'
Ventral plates of head capsule expanded and toothed at anterior ends
Ventral plates not toothed as above
heavily
Ventral plates with four teeth; coloration yellow; spiracular disk large, very
Molophilus Curt. (p. 911)
marked with black
reducedVentral plates with five to eight teeth; coloration green; spiracular disk very
Erioptera {chlorophylla 0. S ) (p. 91•

6.

along their

Genus incertus, Eriopterme No. 1 (p. 95b)
•

C"

7

8.

almost unmarked
Marks of all the lobes solidly black

".

'-'t

Tnmtcra O. S. (p. 9SJ}

Some Ormosia Rond. (p. 9J2)

9.

Some of the marks more or less split by a pale Hne
The three dorsal lobes solidly dark, the ventral pair split by a pale line
All paired lobes split by a pale line

10.

Coloration saturated yellow; anal gills bluntly rounded; lives under bark.

•

^
lU
^ "^

Gnophomyia O. S. (p. 9c5i)
Coloration pale yellow; anal gills elongate; lives in earth.
Helobia St. Farg. et Serv. (p. 928)
surrounding sP'ra(;Jes
11. Lobes surrounding disk very stout, blunt; marks of lateral lobes
Gonomyia Meig., subgenus Leiponeura bkuse (p. 9.i9)and often suffusing disk
12
Lobes surrounding disk longer; marks of lobes not so extensive

910Charles Paul Alexander

12. All13

the lobes with two lines
Dorsal lobe solidly dark
No dirk m..pL-o „n A\^\. K 1

Da^ktXt^tZn
^"*'

" ^'

•'

tSr

'

;
A
i
oome
Ormosia Rond. (p. 922),''^'""'"'^^"^^^"^^ "P^*" "'^ ^'^'^sESnli^r H-t
T^^ T^'"''
Spiracular
disk large;
four or six spots on disk.

^--- «-d- (P- 922)

Erioptera (megophthalma Alex)
(p 915")
Helobia St Farg et Serv. (p. 928)

•

Pupae
1.

2
2.

Rostral sheath very elongated; antennal
sheaths lying across face of eye-\^>-gi^ted; antennal sheaths lying behindl;^'"'""''" "^^ '^ ^'^ ''%
lT'?!
tr*'
Leg
sheaths
very short, barely exceeding wings;
crestof mesonotum smooth or nearly so

Mesonotum and tarsal arrangement not as above
'•

5.
b.

""^SS^^^C'ui;:^^^'* ''''''''''^^''^- ^^^^-^j"""""^^^ ^'"^- ^P" ^^^Mesonotum and breathing horns' not asabove
Mesonotum at crest with four sharp spines or two
phtes produced into spines
6
Mesonotum at crest without distinct prominent

Alesonotum at crest with four spines

Mesonotum at crest with a plate on either side

8.

^^
'

'-^^ with numerous-se^e; b^athin^^

spLaeso^'split at "tip" into three or' four teeth'

Abdominal pkurites with 'a ,ra„sver« .„bte™i;u>lrow o, ,roJ^e°^t'^''^J;We±l"lstout setae; dorsum of segment 8 witha pentagon of five lobes.

Enopiera RIeig, (p. 914)

The most important literature on the tribe
Mdopha,^bifilatu,

J/«(

pAito ob^rmfS™?!".2?!"°&*ttT:So.:.*.-::::;;;:

Keilin. 1613:4.

GeneralBeftaflSTQ 56

G"-"'^^™' "-p^SzSLTr*-'^-.:;::::::; Jfe™r---

Ormosia nodulosa
Ortnosia nodulosa

Ormosia vanaurmosmvaria

Eriopterini is as follows-

Larva, general

GeneralGeneral--"^.c*.
Larva, pupa:

(Hyp«lermal

De iTeiierlfigM:?^

b*., ,886:192- ^' ^^-'^'
BeJing, 1886:202.
Beling, 1886:202.

Cameron, 1917:
i^ameron,
1917:65.
65.
De Meijere, 1916: 201-204

The Crane-Flies of New York
Helobii hybrida
Helobia hybrida
Helobia hybrida
Trimicra pilipes
Gnophomyia rufa
Gnophomyia tripudians
Gnophomyia tripudians

Gnophomyia tripudians— Part911

II

Larva, pupa

Beling, 1879:50-51.

Larva
Larva, pupa
Larva, general
Larva, pupa
GeneralGeneralHart, 1898 [1895]; 199-200.
Malloch, 19 15-17 b: 229-230.
Gerbig, 1913: 161-163.
Hudson, 1920: 32-33.
Gamkrelidze, 1913, a and b.
Keilin, 1913:3.
(Hypodermal

Larva

glands.)
Keilin, 1913:4.

(Hypodermal

glands.)Gnophomyia tripudians
Gnophomyia tristissima
Gonomyin lenella

GeneralLarva, pupa, general.
GeneralLarva, pupa
Larva, pupa, general.
GeneralLarva, pupa, general.
GeneralLarva, general

Rhabdoma.ttix schistacea
Trentepnhlia hromeliadicola

TrentepohUa leucoxena
Trentepohlia pennipes
Teucholabis complexa
Chionea araneoides

.

.

.

Edwards, 1919 b.
Malloch, 1915 17b: 230-231.
Beling, 1879: 56.
Beling, 1886: 195.
Picado, 19 13:.3.56-3.57.

Alexander, 1915 b.
De Meijere, 1911 50-51.
Johnson, 1900.
Brauer, Egger, and Frauenfeld,
:

1854.

Subtribe Eriopteraria

Genus Molophilus Curtis (derivation obscure)
1833

Molophilus Curt.

Brit. Ent., p. 444.

— Form long and slender.

Larva.'

Spiracular disk squarely truncated and surrounded by

five subequa! lobes; paired lobes of disk heavily lined with black; a black mark running proxi-

mad from spiracles; dorsal lobe with an oval black area.
ventralrods

flattened;

anterior

Labrum narrow, densely hairy.

Head capsule long and narrow;to form mental plates.
Mandible blunt at apex; ventral cutting edge with about

ends expanded,

four teeth; a single subapical dorsal tooth.

four-toothed,Maxilla large and blunt.

Antenna large; apical

papilla elongate-oval, sculptured.

Pupa.— Cephalic crest setiferous.
curved.

Pronotal breathing horns long and slender, sinuously

Mesonotum declivitous, at crest with four powerful teeth.

at about midlength of second abdominal segment.

Wing sheaths ending

Leg sheaths ending at about midlength

abdominal segment; tarsi of middle legs the shortest.
on segments 2 to 7; dorsum of segment 8 with four blunt lobes.
of fourth

Abdomen with spiracles

Molophilus is a well-defined genus (including more than fifty species)of small and usually dull-colored flies, which are found practically thruout

the world.

The adult flies frequent shaded situations and may be swept fromrank vegetation in such places. The swarming habits of this group are
discussed later under the specific accounts.

The immature stages of the various species are spent in wet earth.
In Europe, Molophilus ohscurus (Meig.) (Beling, 1879:56) and M. ochraceus(Meig'.) (Beling,

1886:193-194) have been found in wet earth near running

912Charles Paul Alexander

water in shaded woods.

more than ten days.

The pupal duration of tiie latter species is not
Keilin (1913:4) notes the presence of hypodermal

M. bifilatus Verr., but does not mention the larval
Of the American species, M. hirtipennis has been reared
fromglands in larvae of
habitat.

similar situations.

Molophilus hirtipenms (0. S.)
1859

Erioplera hirtipennis 0. S.

Proc. Acad. Nat. Sci. Phila., p. 228.

The little dark-colored cranc-flie- of the species Molophilus
hirtipennis
are common on vegetation in shaded woods in spring andearly summer,or they may be found in small cUuicing swarms in similar
situations.
The larvae are exceedingly al)undant in wc^t organic mud or incool, richwoods in the neighborhood of streams or springs.

The writer has reared

the species very frequently from specimens found in
Needham's Glenand on Bool's hillside, Ithaca, Now York, in April and May,
1914 to1917. The pupal period is prol)al)ly al)out a w.xk, l^ut in all
the rearingsof the writer this could not be ascertained closer than
Larva.

ten days.

— Length, &-10 mm.

Diameter, 0.4-0.5 mm.
Coloration light yellow.

Form long and narrow; body terete, noticeably constricted before
spiracuL^r disk (Plate
XLV, 344). Integument covered with a delicate appre.sed pubescence anda few transverserows of very short, erect setae.

Spiracular disk (Plate

LXV, 351) squarely truncated, sur-rounded by five subequal lobes; ventral lobes on inner facewith two heavy black parallelby a capillary yellow line; lateral lobes with a similar double line
runninginward far beyond spiracle; these double lines not connected at
distal end; dorsal
hues, separated

lobe witha single oval black mark which is less intense outwardly; a black

mark beginning at spiracle

running proximad toward center of disk; lobes with a few shorthairs at tips.
(There is avariation in the degree of intensity, but the general
pattern is as described above.)
Anal gills four, short and blunt.
littleHead capsule (Plate LXV, 345) long, narrow, consisting of six chitinizedrods, the ventral
rods broad and flat, at the anterior end expanded to form
the mental plates (Plate' LXV, 346),
each rod contributing four teeth, of which the outermost is bluntly
rounded, the middle pairthe largest and subequal.
Dorsal bars two on either side, one slender, attheir anterior

ends articulating with a transverse chitinized rod; the various
bars connected by a thinmembranous tissue. Labrum and epipharynx elongate, narrow, occupying the space
betweenmandible and antenna on either side, the ventral face with abundant long
hairs, on sides
margined with numerous long, incurved, flattened setae. Mental
plates as described above;
behind them the hypopharynx (Plate LXV, 347), consisting ofa semicircular cushion provided
with dense, short setae. Antennae (Plate LXV, 34S) rather closely
approximated on dorsum,
promment, each 1-segmented but bearing a long apical papilla; basal
segment moderately
elongated, cylindrical, the apex obliquely truncated, the papillahyaline, gradually narrowed

The Crane-Flies of New York— Part II

913

toward tip, shaped somewhat Hke an ear of corn, the surface delicately sculptured; besides
Mandible (Plate LXV, 349) slender,
this papilla, two or three much smaller cylindrical ones.
ending in a blunt rounded lobe; ventral cutting edge with four blunt teeth, the second fromthe base very tiny and connected with the third from the base (in some specimens the teeth
are very blunt and rounded, so that the cutting edge appears crenulated or wavy); a single

tooth on dorsal cutting edge, immediately behind apex; prosthecal region of mandible witha longitudinal slitlike opening filled with a dense row of long setae. Maxilla (Plate LXV,350) large and blunt, the outer lobe pale, roughly triangular, covered with numerous shorthairs, and with a few small sensory papillae near apex, surrounding palpus; inner lobe alittle shorter, densely clothed and fringed with abundant long yellow hairs.

Pupa.

— Length, 6.2-7.3 mm.
Width, d.-s., 0.7-0.8 mm.
Depth, d.-v., 0.8-1 mm.

Head, thorax, and appendages brown, when fully matured almost black; breathing horns
yellowish; abdomen pale yellowish white.

Cephalic crest of moderate size, consisting of two conical lobes directed forward and bearing

a stout seta on anterior face; immediately in front of these, a smaller transverse crest lying
between antennal bases. Front rather broad. Labrum triangular; lobes of labium triangular,
divergent; maxillary palpi elongate, moderately stout, narrowed at tips and extending beyond
joints

of

fore

legs.

Thorax prominent, carinate medially before

horns long and slender, sinuously curved, apices directed forward.
(Platedeclivity.

BreathingDeclivity of mesonotum

LXVL 352) precipitous, at the crest armed with four powerful teeth, median pair thelargest, somewhat divergent, separated by median line; a few tiny setae on mesonotum, includ-ing four in a transverse row at about the level of wing root.

Wing sheaths ending opposite

apex of second abdominal segment.

(In fully matured pupae the characteristic venation ofthis genus shows on the wing pads.)

Leg sheaths rather short, ending about opposite mid-length of fourth abdominal segment; hind legs slightly the longest, fore legs a little shorter,

middle legs conspicuously shorter, ending about opposite apex of fourth tarsal segment offore legs.

Abdominal segments with two very narrow basal rings and a much broader posterior
Abdominal spiracles distinct on segments 2 to 7. Petae as follows: on pleura, one
immediately caudad of spiracle, a second caudad and somewhat dorsad of spiracle opposite
Tergum with two setae on sides of posterior
posterior ring, another opposite anterior ring.
ring.
Segment 8 with four blunt lobes on dorsum. Male cauda (Plate LXVI, 353 and 354)with ventral lobe large, bluntly rounded at tips; dorsal lobes ending in two acute, chitinized
points which are widely separated and directed dorsad and slightly caudad, on outer facewith a small seta; near base of cauda on dorsum, surface tumid and bearing a small seta on
either side.
Female cauda (Plate LXVL 355 and 356) with sternal acidothecae much shorter
than the very long tergal valves, these latter, just before apex, with an acute spine which is(When the pupa is nearly ready to transform
directed dorsad and with a seta on side.
to the adult, the long, coarse bristles covering the body of the adult, and the chitinized
genitalia, show thru the pupal integument.)
ring.

Nepionotype.— Ithaca, New York, May 11, 1917. No. 33-1917.
Ithaca, June 1, 1917.
About one hundred larvae and pupae from type locality.

——

Neanotypp.
Paratypes.

8

ICharles Paul Alexander

914

Molophilus ursinus (O.
1850

S.)

Erioptera ursina 0. S.

Proc. Acad. Nat. Sci. Phila., p. 228.

Molophilus ursinus is probably the smallest crane-fly in North America.

The following notes on the swarming were made along Power House
Creek, Gloversvillc, New York, on June 27, 1915:
This species occurred in small dancing swarms over the little lateral streamlets that poured
When not swarming, they rested flat on the upper surfaces of leaves, their wings folded over the abdomen.
Crane-flies associated with this species today included the following:
Biltacomorphellajonesi, Limnophila toxoneura, L. areolatn, L. (Lasiomoftiix) tenuicornis, Liogma nodicornis,
Dolichopeza nmericana, Tipula collaris, T. vohilis, T. iroquois, T. hermannin, T. macrolahis,
T. submaculata, T. fuliginosa, Nephroloma macrocera, N. tenuis, Longurio tesiaceus, and others.

down the steep slope, some of the swarms influding scores of individuals.

Genus Erioptera Meigen (Gr. wool + witig)
1800
1803
1818
1854
1856
1856
1856
1861
1863

1864
Larva.

Polymeda Meig.

Nouv. Class. Mouch., p. 14 (nomen nudum).

Erioptera Meig.

Polyraphia

Illiger's Mag., vol. 2, p. 262.
Meig. Syst. Beschr. Zweifl. Ins., vol. 1, p. 88.

Ann. Soc. Ent. France, p. 474.
Chemalida Rond. Dipt. Ital. Prodr., vol. 1, p. 180.
Limnaea Rond. Dipt. Ital. Prodr., vol. 1, p. 181.
Ilisia Rond.
Dipt. Ital. Prodr., vol. 1, p. 182.
Limnoica Rond. Dipt. Ital. Prodr., Corrigenda, vol. 4, p. 11.
Octavia Bigot.

,

Wien. Ent. Monatschr., vol. 7, p. 221.
Atti dell' Institut Veneto, ser. 3, vol. 10, p. 42.

Trichosticha Schin.

Platytoma Lioy.

— Form elongate,insome cases very slender, terete.

be reduced, in some species (as E. chlorophylla) very small.

Spiracular disk tending toAnal gills blunt.

Head capsuleand two ventral; ventral bars in some species(as E. chlorophylla), at least, with five to eight teeth at their expanded anterior end, forming
the mental plates. Labrum narrow, densely hairy. Mandible small, bluntly toothed.
Maxilla blunt, hairy. Hypopharynx cushion-like.
Pupa.Cephahc crest consisting of blunt or acute (in E. chlorophylla and E. vesperiina)
slender, consisting of six bars, four dorsal—

lobes.

Pronotal breathing horns elongate, the length

many times the diameter, usuallystraight and slightly divergent {E. megophthalma, E. septemtrionis)or acutely pointed andMesonotal declivity along crest with very numerous
setiferous tubercles or (in the European E. lutea) with four prominent teeth.
Leg sheaths
moderately long, reaching fourth abdominal segment; middle tarsi conspicuously shorter,

directed forward {E. chlorophylla).

than the others.

Subapical armature of abdominal segments strong on stornites and tergites

but lacking on pleurites.

Spiracles distinct, tubular.

Lobes on dorsum of eighth abdominal

segment not forming a distinct pentagon.

Erioptera is a large genus of small flies, including more than one hundred
described species, chiefly from the temperate regions.
all species of whichin

wet earth.

In the case of

the immature stages are known, these stages are spent

In Europe, Erioptera (Acyphona) maculata (Beling, 1879.

The Crane-Flies of New York— Part II

915

was found under leaves in damp earth in woods. Pupae taken on
June 8, 1874, emerged as adults on the 13th. The species was foundE. (E.) flavescens (Beling,also in the sandy mud of a small brook bed.
1879:50) was found in sandy, damp earth, E. (E.) sordida (Beling;, 1879:
56, mention only) in wet, sandy earth along brooks, and E. (E.) lutea(Beling, 1886:192-193) in woods, in damp hollows, and along the margins
Edwards (1919 a) has recorded aof brooks, usually under leaf mold.
species of Erioptera as being associated with the larvae of the mosquito
49)

Taeniorhynchus

at

the

roots

of

water

grass

{Glyceria

fluitans)inEngland.
E. chlorophylla, E. vespertina, E. caloptera, and otherIn America,

species inhabit wet mud in open swamps; E. septemirionis and E. megophrich organic mud in cool, shady woods; E. arniata, E. nearand other species, the soil along the sandy banks of streams. E.
(Acyphona) graphica was considered by Hart (1898 [1895]: 197) to bethalma,knabi,

semiaquatic.

The writer has before him the larvae of only two species of the genus,
These are readily separated by

E. chlorophylla and E. megophthalma.
theircolor,

E.

chlorophylla

being

pale

green,

E. megophthalma pale

The pupae of the Nearctic species of thegenus may be separated by the following key:yellow and more elongate.

Breathing horns and cephalic crest strongly pointed and curved forward at their tips;
coloration light green
E. chlorophylla O. S. (p. 918)
Breathing horns and cephalic crest not as above; coloration not green2
2. Breathhig horns elongate-cylindrical to slightly flattened, almost straight but slightly
divergent, diameter subequ::! for entire length3
Breathing horns not as above, more or less curved, or else blunt at their tips
4
3. Cephalic crest sharply pointed; ^yes large
E. megophthalma Alex. (p. 915)
Cephalic crest not sharply pointed; eyes small
E. septemirionis O. S. (p. 920)
4. Breathing horns almost straight, enlarged distally, tips blunt.
E. sp. (near knabi Alex.) (p. 922)
Breathing horns curved, narrowed toward tips
5
5. Breathing horns widely separated at base, bent strongly proximad, contiguous medially;
lobes of cephalic crest acute, vertical; eyes large
E. vespertina O. S. (p. 910)
Breathing horns curved gently forward; cephalic crest with lobes directed laterad, divergent; eyes of moderate size
E. armata O. S. (p. 921)
1.

Erioptera megophthalma Alex.
1918

The
cool,

Erioptera megophthalma Alex.

adult

shaded,flies

of Erioptera

Can. Ent., vol. 50, p. 60-61.

megophthalma

are

not

and boggy woods during the monthsofuncommonearlyinspring.

Charles Paul Alexander

916

They may be swept from rich vegetation in company with suchchar-

acteristic species as Limnophila hrevifurca, L. fuscovaria, L. suhtenuicornis,

Adelphoniyia

Rhaphidolabina

minuta,flaveola,

Gonomyia

Molophilus

hirtipennis,

andE.
Tipula oropezoides.
The elongate larvae of this fly were very common in rich organic mud
taken from Bool's hillside, Ithaca, New York (as discussed under the
Erioptera venusta,

stigmatica,

Jlorens, G.

suhcinerea,

account of Bittacomorphella jonesi, page 780), where the}^ were associatedwith a crane-fly fauna characteristic of such situations. The larvae, inlife, are pale yellowish, with the food contents, of a chalky white color,showing thru the integument. The head capsule and the spiracular disk
are very small the inner face of each lobe of the latter is very narrowly
The species was reared many times during late May
lined with black.
;and early June, 1917, the length of the pupal existence indoors being sevenor eight days.
Larva.

— Length, 10.4-11.6 mm.
Diameter, 0.7-0.75 mm.

Coloration very pale yellow; contents of alimentary canal chalky white.

Form terete, elongated, body tapering gradually to the posterior end, just beyond gills
LXVII, 358) suddenly constricted; last segment elongate-cylindrical, tapering graduBody covered with a short, appressed pubesally to the very small spiracular disk.
(Plateand more erect, with a few elongate hairs
body at caudal margins of segments with short transverse

cence, on last segment this pubescence coarser

interspersed;

lateral parts oflines of small, erect setae;

segments.

Spiracular diska few other similar rows at about midlength of certain of the(Plate

LXVII, 357) very small, tending

to be eliminatedbyreduction; lobes short and blunt, dorso-median lobe the smallest; ventral lobes with two
short brown lines, not connected distally, the proximal line a little longer than the lateralline ofeach lobe; the pale space between these lines alaterallobes withtwo

similar divergent

lines,

little lessthe dorsal one

than diameter of one;

attaining

inner level of

two small, indistinct, brown lines; on disk between spiracles two
small round spots which do not touch spiracles; lobes fringed with short hairs near tip,and capable of close approximation so that disk is often entirely closed. Spiracles large,

spiracles; dorsal lobe withnearly circidar.

Head capsule small, very long and slender, greatly dissected, the three bars of either side
long and delicate; dorsal bars at their articulation joined with a short longitudinal bar nearwhose anterior end the antennae are inserted; ventral bars of capsule not conspicuously
expanded at their anterior end, and apparently not toothed as in other species of this gcnu.i
and in Molophilus. Labrura and epiph:.rynx long and narrow, lying between antenniil
bases; epipharyngeal region densely clothed with short setae at tip

brushes on ventral face.

and with two parallel

Mentum apparently not formed as in E. chlorophylla, a slightly

The Crane-Flies of New York— Part917

IIarched transverse chitinized bar. Hypopliarynx about as in Molophilus. Antennae rather
closely approximated, directed cephalad; basal segment moderately elongated, cylindrical;
apicalpapillarelatively

small,

Mandible very small, with blunt teeth;
row of teeth about three in number, often very blunt.

elongate-oval.

apical point short, blunt; ventral

Maxilla as in Molophilus, but outer lobe with the vestiture of hairs rather longer.

Pupa.— Length, 7.2-S.2 mm.
Width, d.-s., 0.8-0.9 mm.
Depth, d.-v., 1-1.1 mm.

Head light brown; thorax anterior to declivity conspicuously darker brown; remainderof bodyhght yellowish brown; breathing horns hght yellow.

(In fully colored individuals

the head and the thorax with their appendages become much darker, almost black, but thebreathing; horns retain their conspicuous yellow color.)

Head short, face tumid.
female smaller.

Eyes of male very large, widely separated by front;eyes of

Cephalic crest consisting of two prominent lobes; viewed from side, theselobes sharply pointed and directed slightly forward, with a seta on outer ventral face before
tips;

viewed from front, lobes rounded, ending in acute tips, separated by a deep, rather
Front between eyes narrowed toward labrum, which is rather

narrow, V-shaped notch.
sharply pointed.

Labial sheaths small, the lobes contiguous with their apices truncated,

the lateral angles obtuse or produced into a tiny lobe.

Sheaths cf maxillary palpi shortand stout, tapering gradually to tips. Antennal sheaths moderately elongated, angulatod
Pronotal breathing horns stout, expanded
at segments, ending just beyond base of wing.
at base, almost straight and only slightly divergent, somewhat compressed, transversely
wrinkled basally; a small setiferous tubercle in front of base of breathing hern.

Mesonotum

precipitous, at crest (Plate LXVII, 359) on cither side of median line with abundant tiny
setiferous tubercles bearing long, pale hairs, these tubercles continued back along shoulder.

Lateral margin of thorax with two small setae.

abdominal segment.

Wing sheaths ending before tip of second

Leg sheaths (Plate LXVII, 360)

short,

attainingbase offourthabdominal segment; tarsi of hind legs the longest, those of middle pair the shortest; fore legswith femora and tibiae very short.

Abdominal segments (Plate LXVII, 361)

divi.led into two annuli by a constriction near

midlength, the anterior ring very indistinctly subdivided further into two lesser annulets; onsegments 4 to 7, before caudal margin of po: terior ring on both dorsum and sternum, a transverse row of small, conspicuous, blackened, setiferous tubercles, which are more distantfrom one another near ends of rows; on basal abdominal segments t'.icse tubercles less evident

but still present.

Pleura with small but probably nosi-functional spiracles, which are very

indistinct in young pupae but are more evident in fully colored individuals; these spiracles

Setae on abdomen as follows: on sternal segments,
one seta just caudad of end of row of spicules, a second at lateral end of this row, interlocated near base of posterior ring.

mixed with spicules, two on posterior ring on a level with spicules; on tergal segments, astrong seta on a line with spiracles, another seta below end of row of spicules; on pleura, oneseta just ventrad of spiracles, and two post-spiracular and one ante-spiracular setae.

I\^a'eCauda (Plate LXVII, 362 and 363) with ventral lobes obliquely truncated, blunt at tips,
projecting beyond level of subacute dorsal lobes; dorsal lobes slender, slightly divergent
apically,

blackened before tips and with two setae at tips on outer face; at base of cauda,

Charles Paul Alexander918

on dorsal face of eighth segment, a close quadrangle of four rounded tubercles, placed on
Female cauda with tergal acidothecae elongate, subacute at tips; sternala slight elevation.
valves short, blunt; quadrangle of tubercles on dorsum of eighth segment more distinct
than in the male.

———

Nepionotypc.
Ithaca, New York, May 14, 1917.
Neanotype.
Ithaca, June 5, 1917.
Paratypes.
Larvae and pupae in large numbers from type locality,

May 14 to June 5,

1917.

Erioptera chlorophylla O. S.
1859

Erioptera chlorophylla 0. S.

Proc. Acad. Nat. Sci. Phila., p. 226.

is conspicuous by its
pale green color in theand adult stages. It is a common and widely distributed
species thruout eastern North America.
Several larvae were found inorganic mud at Orono, Maine, on June 13, 1913, one of which pupatedon the 21st. When the insect is dropped into boiling water, the green
The associates of this species are discussed
color immediately disappears.
under the account of Ptychoptera rujocincta (page 775).

Erioptera chlorophylla

larval, pupal,

Larva.

— Length, 9-10 mm.
Diameter, 0.7-0.75 mm.

Color uniformly pale green, fading to a pale yellow after death.

Form moderately elongated, last segment of body elongate, gradually narrowed to bluntly
Body clothed with numerous appressed hairs. Spiracular disk somewhat

rounded apex.

more reduced, disk usually entirely closed, lateral lobes
on either side capable of close approximation, tracheae before the opening into spiracles

as in E. megophthalma but evenvery large.

Head capsule (Plate LXVIII, 364) of the Molophilus type, but longer and slenderer; thetwo dorsal bars of each side very delicate, the ventral bars broader and flattened. Mental
plates (Plate LXVIII, 365 and 366) slender; anterior end of each ventral bar widely
expanded and provided with several teeth; these teeth varying in number, in some specimens there being only five, in others eight, teeth to each plate; in the latter case the thirdfrom either side is larger, with two smaller teeth between. Hypopharynx about as in
Molophilus. Antenna (Plate LXVIII, 367) large, basal segment stout, cylindrical; apical
papilla elongate-oval, with apex bluntly rounded and surface weakly sculptured; laterad of
Mandible (Plate LXVIII, 368) rather large;
this papilla a tiny cylindrical hyaline peg.
cutting edge with about four slender teeth, the second from base the smallest; dorsal faceof mandible with a blunt subapical tooth and an oblique comb of about six stout setae or
Maxilla similar to that of Molophilus, but the hairy vestiture longer andchitinized teeth.
coarser.

Pupa.

— Length, 8.8-9 mm.
Width, d.-s., 1.2 mm.
Depth, d.-v., 1.2 mm.

The Crane-Flies of New York— Part919

II

Breathing horns reddish brown; thoracic dorsum green, with a brownish tinge;
abdomen uniformly pale green, posterior half of each dorsal segment a little darker.
Cephalic crest small and compact, consisting of two prominent but closely approximatedlobes which are separated by a deep U-shaped notch, the tips acute and directed forward;

on outer face before apex a short seta.
triangular, divergent.

Labrum broadly obtuse at tip.

Labial lobes roughly

Maxillary palpi short and stout, narrowed towardtip.

Antennal

sheaths with a slender tubercle at base above eye.

Pronotal breathing horns broad at base, narrowed to the acute tip which is directed almost
ventrad; viewed from above, horns very broad basally and with a dorsal carina; ventral
side at base transversely wrinkled; a small setiferous lobe just in front of breathing horns,

directed laterad.

INIesonotum behind breathing horns with a high compressed carina, on

either side of this produced into a lobe directed cephalad and laterad; mesonotum moderately

decHvitous (Plate

LXIX, 369), at crest with numerous setiferous tubercles which are fewer

in number and more widely separated along shoulder.
of fourth abdominal segment;

Leg sheaths reaching to about middle
hind legs a little longer than fore legs; middle legs very short,ending opposite base of last segment of fore legs.

Abdominal segments divided into two narrow basal rings and a broad posterior ring;on pleura a distinct spiracle, opposite posterior annulus and nearer dorsal margin; posterior
annulus, before caudal margin, with a dorsal and a ventral row of long, stout setae.
Setae
on abdomen as follows: on pleura, a seta opposite second basal ring, a second ventrad ofspiracle, and two setae caudad of spiracle, the posterior one a little more dorsal in position;
on tergites, two stout setae lying transversely on the margin opposite spiracle, a third setaat end of terminal rows of bristles; on sternites, a group of two transverse setae on posterior
below level of spiracle and rather widely separated by the broad midventral
Female cauda (Plate LXIX, 370) with dorsal acidothecae short, distinctly upturned,
and ending in a small, subacute tip; before apex with two very short setae; a short blunt
tubercle near base of valves; ventral lobes short, their tips very blunt; dorsum of segment 8
with two blunt median tubercles, one immediately behind the other.
(Described from larvae taken in the Basin Swamp, Orono, Maine, June 13, 19L3; one pupa
ring, slightlyarea.

with the larva, July 5, 1913.)

Erioptcra vcspertina 0. S.
1859

Erioptem vespcrUna 0. S.

Proc. Acad. Nat. Sci. Phila., p. 220.

Erioptera vcsperlina is a characteristic inhabitant of open swan\ps andwet meadows. It has not been reared, but a pupa found in organic mudin the Basin Swamp, Orono, Maine, on June 24, 1913, undoubtedly belongs
to this species.
The associates are discussed under the account of Bittacomorpha clavipes (page 785).
Pupa.

— Length,

8. .5

mm.

Depth, d.-v., 1.1 mm.
Breathing horns reddish brown; thoracic dorsum reddish brown, with an interrupted
down posterior half of mesonotum; sheaths of wings and legs pale

whitish line running

Charles Paul Alexander

920

brown; abdomen whitish yellow, posterior half of each sternite and tergite dark brown,basal half with two narrow transverse lines of same color.
Lobes of cephalic crest widely separated, subtriangular, acutely pointed at tips. Pronotal
breathing horns cylindrical, transversely wrinkled, tapering gradually to tips.

(In the only

specimen at hand, the horns are v/idely separated at the base but soon bend proximad andt'lenceoutward and ventrad, so that the two horns are closely approximated or almost
Behind breathing horn-, mesonotum with conspicuous

contiguous on their distal part^.)divergent lobes such as are described for E. chlorophylla; mesonotum at crest with abundant
black setiferous tubercles, which are fewer in number, smaller, and more scattered along

Hind legs a little longer than fore legs, which,s'loulder, interrupted at mid-dorsal line.

in turn, are a little longer

than middle legs.

Abdominal segments with subterminal armature of posterior ring more spinous than in
E. chlorophylla.

Distribution of setae about as in E. chlorophylla.

(Described from a pupa taken at Orono, Maine, June 24, 1913.)

Erioptera septemtrionis O. S.
1859

Erioptera septemtrionis 0. S.

Erioptera septemtrionis

isProc. Acad. Nat. Sci. Phila., p. 226.

a widely distributed species thruout thenortheastern United States and Canada.
inrich organic

Tho larvae are not uncommon
The writer found them

mud in cool, shaded woods.

CDmmonly in the Standpipe Woods, Orono, Maine, in July and August,1913.
A larva placed in rearing on July 3 transformed to an adult female
on the 16th. Other larvae placed in rearing on July 15 transformed toadult males on the 25th. This limits the pupal duration to not more thanton days, but it is probably much less, presumably about one week.

— Length of cast skin, 7 mm.

Pupa.-

Cephalic crest low and flat.

Antennal sheaths moderately elongated, individual segments

showing clearly thru sheaths.

Pronotal breathing horns elongate, cylindrical, almost straightbut slightly diverging, transversely wrinkled, paler at tips, with a row of small breathing
pores along apical margin; a few small setiferous tubercles before base of breathing horns.
Declivity of mesonotum rather steep, at crest with numerous small tubercles and abundantpale yellow hairs which are less numerous along shoulder.

Leg sheaths with middle pairconspicuously the shortest, as in the genus.

Abdominal segments with subterminal armatureof posterior ring consisting of stout,

pale setae; arrangement of these setae about as in E. chlorophylla.

Lateral spiracles distinct.

Male Cauda (Plate LXIX, 371 and 372) with the ventral lobes blunt at tips, obliquely
truncated, separated by an acute V-shaped notch; dorsal lobes separated by a U-shaped
notch, each lobe terminating in a small

tip,with two

tiny

setae

on

lateralface before

apex; dorsum of eighth segment with four prominent, pale, fleshy lobes which are closely

approximated, the anterior pair directed laterad and a little more distant from each otherthan the posterior pair, which are directed more dorsad.

Neanotype.— Orono, Maine, July 25, 1913.

No. 105-1913.

The Crane-Flies of New York— Part II

921(Subgenus Hoplolabis Oston Sackcn)
1869

Mon. Dipt. N. Amer,, part 4, p. 160.

Hoplolabis O. S.

The subgenus Hoplolabis includes but three known species— the typeof the group,

Erioptera (Hoplolabis) armata, discussed below; E.

bipartita 0. S.,of western

North America; and E. (H.)(H.)

asiatica Alex.,of Japan.

Erioptera (Hoplolabis) armata 0. S.
Proc. Acad. Nat. Sci. Phila., p. 227.

Erioptera armata O. S.

1859

Erioptera armata

commona ratheris

flythruout the northeastern

A larva was found in the sand along the banks of Fall
Creek, Ithaca, New York, on May 16, 1917. This larva was of theUnited States.

and with the spiracular
was placed in rearing andThis larva was found associ-typical eriopterine form, being elongate, terete,disk surroundedby five subequal lobes.

Ittransformed to an adult female on May 31.
ated with numerous hexatomine larvae, such as Eriocera spinosa, E.
longicornis,

and E.

cinerea.

The following description is from the cast pupal skin.
Length, about 7 mm.
Cephalic

crest

consistingof

two moderately large, slightly divergent lobes which areacutely pointed at tips; lobes directed strongly outward and bearing a seta on outer face.

Labrum acutelj^ pointed.

Labial lobes large divergent, almost straight across caudal margin.
,

Sheaths of maxillary palpi stout, rather pointed at tips.
at base,tapering

tubercle before

to slenderbase ofeach

apices,bases

breathingconspicuously

horn.

Pronotal breathing horns broad
wrinkled;

a

small

setiferous

Mesonotum at crest (Plate LXIX, 373)

with numerous short, chitinized points.
seta above

wing axil.

Lateral angle of thorax with two setae; a strong
Leg sheaths with middle tarsi ending conspicuously before tarsi ofother legs.

Each abdominal segment before posterior margin with a transverse row of slender blackon tergum and sternum, and smaller areas on pleura. Lateralabdominal spiracles distinct. »Setae on pleura just ventrad of spiracle, and on tergum justabove transverse row of spicules. Female cauda with tergal valves elongated, gently
upcurved, unarmed; sternal valves shorter, blunt; at base of cauda, on dorsum of eighth segspicules or short bristles

ment, four small darkened tubercles which are produced into slender tips.
Neanoiype.

— Cast pupal

skin, Ithaca,

New York, May 31, 1917.

(Subgenus Mesocyphona Osten Sacken)
1869

Mesocyphona 0. S.

Mon. Dipt. N. Amer., part 4, p. 161.

Mesocyphona is one of the larger subgenera of Erioptera, reaching its

maximum of specific development in the Tropics of the New World.

Charles Paul Alexander

922

The immature stages
E. (M.) parva 0.

Erioptera (Mesocy phono)

ofS. arespent in wet

caloptera

(Say)

and

mud along the banks of streams

and other bodies of water.

The species discussed below as Erioptera(Mesocyphona) species (n(^ar knabi), was reared from the sandy margins
of a small prairie stream in Kansas.
Erioptera (Mesocyphona) species (near knabi Alex.)

Adult flies of a small species of Mesocyphona which is close to E. (M.)
knabi Alex., of Mexico, were not uncommon along Buckner Creek, a small
stream flowing thru Jetmore, Kansas.

prairie

These adults, especiallythe females, were photophilous, appearing in considerable numbers around
lanterns which were hung in tents pitiched along the banks of this stream.

A single pupa found in the muddy sand along the bank of the creek on
July 20, 1917, emerged as an adult on the 22d. The following generalcharacters of the species

may be noted:— Labrum small, apex rather sharp.

PuTpa.-

angles subacute.
long, cylindrical,

Labial lobes squarely truncated, with lateral

Sheaths of maxillary palpi slender.

Pronotal breathing horns moderately
curved slightly forward, enlarged outwardly, blunt at tips. Mesonotumat crest rather tumid and with a few long hairs inserted on stout black tubercles.

Leg sheaths

with middle tarsi the shortest, hind tarsi the longest.

— Jetmore, Hodgeman County, Kansas, July 22, 1917.

Neanotype.'

Genus Ormosia Rondani (Gr. chain)
1856
I860
1863

Ormosia Rond. Dipt. Ttal. Prodr., vol. 1, p. 180.
Rhi/pholophus Kol.
Wien. Ent. Monatschr., vol. 4, p. 393.
Dasypiera Schin. Wien. Ent. Monatsclir., vol. 7, p. 221.
.

— Form terete, moderately elongated.

Larva.'

Spiracular disk squarely truncated, sur-rounded by five subequal lobes which are lined with double marks of brown.

Anal gills

Head capsule slender, very dissected, of six narrow bars, four dorsal and two ventral,
ventral bars broader.
Labrum narrow, epipharynx hairy. Mandible with teeth

blunt.

themoderately elongated.

AntennaoftheMolophilus type.

Men turn

without chitinized

teeth.

— Cephalic crest setiferous.

Pupa.'

Pronotal breathing horns rather short and stout,

more or less flattened and with a row of tubercles along posterior margin.

Mesonotum

declivitous, at crest with a flattened, toothed, chitinized plate on either side (in 0. nuhila)

Wing sheaths ending opposite or just beyond tip
Leg sheaths varying in length with the different species,
middle tarsi the shortest. Abdominal segments with a subterminal transverse row of spines
or setae, these occurring on pleura (as small groups of eight to twelve) as well as on tergites
and sternites. Lateral spiracles distinct, on segments 2 to 7; dorsum of segment 8 withor with abundant setiferous tubercles.
of second

abdominal segment.

The Crane-Flies of New Yorka pentagon of five lobes, these being— PartII

923

unarmed (0. innocens, 0. meigenii) or spinous-tipped(0. nubila, 0. nigripila).

Ormosiaisa large and rather

difficultgenus (including more thanseventy-five species) of small crane-fli(^s whichai'ccharacteristic of sub-arctic and temperate regions and apparently rare or lacking in the Tropics.

The adult flies occur in small dancing swarms, usually in cool, shaded
woods or along brooks. They are most numerous
in early spring and in late summer or early autumn, many of the species
situations in or near

being apparently double-brood 3d.

The innnature stages are spent in moist organic mud near water. In
Europe, Beling records Ormosia haernorrhoidalis (Zett.) (Beling, 1879:
48^9, and 1886:191-192), 0. lineata (Meig.) (Beling, 1886:202), and0. rwdulosa (Macq.) (Beling, 1886:202), as being found in wet earth inwoods, usually beneath leaf mold.

Cameron (1917 65) likewise records the:last-named species as living in, mud. 0. varia (Meig.) was found by De
Meijere (1916:201-204) among decaying leaves in a wet spot near a ditch.
Of the American species, the writer has bred Ormosia innocens, 0.

and 0. nigripila from larvae or pupae in entirely
European species.
The larvae of only two of these species, 0. nubila and 0. meigenii, are
0. nubila is a large brown species;
available to the writer at this time.
0. meigenii is much slenderer, and is light yellow in color.
The pupae of the known Nearctic species may be distinguished asnubila,0. meigenii,

similar situations to those given above for thefollows:1.

Dorsum of eighth abdominal segment with a pentagon of five lobes which are spinousat their tips

Dorsum of eighth alxlominal segment with a pentagon of five fleshy lobes
2. Crest of mesonotum with a flattened chitinized plate on either side of median line.

3.

2
3

0. nubila (O. S.) (p. 923)
Crest of mesonotum tumid, with abundant coarse, yellowish setae on either side of median
0. nigripila (O. S.) (p. 927)
line
Abdomen with large, rectangular, dusky areas on posterior annuli of segments 2 to 8,
giving abdomen a banded appearance; pleural setae rather numerous.
0. innocens (O. S.) (p. 925)
Abdomen without such dusky areas; pleural setae few in number.
0. meigenii (0. S.) (p. 928)

Ormosia nubila (0.
1859

S.)

Erioptera nubila 0. S.

Proc. Acad. Nat. Sci. Phila., p. 227.

Ormosia nubila is probably double-brooded, since the flies are on thewing in the spring and again in the fall. The immature stages live inCharles Paul Alexander

924

organic mud that is usually covered over with a layer of leaf mold.

On
March 27, 1914, the writer found four lai'ge l)rown eriopterine larvae infrom Needham's Glen, Ithaca, New York. An adult female
emerged on April 9. This makes the maximum
pupal existence less than two weeks, bu,t it is undoubtedly very muchless than this.
rich earthof the present species

Larva.

— Length,

11. 5-11. S mm.
Diameter, 1.1-1.2 mm.

Color a deep reddish or cinnamon brown; incisures of segments paler.

Formrather stout,

Spiracular disk (Platebody

terete.

Skin covered with a short, appressed pubescence.

LXX, 375) squarely truncated, surrounded by five subequal lobes;

ventral lobes with two parallel dark brown lines which are narrowly united distally, the pale
stripe between rather broad, distinct, especially near center of disk; lateral lobes withparalleldark brown stripes which are

indi.stinctly

connected

distally,

two

the space between

dusky with numerous brown spots; dorsal lobe with an elongate-oval mark inclosing a linear
yellow center; lobes fringed with long hairs which are longest at tips, shorter toward base,and narrowly interrupted between lobes; di.sk between spiracles unmarked. Spiracles large;
middle piece and extreme outer margins of ring blackish; spiracles separated by a distanceabout equal to one and one-half times diameter of one.
Head capsule about as in Molophilus, but ventral bars of capsule not toothed to form
Labrum and epipharynx about as in Molothe characteristic mental plate of that genus.
Hypopharynx broad, flattened, provided with numerous transverse rows of shortphilus.
Antenna short; basal segment stout, cylindrical; apical papilla rather small,setae.
Mandible (Plate LXX, .374) ending in a rather long apical point, with about
elongate-oval.
four long, flattened teeth along ventral cutting edge, the second from base very small; a short,recurved hook at prosthecal region and a den.se tuft of long yellow hairs in prosthecal slit; aMaxilla about as in Molophilus.
dorsal appendage at heel of mandible.
Pupa.

— Length, 7.5 mm.
Width, d.-s., 1.2 mm.
Depth, d.-v., 1.4 mm.

Head, thorax, and appendages pale yellow; breathing horns yellow; chitinized plates on(In mature pupae, the sheaths of the appendages

mesonotum dark brown; abdomen brown.
are probably darker.)

Cephalic

crest(Plate

LXXI,

381)

low;

lateralanglesproduced into conical,

spinous tubercles, each bearing a stout seta on outer ventral face.

erect,

Antennal sheaths very

Opposite each segment of antenna on basal half of organ, aangulated, almost serrate.

conspicuous blackened tubercle, tho.se at base larger and more con.spicuous, the lateral one
directed outward, aboveit

a second tubercle

directed cephalad

Antenna extending to just beyond base of wings.

and simulating a crest.

Front broad; a blackish area on eitherside near inner margin of eye, probably indicating point of attachment of tentorium.

triangular,

apex subacute.

Labial lobes triangular, tips blunt.

rather long, narrowed to the slender tip.

LabrumSheaths of maxillary palpi

Pronotal breathing horns short and slender, some-

The Crane-Flies of New York— Part II

925

what flattened, pale yellow; a stout seta just before base of each breathing horn.
precipitous; at crest (Plate LXXI, 380) on either side of medianflattened plate projecting dorsad; this plate, atlateral

its

tip,

line,

Mesonotuma heavily chitinized

forking into two lesser teeth, theone of these still further subdivided into two still smaller teeth(in

one specimen

the left plate is divided dichotomously into four teeth, the inner primary tooth being further

subdivided); on shoulder laterad of these plates a slightly swollen plate which is parallelly

grooved; two spines on lateral angle of thorax above base of wing; mesonotum with a few
setae behind crest, as follows: an anterior solitary seta on either side of median line and close
to it, a solitary stout seta above wing axil, two groups of paired .setae, one just dorsad of baseof wing and the other midway between this group and the anterior seta first described.

sheaths rather short, ending opposite base of third abdominal segment.

Wing

Leg sheaths rather

long for this genus, ending opposite midlength of fifth abdominal segment; fore tarsi a little
shorter than hind tarsi, middle tarsi very short, ending opposite or just beyond end of third
tarsal segment of fore legs.

Abdominal segments near posterior margin with transverse rows of short black spineg
which are much smaller and more widely separated near ends of row; at intervals alona
row a few elongate setae; on pleura a similar area of eight to eleven spines with a single sets
Chaetotaxy as follows: on pleura, justin row; tubular lateral spiracles on segments 2 to 7.
above dorsal end of row, one seta, another solitary seta cephalad of spiracle on anterior
on tergites, a solitary seta on posterior ring about opposite spiracle; on sternites, two
Female Cauda (Plate LXXI, 3S2)

ring;small setae on either side of median line of posterior ring.

with tergal valves long and slender, almost straight but slight^ upcurved near

tips;

on

dorsal lateral margin, just before tip, a prominent tooth directed laterad and slightly caudad;

about midlength of valves a somewhat similar blunt tubercle on either side near dorsal margin;
at base of eighth tergite a pentagon of five chitinized lobes, the anterior one rudimentary,and each bearing a subapicalmore widely separated and bearing on lower side near base a

the four developed lobes ending in acute chitinized points
seta; anterior pair of lobes

small, slender lobule.

— Ithaca, New York, March— Ithaca, with type
April
— With types.

Nepimwtype.

27, 1014.

Neanotype.
Paratypes.

Ormosia innoccns (0.
18G9larva,

9, 191-4.

No. 2-1914.

S.)

Rhypholophus innocens 0. S.

Mon. Dipt. N. Amer., part 4, p. 142.

Ormosia innocens is a chai'acteristic early spring species, the adultbeing found in late April and during May. On May 12, 1917, a
few pupae in an advanced stage of development were sifted from organic
mud from Bool's hillside, Ithaca, New York. One of these emerged as
an adult on the following day. The associated crane-fly larvae that
occurred with this species on the date named were as follows: Bittacomorphella jonesi, Dicranomyia stidta, Limnoj)hila adusta, L. fiiscovaria,
JJlomorpha pilosella, Penthoptcra albilarsis, Rhaphidolabina Jlaveola,flies

Charles Paul Alexander

926

Molophilus hirtipennis, Erioptera megophthalma, Ormosia nigripila, Tipula
oropezoides, T. collaris, T. cayuga.

Pupa.

— Length, 7.5-10.5 mm.
Width, d.-s., 0.8-1 mm.
Depth, d.-v., 1-1.2 mm.

(The smaller measurements are those of males, the larger those of females.)Head and thorax brown; mesonotum before declivity darker brown; abdomen pale yellownsh
white, the sternal and tergal sclerites with broad, rectangular, darker areas, producing abanded appearance; breathing horns pale yellow. (In older pupae, the coloration of thehead and the thorax is much darker.)
Cephalic crest of moderate size, bilobed; viewed from side, lobes blunt, with two lateral
Between antennal
setae; viewed from front, lobes separated by a very broad, V-shaped notch.

forehead longitudinally grooved,bases,

these

lines

converging between eyes.

triangular, subacute, separating the small labial lobes.

tapering to blunt apices.

Ivabrum

INIaxillary palpi of moderate length,

Antenna of moderate length, reaching to just beyond wing base.

Pronotal breathing horns rather short and stout, almost straight and only slightly divergbase.
Mesonumerous palehairs, these narrowly interrupted on median line; lateral angle of thorax with two tiny setae;
a strong seta slightly dorsad and cephalad of wing root; about four small setae in alinement
Wing sheaths moderately broad, ending at or just before tip of second
across mesonotum.
abdominal segment. Leg sheaths short, tips of all the tarsi ending about on a level, or
those of fore legs a little longer, terminating just before end of third abdominal segment.
Abdominal segments divided into two distinct annuli by a constriction at about midlength of segment; sternum and tergum of posterior ring of segments 2 to 8 near caudal
margin with large, rectangular, dusky areas which appear subchitinized; caudal margin of

ing,

subcylindrical,

notum

compressed,

precipitous, carinate

andslightly

medially; atcrest

constricted(Platejust

LXX, 376)

beyondwiththis area with a fringe of long, black hairs, there being about seventy-five of these on intermediate segments; on pleural membrane a very small, similar area bearing from twenty-fiveto thirty-nine hairs, and near its dorsal margin a distinct black tubular spiracle on segments
2 to 7; the following additional setae on segments: on dorsal segments, rectangular darkened
areas with sparse scattered elongate hairs ov'er surface, and two setae on each anterior ventral
angle, their arrangement oblique; sternal segments similar, but the two setae on rectangulararea arranged transversely; just caudad of ends of fringe of hairs, one or two isolated hairs;
pleural membrane opposite basal annulus with a single seta; another seta opposite posterior

annulus near ventral margin; a third seta on ventro-cephalic angle of pleural setiferous
area just above level of spiracle; segment 8 on dorsum with five pale, gill-like lobes arranged
in a quadrangle or apentagon, the anterior lateral pair the longest, the median one rather thetwo setae on cephalic face; at base of posterior pair, aMale cauda (Plate LXX, 377 and 378) terminating in two verysmallest; at base of anterior pair,single setaon lateral face.

blunt ventral lobes and two separated, slender, dorsal lobes projecting caudad and dorsad.

Female cauda similar to male cauda, but tergal acidothecae very elongate, much longer thansternal valves.

— Tthaca, New York, May— Seven pupae, with type.

Neanotype.
Paratypes.

12, 1917.

The Crane-Flies of New York— Part927

IIOrmona nigripila (O. S.)
1S69

Rhypholophus nigripilus O. S.

Mon. Dipt. N. Araer., part \, p. 142.

The larvae of Orniosia nigripila are common in rich organic mud inshady places. Larvae found in Needham's Glen, Ithaca, New York,on INIarch 27, 1914, emerged on April 18. Other larvae from Coy Glen,
found on April 17, emerged on May 1. At Orono, Maine, large larvae
were found on July 14, 1913, and were placed in rearing, emerging on
the 26th as adult males. This limits the pupal duration to not more than
two weeks, but it is undoubtedly much less, probably not more than afew days or a week.
Pupa.

— Length,

5.2 mm.

Width, d.-s., 0.9 mm.
Depth, d.-v., 1 mm.

Head, thorax, and appendages hght brown, becoming darker in maturity; breathing horna
and al:)domen pale whitish.
CephaHc crest with lobes small, low, and rounded, with a powerful seta at tip, directed
forward.

Labrum blunt at apex.

pointed.

Sheaths of maxillary palpi stout, rather elongate.

Labial lobes large, subtriangular, lateral angles obtusely-

Pronotal breathing horns(Plate LXXn, 3.S4) rather short and flat, compressed, slightly expanded beyond base; outer,or posterior,margin with about five small tubercles, at

least

one of whichissetiferous.

Mesonotum very steep and precipitous, as in this group of species; crest (Plate LXXII,383)
tumid, extensive, on either side with abundant coarse yellow bristles; caudad of these, four
stout setae in a quadrangle, two on either side of median line; lateral angles of thorax with

two stout setae; a strong seta just above wing base. Wing sheaths reaching base of third
Leg sheaths mod?rately long, extending almost to midlength of fourth
abdominal segment; tarsi of middle legs much shorter than the others, hind legs a very little
abdominal segment.

longer than fore legs.

Abdomen with segments (Plate L XX 1 1, 38.5) before their caudal margin bearing transverserows of slender spines; pleura with a small area lying a little cephalad of tergal and sternalrows and margined behind with eight to ten spines. Spiracles distinct, tubular. Setae
as follows: on pleura, a seta on anterior annulus, a stout seta just ventrad of spiracles, anda third lying a little ventrad and cephalad of spiracle; on sternites, two stout setae nearbase of posterior annulus; on tergites, two setae on posterior annulus, lying transversely
at level of spiracle, and a third seta just cephalad of end of row of spines.

Male cauda (Plate
LXXII, 386) with ventral lobes rather slender, narrowed outwardly, and somewhat pointed

at apex; dorsal lobes curved strongly backward, terminatingin acute,

chitinized points

with a strong seta on outer face before tip and a second seta nearer base; dorsum of segment 8
with five brown, chitinized lobes which are crowned at their apices with a circlet of spines;
posterior pair the longest and stoutest, broad at base,more slender outwardly, with a large
and about three smaller inner sp'nes; anterior pair more slender; median lobe
slender, crowned with a circlet of about six small, subequal spines.
Female cauda (Plate

lateral spine

Charles Paul Alexander

928

LXXII, 387) with dorsal terebra elongate, almost straight or very slightly upturned; eighth
segment with dorsal pentagon of spine-tipped lobes quite as in male.

Ormosia meigenii (0. S.)
1859

Erioptera meigenii 0. S.

Proc. Acad. Nat. Sci. Phila., p. 226.

Ormosia meigenii is one of the commonest species of the genus, occmring
swarms in early spring. Larvae of this species weretaken on April 10, 1914, in organic nmd from Needham's Glen, Ithaca,
New York. An adult female emerged on May 4.
in small dancing

Larva.

— Length,

6.5 mm.

Diameter, 0.5-0.6 mm.
Color, light yellow.

Form elongate, terete.

Spiracular disk (Plate

LXX, 379) about as in 0. nubila, dorso-

median lobe the smallest, ventral lobes with brown lines rather close together, the distalline the broadest; lateral lobes

with lines rather short, not contiguous at their distal ends

and not extending past midlength of spiracles; dorso-median lobe with marks oval, solidly
dark brown.
atextremeLobes fringed with moderately long, yellowish setae, which are almost lacking
tips;

disk

between spiracles unmarked.

Spiracles large,transversely oval,

separated by a distance a little greater than the long diameter of one.

Head capsule and mouth parts almost as in 0. nubila; mandible with the long, slender
teeth of that species.

Pupa.

— (Described from a castskin.)

Cephalic crest consisting of low, rounded lobes, each with a seta on anterior lateral face.
Pronotal breathing horns of moderate length, flattened, a little narrowed toward tip, outermargin with fine tubercles.

Mesonotum not so declivitous as usual in the genus, with tiny

roughenings at crest; the usual two setae at lateral angle of thorax present, another above
each wing, and four more in a transverse row at level of axilla of wing; a seta on mesonotum,
close to median line, just below crest.

Middle legs much shorter than fore and hind legs.

Subterminal rows of setae on abdominal segments consisting of slender, acute spines,
with a few setae interspersed; on pleura the spines few in number. Spiracles elongate,

Female cauda with tergal sheaths very long, sternal sheaths short, their tips
dorsum of eighth abdominal segment five small unarmed tubercles, which areblunt or nearly so, anterior pair more widely separated than posterior pair.
(In the shapeof the cephalic crest, the breathing horns, and the declivity of the mesonotum, this species
resembles 0. nigripila; but the arrangement of setae on the abdomen, and the great reduction
and unarmed condition of the lobes on the eighth abdominal tergite, are distinctive.)tubular.

blunt; on——

Nepionotype.
Ithaca, New York, April 10, 1914.
Neanotype.
Cast pupal skin, Ithaca, May 4, 1914.

Genus Helobia St. Farg. et Serv. (Gr. marsh + / live)
1825
1830

18%
1886

Helobia St. Farg. et Serv. Encyclop. Method. Ins., vol. 10, p. 585.
Symplecta Meig. Syst. Beschr., vol. 0, p. 282.
Idioneura Phil. Verb. Zool.-Bot. Ges. Wien, vol. 15, p. 615.
Symplectomorpha Mik. AVien. Ent. Zeitung, vol. 5, p. 318.

The Crane-Flies of New York
Larva.

— Form moderately elongated, body

terete.

— Part929

IISpiracular disk surroundedby five

subequal lobes which are marked with V-shaped brown lines, in some specimens the inner

Anal gills moderately
Antenna with the apical papilla very short,faces of the three most dorsal lobes being entirely brownish black.

Head capsule as in the Eriopterini.

elongate.

subpyriform.

Pupa.— Cephalic crest setiferous.

Pronotal breathing horns elongate-cylindrical, directedventrad and cephalad, with rows ofbreathingpores

along dorsal face.

Rlesonotum

declivitous, at broad crest armed with numerous chitinized spines;

an arcuated longitudinalrow of six small pits extending from wing axil toward crest of thorax. Wing sheaths ending
before tip of second abdominal segment.

abdominal segment;

tarsi oflegs intermediate in length.

Leg sheaths ending about opposite tip of third

hind legs the longest, those of middle legs the shortest, fore

Abdominal segments with a strong subterminal armature on

ventral segments, much weaker to lacking on dorsal segments; lateral spiracles distinct on

segments 2 to 7; dorsum of segment S with five blunt lobes.

Helobia is a small genus (five species) of common and sometimes very
The only Noith American species, Helobia

widely distributed crane-flies.
hyhridn,isapparently the most widespread tipulid known, ranging overpractically the entire Holarctic region,

and,India,

inthesouthward in the mountains to

New World, to Central America. The immature

stages of the known species are spent in moist earth near water.

Bruch{in litt.) mentions the rearing of H. macroptera (Phil.) in Argentina.

Helobia hybrida (Meig.)
1804
1818
1830
1848

Limonia hybrida Meig. Klass., vol. 1, p. 57.
Limnobia punctipennis Meig. Syst. Beschr. Zweifl. Ins., vol. 1, p. 147.
Si/mplecla punctipennis Meig.
Syst. Beschr. Zweifl. Ins., vol. 6, p. 283.
List Dipt. Brit. Mus., vol. 1, p. 48.

Limnobia cana Walk.

Helobia hybrida is undoubtedly the most widely distributed North
American crane-fly. Beling (1879:50-51) found larvae and pupae atthe end of July, 1876, in wet, sandy earth along the margins of small
brooks in deciduous woods. Adults emerged in his breeding cages on
July 27 and August 6. Hart (1898 [1895]: 199-200) found the larvae insimilar sandy situations along the Illinois River, associated with the
He suggests that it may serve as foodlarvae of Tabanus atratus Fabr.
for this horse-fly larva.
Larvae of Helobia were especially abundant on
May 17, and these transformed to adults within a month. Females

wereobserved ovipositing along the shore, patting the valves of theagainst the moist sand.
Malloch ( 1915-17 b: 229-230) has

ovipositor

given additional notes on the structural details of Hart's material.

Charles Paul Alexander

930

Dr. Adam Boving found this species in Iceland and made careful noteson the burrows made h_y the larvae. Thru the kindness of Dr. Boving,
the writer is able to include a translation of his manuscript.
is indebted

The writeralso to Dr. Lundbeck, director of the museum at Copenhagen,for the loan of this material for study.
later in this paper.

Iceland, in 1908.

These are the specimens discussed

Boving's notes were made at Fell Station, southeast

The translation follows:

Inside the moraine of 1S77, in the low land where ice was standing in 1886, quantities ofdipterous larvae were found in the moist sand on the bottom of flat hollows which at times
are flooded by water and at times are partly drained, as was the case on the day when thefollowing observations were taken.
The whole dark, moist surface of the bottom was covered by an irregular system of slightly
elevated, long, tubular galleries, some of which were rather straight, some formed brokenlines, some peculiar arabesques, and some plain spirals.
The width of the galleries was about
the size of an ordinary pinhead, some a trifle larger, some a little smaller.
In the anterior
part of each gallery was found either a cylindrical white tipulid larva (llelobia) about one
centimeter long, or another dipterous larva of the same general size and appearance. The
larvae were found just below the surface.
It was not always easy to capture them, for when
I pushed my knife under the mouth of the gallery they moved quickly backward, and then,digging deeper into the soil, made a new gallery that branched off from the main one.
It
was not possible to distmguish the galleries of the crane-fly larvae from those of the other
dipterous associate.
Very often, from the mouth of the spiral galleries, one-third of abroken pupal skin stuck out; but larvae were found also in many of these galleries.
The imagines of t!ie two Diptera were present in large numbers, some flying close to theground, others resting on it. Both forms were long-legged and capable of running over
the water film.
I secured a pair of both in copulation.
The eggs were foimd on the moist
surface, singly or in small masses of two or three together.
The larvae feed, of course, on organic particles in the sand. The imagines were not observed
to take any nourishment at all; they copulated as soon as they had left the pupal skins, andI did not find them in any other place than on the bare, moist soil where the larvae lived;
not, for instance, on flowers growing near bj'.
A small carabid (probably Bembidion grapii Gyll.) was present in the locality in comparatively large numbers, evidently preying on the larvae of the Diptera.
A single carabid larva also was found; from its size and habitus it may very well be the larva of theBembidion.

A small black spider was probably feeding on the imagines of the Diptera. It did not makea regular web, but spun a number of single threads, each about two feet long, attaching them
to a piece of gravel and proceeding from this as a common center, spreading the threads
close to the ground like radii, and finally fastening the ends to small grains of sand.

The adult flies of Helobia hybrida are very common.

They are theon the wing in spring, some appearing in early
March or, in open winters, even in late February, They remain until
The writer has noted the females running about on tliclate in the fall.
wet sand along the l)anks of the Kaw River at Lawrence, Kansas, andovipositing quite as described by Hart (1898 [1895]: 199-200).
first

tipulids to appear

Larva.

— Length, 7.8-10 mm.
Diameter, 0.6-0.7 mm.

The Crane-Flies of New York— Part931

II

Coloration, pale brownish yellow.

Form moderately elongated. Body covered with a sparse appressed pubescence. SpiLXXIII, 388) surrounded by five lobes, the dorsal one notably smaller

racular disk (Plate

than the paired lobes; inner face of lobes with two subparallel brown
their outerlines,

connected atends to form narrow V's; disk marked with about six spots Ijetween spiracles,

the largest at base of ventral lobes; disk fringed with short hairs, which are interrupted fora short distance between lobes.

Anal gills moderately elongated, pale.

(Beling describes

the three dorsal lobes of the disk as having the entire inner face shiny blackish brown, andthe ventral lobes merely margined with brownish; there would thus seem to be some variation in the character and degree of markings in this species.)

Head capsule as in the tribe.

Antenna with the basal segment stout, cylindrical, theMandible flattened; teeth large but very bluntly

apical papilla very short, subpyriform.

rounded; apical tooth the largest, with a slightly smaller denticle cu either side.

Pupa.

— Length, 7-9 mm.

Cephalic crest small; lobes pointed, directed ventrad, each bearing a short, stout seta.

Labrum narrow.

Labial lobes large, divergent, caudal margin almost straight across.

Sheaths
narrowed at tip (Plate LXXIII, 390). Antenna moderately
angulated, ending just beyond base of wing. Pronotal breathing horns elongate-cylindrical,
of maxillary palpi slender,

directed ventrad and cephalad, with rows of breat'hing pores along dorsal face; ventral facetransversely wrinkled.

jNIesonotum moderately declivitous, at broad crest (Plate LXXIII,389) armed with numerous black chitinizedspines directed backward;these spines

most

num rous proximally, interrupted by a mediamspace, less numerous along shoulder; four setaeina transverse row across mesonotum, two on either side of medianline; lateralangle ofthorax broad and blunt, with two small setae, the outermost one the larger, the inner one

about half its size; an arcuated longitudinal row of about six pits extending from above

wing toward crest of mesonotum. Wing sheaths short, ending before tip of second
abdominal segment. Leg sheaths ending opposite or slightly beyond tip of third abdominal
segment; hind legs much the longest, middle legs much the shortest, fore legs intermediate
axil of(PlateLXXIII, 391).

Abdomen with a rather strong armature of stout black spines on sternal segments, onsegment 3 this appearing as a small area of about ten spines, on either side of tips of hindtarsi; tergal armature much weaker or lacking; lateral spiracles very distinct, tubular, inFemale cauda (Plate
LXXIII, 393) with tergal valves the longest, terminating in sharp cylindrical points; sternalvalves much shorter, blunt at tips; dorsum of segment 8 with five blunt lobes. Male cauda(Plate LXXIII, 392 and 394) with dorsal valves the longest, each terminating in a long,subacute, chitinized spine directed dorsad and laterad and bearing before its tip two stout
setae; ventral lobes stout and blunt, longer than dorsal lobes.
cast pupal skin the principal tracheal trunks being very conspicuous.

Nepionotype.
Neanoti/pe.
Paralypes.

—

Fell,

between Oracfa and Heineberg, southeastern Iceland.

— Cast pupal skin, with type.
•— Numerous cast pupal skins, with types

Museum).

(in the collection of the Copenhagen

Charles Paul Alexander

932

Genus Trimicra Osten Sacken (Gr. three + small)
Trimicra 0. S.

1861

Proc. Acad. Nat. Sci. Phila., p. 290.

Trimicra is a small genus of crane-flies, including about fifteen described
species found in most parts of the world, almost all being forms of moderate
size

and obscure coloration.

These various species bear a close resem-

blance to one another and are hard to distinguish specifically.

Bergroth and other European writers consider the genus Trimicra as
being the same as Psiloconopa Zetterstedt, but at this time the writeris not entirely willing to accept this view.
Beling (1879:48) described what he took to be the larva of one oftheseflies,

and

his

in

key to the larvae of

included it in close proximity to the Pediciini.

crane-flies(1886:206) he

As mentioned elsewherein this paper, it is highly probable that Beling described a pcdiciine larva,

but by an accident had larvae of Trimicra in his breeding jars, the latterlarvae emerging first and confusing the author.

Gerbig (1913:161-163) describes the real larva of Trimicra pilipes
the best-known species of the genus. It is found along themargins of flowing streams with muddy banks. It is a dark-colored(Fabr.),

larva,

about

diameter.

15

in length and about 2 millimeters inbody is similar to that of the larvae in the

millimeters

The skinof thetypical subgenus of the genus Limnophila, being covered with chitinous,
hairlike projections, which in Trimicra are longest on the dorsal surface of

On each segment^ there are solitary elongate bristles, above andbelow each of which is a gland. The spiracular disk (Plate LXXIII, 395)

the body.

surrounded by five nearly equal lobes, which have the inner faces
marked with equal blackened, chitinized areas. On the lateral margins
of each lobe, but occupying only the distal part of the lobe and not continuous around the disk, is a fringe of moderately long hairs. At the
tip of each ventral lobe are two bristles, and at the tip of each lateralislobe is a single bristle, these being inserted outside the line of hairs andsurrounded by a bright circular area.
in

some

crane-fly larvae(asThe ventral sensory bristles foundTipula variipennis) are lacldng.

At thebase of the lateral lobes are the spiracles, which are generally similar to
those in the subgenus Limnophila.
of the spiracles, the feltof the body.

Gerbig discusses in detail the structurechamber, and the musculation of this region

The Crane-Flies of New York— PartII

933

Briich (ui lilt.) mentions the rearing of Trimicra redproca (Walk.) in
Argentina from larvae very similar to that described above, occurring inthe same type of habitat.

Genus Empedomorpha Alexander (Gr. Empeda + shape)
1916

Empedomorpha Alex.

Proc. Acad. Nat. Sci. Phila., p. 507-508.

Empedomorpha isa monotypic genus of flics, evidently related to
Trimicra.
It is very curious in its marked sexual dimorphism, the malehaving an extremely largo, hairy stigma, which encroaches upon theadjoining veins and often distorts them. The fly is still very insufficiently
known. It is a prairie-inhabiting species, occurring from South Dakota
Nothing is known concerning the immature
to Texas and New Mexico.
stages, and the following observations on the habits and occurrence of
the adult flies are all that are available.

Empedomorpha empedoides (Alex.)
191G(?) Trimicra

empedoides Alex.

Can. Ent.,

vol. 48, p.

44-45.

Empedomorpha empedoides, as stated under the generic account above,a prairie-inhabiting species. Adult flies were found running about
on the sand flats of the Arkansas River, near Cimarron, Kansas, from
July 13 to 15, 1917, l)y H. L. Fackler and the writer. The vegetationof the sand bars here is very sparse to almost lacking, a few psammophytic
grasses being the main element.
Associated with these flies on the sandy
surface were a characteristic group of sand-loving insects, of which thefollowing were the most constant: larvae and adults of tiger beetles,
representing at least three species of Cicindela; ground beetles, Carabidae,
including such genera as Omophron, Dyschirius, Bembidion, Tachys, anda few others; rove beetles, Staphylinidae, such as Stenus; Heteroceridae,
Psammocharidae, Asilidae, Saldidae, and similar groups. In the cool ofearly evening, numerous small spiders that lurk in hollows and in deserted
insect burrows during the day emerge from hiding and become active.
The crane-flies were observed during the hours of bright sunlight, when
the temperature registered over 100° F. in the sun.
They run rapidly
over the moist sand, their course being very shifting and zigzag, quite
They fly readily but only for short distances,like that of tiger beetles.
and prefer to alight on the sand rather than on the vegetation. They
walk awkwardly over the sand, but are able to crawl up grass blades orisCharles Paul Alexander934

Tho habits of the adult flies are strikingly like thosesimilar objects.

of Helobia (page 930).

Th(^ wiitcr is sure that the larvae are to be foundin the sand in these same haunts.

Genus Qnophomyia Ostcn Sackcn (Gr. darkness + fly)
1859
1867
1911
Larva.

Gnophomyia O. S. Proc. Acad. Nat. Sci. Phila., p. 223.
furina Jaenn. Abhandl. Senkenb. Ges., vol. 6, p. 318.
Dasymallomyia Brun. Rec. Indian Mus., vol. G, p. 304.

— Body slender, tapering toward ends, with transverse welts on intermediate

abdominal segments.

Spiracular disk surroundedby five subequal lobes.

Anal gills con-sisting of four blunt, rounded lobes, constructed for propulsion rather than for a respiratory

function.

Head capsule moderately elongated, rather compact for the Eriopterini.

rather small, apical papilla elongate-oval.

Antenna

INIandible slender, with a long apical point andthree teeth along ventral cutting edge.

Pupa.

— Cephalicwith a long seta.

crest a low, blunt tubercle

on either side of median line, each tipped

Pronotal breathing horns small, narrowly trumpet-shaped.

Mesonotum

moderately declivitous, at crest practically unarmed; a strong seta at lateral angle of thorax
and another on either side of mesonotum behind crest. Wing sheaths short, ending before
tip ofsecond abdominal segment.

Leg sheaths very short, reaching just beyond wings,

attaining end of second abdominal segment; all the tarsi ending about on a level.

Abdomenprovided with long setae; spiracles on abdominal segments 2 to 7.

Gnophomyia is a small genus including about forty described species
which are most abundant in the Tropics of the New World. The European
Gnophomyia tripudians Bergr. has recently been reared by Gamkrelidze
The former found(1913, a and b) and l)y KeiHn (Edwards, 1919b).
larvae in large numbers in the viscous, semi-decomposed mass of tissue
beneath the bark of a fallen Carolina poplar, associated with Miastor
These larvae were found near Paris, France, in
metraloas Meinert.
March, 1911. Gamkrelidze records a gregarine parasite in the intestine
and a nematode worm in the body cavity. The species was later rearedin England from d(*ad oak by Kcilin, who has discussed and figured
glands in the larvae (1913:3). The only American species that has beenreared is the common Gnophomyia tristissima, discussed later.
Gnophomyia rufa Hudson, of New Zealand, has recently been discussed
It is occasionally found in dense
in some detail by Hudson (1920:32-33).
ofWellington.
The
larva lives in and feeds on thevicinity
theforests in
which
detritus
accumulates
in large quantities atvegetablesemi-liquidof
AdeliaSolayidri,
leavesacommonand very conspicuousofthethe bases
epiphytic plant in most of the primitive native forests. The length of4

The Crane-Flies of New York— Part II

the fully grown larva is slightly over 30 millimeters.

935

It is subcylindrical,

Oval warts armed with minute teeth are situatedon both surfaces of abdominal segments 2 to 7. The larva is very dark
The spiracular disk
slaty gray in color, darker toward the extremities.
Apparently only
as shown by Hudson's colored figure is very small.
one larva inhabits the space between the two sheathing leaves of theAstelia, and only those leaves which are full of a thick, brown, coffeeconsiderably flattened.

likeliquidare

The pupa

frequented.

isinclosedina rather tough,extremely elongate, silken tube situated between the sheathing leaves.
The
It rests in an upright position in the midst of the semi-liquid mass.

pupa measures about 38 millimeters in length, being very elongate with
The two pronotal breathing
the head and the thorax unusually small.
horns are shaped somewhat like a bivalve shell. Abdominal segments
3 to 6 at the base on the dorsal side have finely-toothed warts; the ventral
surface has plain ridges.

It isprobable that the present species is not atrue Gnophomj'ia, but until more is known of this species and its relatives
itshould be referred to this genus.

The Gnophomyia pilipes referred to by Beling (1879:42) and by Gerbig(1913:161-163) pertains to Trimicra (page 932).
G. tristissi7n.ah.SiS been recorded by Malloch (1915-17b: 230-231) as livinginwet mud, but this is an error.

The writer has material from exactly

the same source as Malloch's, received from James A. Hyslop, and this—shows that the haunt of the larvae is beneath the decaying bark of trees
an unusual habitat for one of the Eriopterini, which for the most partThe rearing of this species inlive in damp sand or earth near water.
New York (by Young), Massachusetts (by Johnson), Maryland (byHyslop), Virginia (by Shannon), Kansas (by Alexander), Illinois (byMalloch and Alexander), and Texas (by Mitchell), leaves no questionthat the immature stages are to be found beneath the decaying bark of
the larger hardwood trees, the tuUp tree, Liriodendron Tulipifera Linn.,being often preferred.

CnophoTnyia iristissima O.
1859

S.

Gnophomyia tristissima 0. S.

Gnophomyia
vellow halteres.

tristissimaisProc. Acad. Nat. Sci. Phila., p. 224.

an interesting

1)1

ack

fly

with conspicuous

It is common and widely distril)uted thruout the eastern

Charles Paul Alexander

936

United States and Canada. As stated a])ove, the species has been rearedon at least seven different occasions in as many States, the most complete account being that by Hyslop, whose specimens and manuscript
notes were kindly placed at the writer's disposal. His data on this species
are as follows:

May 14, 1914. Wolfsville, Maryland. l^ndiT the b;irk of a rotten stump of a tulip
poplar (Liriodendron) on the roadside near Wiirrenfeltz .schoolhouso, field on the left going
I found a great number of amber-yellow dipterous hirvae (three in alcohol), andto town.
also three pupae slightly shortened and with the thorax and lc>!?;s»ferruginous and the abdomen pale amber. They were in a very moist nidus of rotted inner bark; placed in rearing ina tin box.
May IG. One adult emerged today (pinned); pupal case in alcohol.
May 18. Three adults emerged today (pinned); pupal cases in alcohol. Observed
emergence of one adult. The swaying motion observed in Tipula infuscata was not observed,
but the adult simply glided straight out of the pupal case by a wavelike contraction of theabdomen. The whole emergence took only about eight seconds. A larva pupated.
May 23. Adult emerged. Pupal stage five days. Placed the remainder of the pupae
The pupae are quite active and move(r41 had transformed from larvae to pupae) in alcohol.
under a shelter if exposed. Just before emerging, the pupa takes on a black color on thethorax and smoky yellow on the abdomen.
Shannon's material was reared from pupae taken under the bark of adead tulip tree at Dead Run, Fairfax County, Virginia, on May 5,
A larva that is undoubtedly this same species was found beneath1913.
the same tree on April 17, 1913. Johnson bred this species from larvae
found beneath bark at Riverside, Massachusetts, on April 24, 1905.
The Texas specimens were found l^y Mitchell beneath the bark of cottonwood (Populus) at Victoria, on June 30.
At Lawrence, Kansas, in 1919, the writer found a fow larvae underthe bark of a box elder, associated with the following dipterous larvae:
Pterocalla strigula Loew, Lonchaea laticornis Meig., Phaonia harti Mall.

These associated species were kindly determined by Mr. Malloch. In
1920 the flies were bred from under the bark of several deciduous trees at
Urbana, Illinois, by Mr. Malloch and the writer.
Well-preserved specimens of the larvae are not available to the writer,
and the following description is taken direct from Malloch (1915-17 b:
230-231)—

Larva.
Length, 9-11 mm. Slender, slightly tapering toward both extremities, more
Body yellowish testaceous, covered with dense decumbent
decidedly towards the cephalic.
pile.

Head [Plate LXXIV, 39.1] more compact than that of Helobia, the lateral rods stouter;antennae very small; maxillie large, produced beyond the apex of the narrow labrum, the palpi
stout labium not chitinized; mandibles slender, with a long sharp apical tooth and about three
Locomotor organs consisting of rather
poorly defined teeth along the lower lateral margin.
;

The Crane-Flies of New York— Part II

937

broad fusiform areas on anterior portion of abdominal segments except basal and apical;
hairs alono; mirgins of segmental incisions more distinct than elsewhere because of theirb^in"; sli'^htly curved upward; apical segment with five processes, their structure and markings as in figure [Plite LXXIV, 397]; anal ventral blood-gills in the form of four shortrouided protuberances.

The pupa is described from four
Mr. Hyslop:
Pupa.-

castskins,

kindly presentedby— Length, 8-10 mm.

Cephalic crest a low, blunt tubercle on either side of median line, each tipped with a long,stout seta.
lobes.

Labrum broad, elongate, obtuse at tip, completely separating triangular labial

Sheaths of maxillary palpi moderately slender, tapering gradually to tip.

Antennal

sheaths moderately elongated, extending to about opposite wing root; basal segments angulate.

Pronotal breathing horns (Plate LXXIV, 399 and 400) small, trumpet-shaped, very flattened,
median area lacking and hence the margins contiguous; just proximad of breathing hornsa large, roughly triangular lobe on either side, immediately behind which are two small

Mesonotum moderately declivitous, at crest rather tumid, but unarmed, with afew parallel grooves on either side of median line; lateral angle of thorax very sharp, before

setae.

two setae, one very powerful, the more dorsal one abortive; a strong seta on either
mesonotum behind crest. Wing sheaths short, ending just before tip of second
abdominal segment. Leg sheaths (Plate LXXIV, 398) very short, ending just opposite tip
tip withside ofofsecond abdominal segment and thus projecting but slightly beyond wing tips; hind legs

a little the shortest, but no striking difference in length of various sheaths.

Abdominal segments subdivided into a narrow basal ring and a much broader posterior
ring; abdominal segments on dorsum with four transverse rows of tiny but stout setae,two

on basal ring and two on posterior ring, one being subbasal, the other subterminal, in position;

stsrnum with only the subterminal row of setae present, but this well marked, the otherrows merely vestigial; on either side of dorsum, just cephalad of ends of subterminal row
of setae, a powerful bristle; in alinement with theseand subequally spaced, two smaller

setae; a strong pleural seta on a raised papilla opposite basal ring and three opposite posterior
ring, the two anterior being larger, the posterior one very small; tsternum with a strong setaon extreme lateral margin of posterior ring; lateral spiracles distinct, on segments 2 to 7.
Male Cauda (Plate LXXR', 401) with ventral lobes small, blunt, rather widely separated
basally but converging apically; dorsal lobes powerful, divergent, and rather acute at tips;
at base of each near lateral margin a shortbi.'id knob sending one arm dorsad, the otheron cephalic angle a short, stout seta; two long, powerful, lateral setae on
either side, and a single powerful seta on either side of dorsum, immediately behind which
is a blunt tubercle.
Female cauda with sternal valves elongate, powerful, at their tips terminating in slender, divergent points; dorsal valves small, blunt, divergent, located at baselaterad; at baseof sternal valves, at their tips

with a short, slender spine directed backward; base of seg-ment about as in male.
iVeanoOype.— Wolfsville, Maryland, May 10, 1913. No. 234.
Two male and one female pupae from type locality,

Paratypes.

—

May 18, 1913.

Charles Paul Alexander

938

Genus Qonomyia Meigen (Gr. angle + fly)
1818
185G
1869
Larva.

Gonomyia Meig. Svst. Beschr. Zweifl. Ins., vol. 1, p. 146.
Taphrosia Rond. Dipt. Ital. Prodr., vol. 1, p. 182.
Goniomyia O. S. Mon. Dipt. N. Amer., part 4, p. 176.

— Form elongate,

terete.

Spiracular disk surrounded by five blunt lobes which areheavily marked with brown, in some species (G. alexanderi) the brown suffusing the disk

between the

spiracles.

Head capsuleof eriopterine type.

IMandible withlateralteeth

Antenna with apical papilla elongate-oval. Alentum not chitinized.
Cephalic crest blunt, the surface with minute roughenings. Pronotal breathing

slender, flattened.

Pupa.-—

horns flattened, fanlike (Gr. sulphurella), or short, trumpet-shaped.

Mesonotum declivitous,

at crest with an interrupted transverse row of six to eight tubercles which are densely besetwith sharp black spicules.

Wing sheaths attaining base of third abdominal segment.

Legsheaths moderately elongated, reaching base of fourth abdominal segment; tips of middle
tarsi ending a short distance before apices of other tarsi.

weak.

Lateral spiracles distinct, tubular.

Armature of abdominal segments

Five blunt, fleshy lobes on dorsum of eighth

abdominal segment.

Gonomyia is a lar^c and diverso gsnus of small crane-flics (including
more than one iiundrcxl known species) described from all parts of theworld.
They are divided into four recent subgenera, of which three
Gonomyia Meig., Progonomyia (new name for Gonomyella Alex., pre-—occupied), and Leiponeura Skuse— occurin the Nearctic fauna.

The immaturo stages of the known species arc spent in moist sand or
earth, usually near water.

In Europe, G. tenclla Meig. (Beling, 1879:56,

mention only) was found in August in damp, sandy earth along the marginof a dried-up brook.

The writer has found the immature stages of Gormmyia {Leiponeura)and G. (G.) kansensis in wet sand near rivers. G. sulphurcUa
and G. suhcinerea 0. S. occur in muddier and more stagnant conditions
near ponds and small streams.
Not enough larvae arc availal)lc for study to require a key at thistime. The pupae of the known Nearctic species may be distinguished byalexanderi

the following kc}':
1.

2.

Pronotal breathing horns narrow at base, expanded distr.lly into a very flattened, fanlike blade with delicate and anastomoi^in.-? narvures
G. sulphurella O. S. (p. £40)
2
Pronotal breathing horns not as above, more earlike or trumpet-shaped
Pronotal breathing horns massive, trumpet-shaped; l^tsril margin of thorax before
wing root produced into an angle; mile cj.uda small, elongate, dorsal lobes a little
shortfir than ventral lobes, with two stout lobes on dorsil side fir removed from their
G. alexanderi (Johns.) (p. 939)
b la'!
Pronotal breathing horns flattened, earliko or narrowly truimet-s'iaoeil; literal marginsof thorax above wing root broad and blunt; male c luda s'lort, stout, dorsil and ventral
lobes subequal in length, the latter closely approximated along median line, the former
G. kansensis Alex. (p. 941)
widely separated, at their base with two acute points

The Crane-Flies of New York— Part939

II(Subgenus Leiponeura Skuse)
1889
1915

Leiponeura Skuse.
Lipophleps Bergr.

Proc. Linn. Soc. N. S. Wales, set. 2, vol. 4, p. 795.

Psyche, vol. 22, p. 55.

Gonomyia {Leiponeura) alexanderi
1912

Elliptera alexanderi Johns.

(Johns.)Psyche, vol. 19, p. 3.

The beautiful crane-fly Gonomyia alexanderi is locally common in theeastern United States. The adult flies may be swept from rank vegetation
When resting, the adults have a charin the neighborhood of streams.
fore
legs
st'anding straight ahead and almost
acteristic position, theextendedlaterally and slightly forward, theparallel, the middle legs
butwidelydivergent, and the wings folded
backward
hind legs directed
This is the characteristic resting position for the genus.
The larvae were found in some numbers in rather coarse sand, aroundsmall pools of water near the Sacandaga River, Fulton County, New York,over the back.

on June 5, 1914. The adults emerged on June IG, giving a pupal period
The descriptionof not more than eleven days and presumably much less.
and figures of the pupa are made from the cast pupal skin of the male.
Larva.

— Length, 8.3 mm.
Diameter, 0.4-0.5 mm.

Coloration very pale yellow or yellowish white.

Form terete, elongated, slender.

Body with a sparse, pale pubescence, at posterior margins

of segments with a transverse erect ridge of stiff hairs.
large,

flattened,almostSpiracular disk (Plate LXXV, 403)

pentagonal in outline, surrounded by five lobes; dorso-medi ai

lobe small, slender; paired lobes very short and blunt; margin between lobes almost straightor but feebly concave;when disk is partly closed, lobes appearing a little more prominent;

ventral lobes a little larger than lateral lobes; lobes heavily suffused with brown; on ventral
lobes a lateral dark brown hne running dorsad to near spiracles, at its dorsal end connectedacross disk by a paler brown suffusion; proximal stripes of ventral lobes shorter and paler,

above their inner ends with a small brown spot; lateral lobes almost entirely suffused withbrown, this entirely surrounding spiracles and in some specimens entirely suffusing disk

betweenspiracles,

this

mark

bifid at its distal end; dorsal lobe indistinctly marked

withvery pale brown; disk margincl with short, pale hairs which are not interrupted r,nd are
only alittlelongerat

tips

of lobes.

Spiracles widely

separated, the distance

betweenthem being three or four times diameter of one; spiracles yellow, centers pale brown.

Head capsule as in the tribe, the ventral bars broader than the slender dorsal bars, theirinner ends not expanded or toothed to form the mental plate.
ately elongate, densely hairy.

Labrum-epipharynx moder-

Men turn not chitinized; hypopharyngeal region a cushion,

covered with delicate, short setae.

Antenna as in this tribe, basal segment moderately elonMandible (Plate LXXV, 402)

gate, densely hairj', apical papilla rather small, elongate-oval.

Charles Paul Alexander

940

moderately large; apical point elongated, slender; ventral cutting edge with three flattened
teeth, gradually smallerfrom outermost toward base; outermost tooth

flattened, a little

enlarged distally, about as long as apical point; basal tooth small, acute; a prosthecal append-

age with a brush of hairs beneath it.
Pupa.'— Length of cast pupal skin, about 5.5 mm.

Cephalic crest of two prominent lobes, blunt at
lobes blunt at tips.
apices.

tips,

their surface granulated.

LabialSheaths of maxillary palpi moderately stout, tapering suddenly to sharp

Antenna with basal segment very angulated.

Pronotal breathing horns (PlateLXXVI, 408 and 409) massive, short, trumpet-shaped, flattened laterally, and here marginedwith an elevated ridge, along which are scattered the rows of breathing pores; mouth of thistrumpet wide. On thorax between breathing horns, large, rounded lobes which are minutely
granulated.

Declivity ofmesonotum(PlateI.XXVI, 407) somewhat precipitous,

at therather narrow crest with about six small tubercles which are densely beset with spicules;

along shoulder a similar, but more elongate, transverse welt; lateral margin of thorax before

wing root projecting out as a sharp angle with a seta at its base.

Wing sheaths reachingend of second abdominal segment. Leg sheaths moderately long, attaining base of fourth
abdominal segment; fore legs a very little longer than hind legs; middle legs much shorter,
ending just beyond base of last tarsal segment of fore legs.
Abdominal segments divided into two narrow basal rings and a much broader posterior
annulus. Armature of abdomen very weak. Male cauda (Plate LXXVI, 410 and 411) small,elongate; ventral lobes a Httle longer than the short, blunt dorsal lobes; on dorsal face nearend of eighth segment, two stout lobes pointed at the tips which are directed dorsad andslightlycaudad; eighth segment -ndth a close pentagon of pale, slender lobes, the posterior

pair larger and closer together than the anterior pair, the median lobe the smallest.

— Sacandaga River, Fulton County, New York, June
— With type
Paratypes. — Two larvae with type.
Ncpionotype.
Nernoti/pe.

5,

1914.

larva.

(Subgenus Gonomyia Meigen)

Gonomyia {Gonomyia) sulphurella O. S.
1859
1869

Gonomyia sulphurella O. S. Proc. Acad. Nat. Sci. Phila., p. 230.
Goniomyia sulphurella O. S. Mon. Dipt. N. Amer., part 4, p. 180-18L

Gonomyia sulphurellaisa handsome

littlecrane-fly

which

is

very

common and widely distributed thruout the eastern and central UnitedStates.

Larvae are not infrequent in mud along the banks of streams.

The writer has bred this species from larvae sifted from sandy mud fromthe banks of Cascadilla Pond,

Ithaca,New York,

where they occur

associated with numerous larvae and pupae of a tabanid (Chrysops itidiis(). S.), a stratiom\iid {Odontomyia sp.), and other forms.
Larvae collected
on May 14, 1913, emerged as adults on June 1. Adults have been reared

as late as October 19 by E. A. Richmond.

The Crane-Flies of New York
Pupa.

— Lengthof cast pupal skin,

Cephalic crest small, blunt.
subcircular, outer941

about 0.5-7 mm.

Labrum triangular, apex obtusely rounded.

margin rounded.

Labial lobes

Sheaths of maxillary palpi short and stout, at apex

LXXVII, 414).

suddenly narrowed (Plate— Part IIorgan extending to beyond base of wing.

Antennal sheaths angulated at segments, thePronotal breathing horns (PlateLXXVII, 413^

with extreme base expanded, the neck short, constricted, soon passing into a greatly
expanded and very compresszd disk, the whole suggesting a fan or certain polypores; marginof this disk entire or gently crenulated, and sloping from ventral side outward; surface finely

nerved and reticulated.

Mesonotum somewhat precipitous, crest (Plate LXXVII,

tumid, v/ith rounded knobs arranged transversely along

it,

412)

there being about eight isolated

knobs and a more elongate one along shoulders, these knobs covered with minute blackened
Lateral margin of thorax above wing root forming almost a right angle. Wing

spicules.

sheaths ending opposite base of second abdominal segment.

Leg sheaths comparatively

short, ending opposite base of fourth abdominal segment; fore legs a little the longest, middle
legs conspicuously shorter

than the others.

Abdominal segments divided into two narrow basal rings and a broad posterior annulus.
Armature of abdominal segments weak, posterior ring with a narrow row of small blackspines before caudal margin; on basal ring a broad band of microscopic scabrous points
arranged in interrupted transverse rows, there being about seven or eight of these rows
to a band.
Spiracles small but distinct.
Female cauda with tergal valves short, but little
longer than sternal valves, slightly upturned, ending in short, cylindrical tips; near basewith a small, blunt tubercle on either side; dorsum of segment 8 with five rather long, palelobes, the anterior pair more slender and more widely separated than the posterior pair, which

are sometimes closely approximated.

— Cast pupal
Ithaca, New York, October— Three pupae with type.

Neanotype.
Paratypes.

skin,19, 1915.

Gonomyia (Gonomyia) kansensis Alex.
1918

Gonomyia kansensis Alex.

Gonomyia kansensisisCan. Ent., vol. 50, p. 158-160.

a prairie species of the cognatella group andappears to be common along the Arkansas and Kaw Rivers in Kansas.
Living pupae were found at Larned on August

1,

1917, in sand along

the banks of the Arkansas River, where they were associated with thetypical sand-loving fauna, including Gelastocoridae, Saldidae, and Carab-

The observation
pupa of this fly is discussed on pageidae (Omophron, Dyschirius, Bembidion, and Tachys).
of a small ant preying on a living

729.

— Length of cast pupal skin, 6 mm.

Pupa.-

Similar to G. alexanderi in mort essentials but showing the following differencea: antennal sheaths strongly angulate at -segments; a tubercle on ventral face of antenna at base, andanother at inner cephalic margin of eye.

Charles Paul Alexander

942

Pronotal breathing horns (Plate LXXVII, 415 and 410) flattened, earlike or very narrowly

trumpet-shaped, with a thick marginal ridge, the disk restricted.

Lateral angle of thorax

above wing root very broad and blunt, setiferous. Leg sheaths of fore and hind legs almost
on a level, those of middle legs shorter, ending a little beyond midlength of last tarsal segment
Male cauda (Plate LXXVII, 417 and 418) with dorsal and ventral lobes veryof fore legs.
short and blunt, subequal in length; ventral lobes closely approximated on median line;
dorsal lobes widely separated at their base, with

two small acute points directed stro: gly

dorsad, divergent at their tips, each with two small setae on outer face near base; dorsum ofsegment 8 with five lobes, the anterior pair a little more widely separated than the posterior
pair, the median lolje slender.
Neanotype.

— Larned, Kan.sas, August

1,

1917.

Genus Rhabdomastix Skuse (Gr. rod + lohip)
1889

Rhabdomastix Skuse.

Proc. Linn. Sac. N. S. Wales, ser. 2, vol 4, p. 828-829.

The genus Rhabdomastix

includes nearly a dozen species, some ofwhich have been previously described as Gonomyiae.
The immature stages of Rhabdomastix schistacea (Schum.) were foundby Beling (1886:195) in wet earth beside a stream in beech woods on
May 6. The larva measures 6 millimeters in length; the greatest diameter
The body is strongly dilated in the anterior part andis 0.8 millimeter.
The integument is deep brownish yellow.
gradually narrowed behind.
The spiracular disk is short and blunt, and has four very small, tuberculate
teeth, the lateral pair lying somewhat more cephalad than the more
powerful ventral pair; spiracles small, circular, yellowish brown, separated
by a distance about equal to four times the diameter of one. The pupa
has the mesonotal declivity provided with an interrupted crossrow ofsmall, unequal, chitinized teeth.

(Subgenus Sacandaga Alexander)
1911

Sacandaga Alex.

Ent. News, vol. 22, p. 349-352.

Rhabdomastix (Sacandaga) Jlava (Alex.)
1911

Sacandaga flava Alex.

Rhahdomasiix flava

isEnt. News, vol. 22, p. 351-352.

a curious

fly

which

is

apparently related toGonomyia l)ut represents a quite different offshoot of the Eriopterini.
The writer believes that the larvae might be found in moist earth along
streams, but at present they are quite unknown.
The following notes

The Crane-Flies of New York— Part II

943

on the swarming habits of this species have been pubHshed (Alexander,
1912 a: 72-73):
On June 13, 1000, T found the species swarminj? [on Sport Island, in the Sacandaga River,
The species came out at about 7.45

New York] and made the following observations.

p.m. and at 7.51 p.m. began its flight in under an elm tree at the northeast end of the
The flight was generally forward, but continually from side to side for a few inches.
island.
The flight was quite irregular, always toward the slight north breeze. The whole swarm
would often move away and return, a little later, to the first place. It swarmed within fourThe
feet of tlu^ ground, generally much lower, averaging, perhaps, two feet.
number of individuals participating in the swarm was about twenty. Other species swarming nearby at the same time were Chironormis hyperhnreua, var. meridional is, Joh., and the
may-flies, Ephemerella excrucians Walsh, and Siphlonisca aerodromia Ndm.
.

.

.

Genus Trentepohlia Bigot (named after J. J. Trentepohl)
1854

Trentepohlia Bi'jot.

1011

Mongomioides Brun.
Mongomella Enderl.

1912

Ann. Soc. Ent. France, p. 474.
Rec. Indian Mus., vol. 6, p. 296.
Zool. Jahrb., vol. 32, part 1, p. Gl.

Trentepohlia is a tropicopolitan genus including about fifty-five described
species arranged in six subgenera

Brun.,— Trentepohlia Bigot, Anchimongoma

Mongoma Westw., Plesiomongoma Brun., Paramongoma Brun.,and Neomongoraa Alex.

Of these species, fifteen are American and theremainder are Old World forms. They are almost all species of delicate,
It is an interesting fact that
ethereal structure, with long, slender legs.

{Mongoma) pennrpes has been observed by Jacobson (DeEdwards, 1912-13:211) to form chains on horizontal spider webs, as is noted herein for Thrj^pticomyia (page 712) andsomewhat similarly for Oropeza (page 982). H. K. Munro has supplied (in litt.) the following interesting notes on the habits of Trentepohlia
Trentepohlia

Meijere, 1911:50, and(Trentepohlia) humeralis Alex, as observed in eastern Transvaal at theend of April, 1920:
Very inconspicuous when flying and resting. When flying resembles very much a small
Invariably settles on under side of twigs, leaves, and similar objects.
When at rest the wings are
Usually found among bushy undergrowth, but also in grass.
folded along back; very often on settling the insect moves itself up and down in the manner
Slow flier.
of the long-legged harvest spiders (Phalangiidae).
piece of thistledown.

A fossil Trentepohlia, T. cruciferella (Ckll.), has been described from theGurnet Bay Oligocene (Cockerell, 1917b: 373-374). Observations on theimmature stages of three species are available.

Charles Paul Alexander

944(Subgenus Mongoma Westwood)
Trentepohlia
1887(Mongoma) pennipes (O. S.)

Mongoma pennipes O. S.

Berl. Ent. Zeit., vol. 31, part 2, p. 204.

The immature stages of Trentepohlia pennipes have been described by

De

Meijere(1911:50-51)

Semerang, Java,asin January,

follows:

Jacobson found the larvae at

1903, in decaying plant stems.

larva sent was 9 millimeters long and almost

1

The only

millimeter in diameter,of cylindrical form, only slightly narrowed behind and brownish in color.

The head capsule was entirely retracted.

The entire body was thickly

beset with fine, short, appressed hairs; in addition to these, on the ventral
side of each of the six intermediate segments were transverse swellings

where the hairs were shorter and even more numerous. Surrounding
the anus were four long, cylindrical, anal gills, each constricted in threeor four places; if bent forward, the anterior pair would reach the middleof the penultimate segment of the body, the posterior pair being somewhat
shorter.
The last segment of the body was truncated, the lower angles
being somewhat produced and provided with a few somewhat longer
hairs; the spiracles, situated in the upper part of the spiracular field,
were relatively small and somewhat elongated.
The pupa (Plate LXXVIII, 419) was about 9 millimeters long, elongate, of a yellowish brown color, the abdomen for the most part brighter.
The thorax was almost smooth, with only a few short, brownish yellow
bristles.
The abdomen, except on the anterior segments, was thickly set
with numerous tubercles. The apex of the abdomen had two short, thornlike projections, curved outwardly; beneath these were two shorter
tubercles, and four short tubercles formed a quadrangle on the dorsumof the last segment.

The pronotal breathing horns

consisted of flat-

tened, leaflike lobes, the upper surface of which was scaly.

(Subgenus Paramongoma Brunetti)
Trentepohlia (Paramongoma) hromeliadicola (Alex.)1912

Mongoma hromeliadicola Alex.

Ent. News, vol. 23, p. 415-417.

Trentepohlia hromeliadicola and T. leucoxena have a larval habitat
which has not been found elsewhere in the family. They live in thewater that gathers in the leaf axils of tropical bromeliaceous plants,k

The Crane-Flies of New Yorkr4

— PartII

945^
spending their immature stages in this habitat, where they are associatedwith a remarkable fauna of other organisms. The adult females havethe valves of the ovipositor grcuitly elongated, and the writer has sugi

gested elsewhere that thismay be an adaptation for laying the eggs in

was reared in Costa Rica by Picado,
whose important paper (Picado, 1913) on the bromeliaceous epiphytes
contains colored figures of the larva, the pupa, and the adult. From
this paper it is seen that the larva (page 356, figure A, and plate 13,
hal^itat.

this

T. hromeliaclicola

figure 4, of reference cited)is

not unlike that of T. pennipes, describedabove, the four prominent anal gills of T. bromeliadicola (Plate LXXVIII,
420, of this paper) being a notable feature incommon, altho here theconstrictions are very numerous, there being twenty-five or thirty shown

The pupa (Picado, 1913:357, fig. 51, and pi. 13, fig. 2)(Plate LXXVIII, 421, of this paper)in the figure.

has the pronotal breathing horns

approximated on the median line, and the sheaths of the ovipositor (PlateLXXVIII, 423) greatly elongated to contain the elongated terebra of theadult within.

According to Keilin (1913), the tegumentary glands of thislarva are a provision against drought, which is the great source of danger
to organisms living in this habitat.

Trentcpohlia {Paramoyigoma) leucoxena (Alex.)
1915

Mongoma leucoxena Alex.

Ent. News, vol. 26, p. 29-30.

was reared by Knabin Mexico, from larvae
found living in bromeliaceous plants quite as in the preceding species.

Trentepohlia leucoxena

Genus Teucholabis Osten Sacken (Gr. weapons +1859
Larva.'

setae.

Teucholabis 0. S.

forceps)

Proc. Acad. Nat, Sci. Phila., p. 222.

— Form elongate, slender, terete. Body practically destitute of pubescence andSpiracular disk surrounded by three very broad lobes, a flattened ventral lobe andtwo shorter lateral lobes at the base of which are the small black spiracles. Gills four,
bluntly rounded and developed for propulsion.
Head capsule of four elongate, slender rods
or plates, interno-lateral pair forked at about midlength.
Mandible rather small, with about
three blunt lateral teeth.
Antenna elongate, two-segmented.
Pupa.

— Cephaliccrest setiferous.

Pronotal breathing horns short, blunt, closely applied

Mesonotum precipitous, at crest with two powerful hooks and smaller serrated
near shoulder. Wing sheaths reaching end of second abdominal segment. Leg

to thorax.
platessheaths reaching end of fourth abdominal segment, middle tarsi the shortest.
with a transverse row of setae before ends of segments.

Abdomen

Charles Paul Alexander

946

Teucholabis is a rather extensive genus of small crane-flies (including

more than fifty described species) which find their center of distribution
A few species occur in Africa and thein the Tropics of the New World.
The genotype, Teucholabis complexa, is the only species
Oriental region.
that has been reared (Johnson, 1900). Johnson's material was kindly
sent to the writer for study, and furnishes the basis for the following
descriptions.

Teucholabis complexa 0. S.
1859

Teucholabis complexa O. S.

Proc. Acad. Nat. Sci. Phila., p. 223.

Larvae of Teucholabis complexa were found by Johnson in considerable
numbers beneath the bark of a decayed oak l^elow Avalon, New Jersey,
on June 8, 1899. They commenced pupating about the 13th, the imagines
continuing to emerge from the 22d to the 27th. This gives a pupal
duration of not more than nine days.
Larva.

— Length, 9 mm.
Diameter, 0.55-0.6 mm.

Coloration pale yellowish white, spiracles conspicuously darker.

Form long and slender, body terete, tapering abruptly to the small prothoracic segment
Sutures between segments indistinct. Body practically destitute of(Plate LXXIX, 424).

pubescence and setae.

Spiracular disk (Plate LXXIX, 427) with a broad, flattened, ventral

lobe, which is very bluntly rounded to subtruncate at apex, and two very short, blunt, laterallobes, at the base ofwhich are the spiracles; spiracular disk without distinct markings.

Spiracles small; middle piece black, ring pale horn-color; spiracles rather widely separated,the distance between them about equal to the long diameter of one.

Anal gills (Plate

LXXIX, 42<S) represented by four blunt, rounded lobes, which are apparently developed for
propulsion rather than for a respiratory function.

Head not easily distinguishable in material available for study.

Head capsule consistingof four long, slender, rodlike plates, the internal lateral pair forked atabout midlength,

so that capsule ends in six rods.

Epipharynx with numerous small spines.

Antenna (Plate

LXXIX, 426)

basal segment elongate-cylindrical, apical segment small,ovate.

two-segmented,

Mandible (Plate LXXIX, 425) rather small, apical point inconspicuous, with about
Lobes of the maxilla blunt, stout, hairy, not extending

three similar lateral teeth below it.
farbeyond tip of mandible.

Pupa.

— Length, 6.5-6.6 mm.
Width, d.-s., 1-1.1 mm.
Depth, d.-v., 1.2 mm.

Coloration pale; head, thorax, and sheaths of appendages darker; eyes black.

Form slender, narrowed behind.

Between antennal bases a prominent, two-parted crest,
Front above eyes

each lobe somewhat truncated behind and bearing a single stout seta.

The Crane-Flies of New Yorkslightly raised, two narrow lines on front,— Partmeeting below at a very acute angle.

in male, smaller in female, the front correspondingly" narrowed or broadened.

subtriangular.

947

II

Eyes large

Labrum short,

Sheaths of labial lobes small, suboval, separated by tip of labrum.

of maxillary palpi short, stout, straiglit.

Sheaths

Antennal sheaths moderately long, extending tobeyond wing base; basal segments prominent, indicated On sheath as prominent elePronotal breathing horns short, blunt, anterior face closely applied to pronotum,
Thorax very deep, precipitous, at crest
outer face free; at base a small rounded knob.
(Plate LXXIX, 429) armed with two strong curved hooks, one on either side of median line;
on shoulder laterad of these hooks, two prominent flattened plates whose margins areminutely serrated, the dorsal, or outer, plate being the larger and terminating in a large
curved hook. Wing sheaths moderately broad, ending just before tip of second abdominal
just

vations.

segment.

endingLeg sheaths (Plate LXXIX, 430) long and slender, outer pair much the longest,
opposite tip of fourth abdominal segment, middle pair the shortest.

about

Mesonotum with two strong setae.
Abdominal segments divided into two annuli by an indistinct suture, anterior ring narrow;anterior ring with a strong seta on pleural region; posterior ring with strong setae on dorsum

and sternum near caudal margin; three other setae on pleural region, two at about midlength
Male cauda (Plate LXXIX,
of segment, the third near base and more dorsal in position.
431) suddenly narrowed, terminating in two blunt ventral lobes and two acutely pointed
dorsal lobes bent strongly dorsad at their tips and bearing a short seta in notch on inner face
before apex; near base of cauda, on dorsum, a broad transverse swelling terminating in two
widely separated, slender tubercles, immediately cephalad and laterad of which is a long seta;
three strong setae on either side near base of cauda; posterior margin of segment 7 with two
powerful, decussate setae on dorsum, and between them two small setae; another powerfulseta near pleural region, and just inside still another smaller seta.

———

Nepionotype.
Avalon, New Jersey, June 8, 1899.
Neanotype.
With type, June 14, 1899.
Paratypes.
One larva and one pupa.

Genus Cladura Osten Sacken (Gr. branch + tail)1859

Cladura O. S.

Proc. Acad. Nat. Sci. PhUa., p. 229.

—

two from eastern North
The genus Cladura includes six known species
America, one from western North America, and three from Japan. The
small Cladura delicatula Alex., of the mountainous regions of the northeastern United States, differs from the genotype, C. flavoferruginea O. S.,
in several important respects

and it is necessary to erect a new group toThis group may, for the present at least, be considered as asubgenus of Cladura,' and the name Neocladura (Gr. new + Cladura) is

receive it.

proposed.

Moreover, the genera Crypteria Bergr. and Pterochionea Alex,are closely related to Cladura, and the entire group are almost certainly the
direct forbears of the. subapterous genus Chionea Dalman.

Neocladura

Charles Paul Alexander

948

curiously combines the structural characters of Cladura, Pterochionea,

and Crypteria. In the long basal fusion-segment of the antennal flagellum
and in the wing venation, it agrees with Pterochionea and, to a somewhat
In the structure of the male hypopygium,
lesser degree, with Cladura.
which has two slender pleural appendages, it departs widely from thetype of typical Cladura, Chionea, and Pterochionea, and agrees betterwith Crypteria, as well as with Conosia v. d. W. and Lecteria 0. S.
Because of this combination of characters, Neocladura must be separated
in some manner from Cladura in the strict sense.
It must be borne inmind, however, that the conspicuous difference in the structure of themale hypopygium probably has a phylogenetic significance, and the twoflies may not be so closely united as this arrangement would indicate.'''^The habits of the adult flies of Cladura and Neocladura are generally
similar.
Both species fly in late summer and in the autumn. They
often occur on dry, wooded hillsides remote from streams and otherbodies of water.

The habits of the adult flies of Cladura flavofcrruginea

have been discussed by the writer in an earlier paper (Alexander, 1910:250,
as C. indivisa).
His observations, made near Gloversville, New York, on
September 22, 1909, are as follows:
I went to Simmon's Woods, southeast of Gloversville, New York, this afternoon, andwas very agreeably surprised at the occurrence, in large numbers, of this usually uncommon
insect.
Near the entrance of the woods, where Simmon's Brook emerges, the insects were
found in numbers. At each step they flew out of the bushes to others farther away. They
are wary insects, and when sitting on the upper side of a leaf, slip over the edge and hanginverted from the lower side when alarmed by an observer.
They present a very characteristic attitude, sitting on the leaf of a tree, with their wings folded flat over the abdomen, andthe SIX long legs stretched out over the leaf. A few were taken in copulation; these werethe under side of a leaf.
Their habit of clinging to the under surface of a leaf
is quite remarkable and I found several by looking for them there.
There were hundreds of specimens in the low bushes of the woods, usually on the broad
Sometimes they would alightleaves of deciduous trees at a height of two or three feet.
on hemlock, and, occasionally, in ferns near the ground. It was the only tipulid observed
here to-day.
all hanging on

The eggs of C. delicatula have been taken from gravid females.

They

are comparatively few in number, but because of their unusual size theyalmost fill the entu^e abdominal cavity of the fly.

These large eggs areelongate-ovate in shape.

Cladura flavoferruginea O. S.
1859
1861

Cladura flavoferruginea 0. S.
Proc. Acad. Nat. Sci. Phila., pi. 4, fig. 34,
Cladura indivisa O. S. Proc. Acad. Nat. Sci. Phila., p. 291.

The Crane-Flies of New York— PartII

949

The immature stages of Cladura flavoferruginea were discovered while
thisA brief account of the larvae and pupae are

paper was in press.

given here in order to complete the data.

The larvae were found in Augurville Woods near Urbana, Illinois.
They occurred in soil which was baked hard and dry and which supported
scarcely any other insect life.
Associated with the larvae when first
discovered were larvae of a scarabaeid, Xylorydes satyrus; a tenebrionid,
Meracantha contrada; a few dipterous larvae of the genera Sciara and Psilocephala; millepedes of the genus Spirobolus; and a few less common formsof animal life.
A layer of dead leaves and other decaying vegetable mattercovered the surface, but this had not prevented an almost complete dryingout of the soil to a depth varying from six inches to more than a foot.
The only other tipulid larvae characteristic of such dry soil are species of
Dicranoptycha (page 828).
The most conspicuous features of the pupa are its cxceedingl}^ small
size as compared with the adult fly that emerges from it, and the entire
lack of protuberant pronotal breathing horns.
Larva.

— Length, 10-10.5 mm.
Diameter, 1.2 mm.

Coloration light yellow thruout.

Form comparatively short and stout.
pubescence; no distinct setae.

Integument provided with a delicate appressed

Basal annulus of abdominal segments 2 to 7 with a transverse

area of microscopic points arranged in long transverse rows; last ventral segment with a flat-

tened lobe covered with short setae, evidently an organ for shoving.

Spiracular disk entirely

without lobes, the spiracles being located on the exposed dorso-caudal surface of the lastHead capsule relatively compact; frontal plate broad, only slightly
Labrum quadrate, with conspicuous oval lateral arms. Antenna two-

abdominal segment.

narrowed behind.

segmented, terminal segment elongate-oval.

Mandibles of a herbivorous type, with an apicalpoint and two incomplete rows of teeth on inner, or cutting, face.

Mental bars widely

separated, each bar provided with two acute teeth at its proximal end.

Pupa.

— Length, 6.7 mm.
Width, 1.4 mm.
Depth, 1.4 mm.

Coloration pale yellow; head, thorax, and appendages darkening in age.

Cephalic crest gibbous, entire or feebly bifid, armed on either side with a single powerful

two bristles on vertex and two on front; labrum with a pair of small bristles at each
Labial lobes subquadrate, weakly .separated by apex of labral sheath.
Lateral margin of eye produced laterad into a
Palpal sheaths short and stout, straight.
Antennal sheaths extending to opposite one-third length of wing sheaths.
digitiform lobe.
Pronotum and mesonotum armed with
Pronotal breathing pores entirely sessile.
bristle;cephalic lateral angle.

C/ZARLES Paul Alexander

950

Wing sheaths ending opposite base of third abdominal segment.
bristles.
Leg sheaths unusually long, ending opposite base of sixth abdominal segment; hind legs theAbdominal tergites with ten strong bristles, eight being
longest, middle legs the shortest.
arranged in a single transverse row along posterior margin; abdominal pleurites with fourstrong bristles, one on each anterior ring, two near caudal margin of posterior ring, one
ventrad of spiracle, the last-named rudimentary, located on segments 2 to 7; sternites unarmed
conspicuouswith bristles.

Genus Chionea Dalman (Gr. snow)
1810
1912

Chionea Dalm. K. Vet. Akad. Hand!., vol. 1, p. 102.
Sph(eco7Wphilus Beck. Aan. Soc. Ent. Bolgique, vol. 56, p. 142.

Chionea is a small genus of nearly apterous crane-flies, found thruout
the North Temperate Zone.

Allof theknown

species(about eight)

have the wings reduced to mere vestiges, this being the only genus of
The South
considerable size in which all the species show this condition.
African genus Platylimnobia Alex., which shows a somewhat similar
condition of wing atrophy, is probably not very closely allied to Chionea.
The adult flies of Chionea are most often found walking awkwardly
over the snow in the late fall or early spring or during warm days in
winter.
A few instances, however, have been recorded in which specimens
were found with the temperature below freezing. At other seasons of
the year they may be found among fallen leaves, under moss and stones,in the nests of small mammals as Arvicola (Schmitz, 1914), or in deserted

subterranean wasps' nests (Schmitz, 1916, and Becker, 1912).

Interestingaccounts of the habits of the adult flies of the commonest local species,

Chionea valga Harris, are supplied by Ainslie (1906), Johnson (1907),
Lugger (1896) gives the following interest-

Washburn (1907), and others.

ing account of the habits of the same species:
As a general rule the wingless flies are found only early in the morning, though in one caseA few winters ago
a female was discovered crawling over the snow in the evening.
the writer discovered a female moving slowly over the snow and by searching he soon detected
a male. Putting both together imder an inverted glass the snow-flies immediately mated,notwithstanding it was several degrees below the freezing point. Soon afterward the female
found a crack in the glassy surface of the frozen enow, and forcing herself into it slowly
disappeared from view. Penetrating for some depth into the snow she deposited a number
Most females foimd seemed to have theof elongated eggs, which, however, did not hatch.
eggs already fully matured and only lacked to be fertilized.

...

Frauenfeld(Brauer,

Egger,and Frauenfeld, 1854:616) believed thatthe eggs are deposited in snow, since they are often laid in January or

February.

He thought the slimy substance secreted by the small lateral

The Crane-Flies of New York— Part951

IIvesicles of the receptaculum seminis of the female acts as a covering toprotect the eggs from wet and cold.

It

seems probable, however, thatthe flies enter some crevice in the snow around the base of trees or shrubs

and reach solid earth, at least in some cases.
the

flies

to be nocturnal in their habits.

Many authors have heldRecently,

Marchand(1917)

has furnished some interesting notes on an alpine Chionea, presumal)ly
His observations and experiments showed him that
C. alpina Bezzi,

Chionea was perfectly adapted to life on the snow, being attracted tomedium by its bright light and white color, the contact of the cold
surface on the feet resulting in a direct stimulus thru the claws.
The
insects drink water by pressing their proboscides against the snow.
Marchand considers the principal reason for these activities' being held on
the snow to be for the purpose of mating, since the insects can cover
thismuch more visible
The copulation of this crane-fly has been

considerable distances over the level surfaces and areto one another at this time.
fully describedby Mik (Osten Sacken, 1887:196) as follows:

The upper valves of the ovipositor prevent the male from getting on the back of the female;
it lies on its own back, in the direction of the longitudinal axis of the body of the female;when the latter is walking it drags the male, who raises himself on his hind legs to an almost
perpendicular position; this serves to explain the unusual incrassation cf these legs.

The genotype, C. araneoides Dalm., is the only species whose immature
stages have been described.

Chionea araneoides Dalm.
1816

Chionea araneoides Dalm.

K. Vet. Akad. Handl., vol. 1, p. 104.

The present knowledge of the life history of Chionea araneoides is due
almost entirely to the work of Brauer, Egger, and Frauenfeld (1854).
In February these investigators brought living adults, taken in copula,into an unheated room and placed them with their natural surroundings,

such as rich, damp, humous earth, rotten grape leaves, and similar substances,

in

glass

containers.

Aftera time a great

numberof small,elongated eggs of a hyaline appearance were noted, laid at random, somebeing deposited on the walls of the container, where they adhered but

soon shrunk and appeared dried out.
egg stage was not ascertained.

foimd in the soil.
color

Unfortunately the duration of theSome weeks later the young larvae were

They were of a cylindrical form, very pale yellow in
They were associated with numerous
active.

and not especiallyChakles Paul Alexander

952

larvae of Sciara longipes Meig.

Toward the end of May only a few larvae

grown but were not carried over into
were left, and these appeared
unknown.
which
is
still
the pupal condition,
fully

Larva.

— Length whengrown, 7.4 mm.

fully

Color light reddish yellow, contents of aUmentary tract showing thru body. Body terete(Plate LXXXII, 443), consisting of twelve segments, there being, besides the head, threethoracic and nine abdominal segments; last segment wedge-shaped (Plate LXXXII, 446

and 447), obhquely truncated so that the surface slopes from behind upward. Two spirMouth
acles on this oblique surface, with a paler brown, pincer-shaped mark between.
Mandible (Plate LXXXII, 444), produced into a long apicalparts powerfully constructed.
point; near midlength on inner edge of mandible a deep incision, distad of this about fiveteeth, basad of it three teeth.

Labrum quadrangular, with two lateral points on anterior part.

Mentum projecting outward as a cuticular rounded lobe. (Brauer shows two other appendages which are toothed on the outer face [Plate LXXXII, 445, of this paper]; these probably
represent part of thementum, but from Brauer 's figures they would appear to lie above

the labrum.)

Subtribe Elephantomyaria

Genus Elephantomyia Osten Sacken
1859
Larva.

Elepha7doniyia 0. S.

— Formterete.

(Gr. elephant+ fly)

Proc. Acad. Nat. Sci. Phila., p. 220.

Segments of body just before sutures with transverse rows of stiff

hairs; abdominal segments 5 to 8 on ventral surface with a mouthlike depression surroundedby long,

stiff

hairs.

Spiracular disk surrounded by four lobes, ventral pair the longest,

each of the latter bearing at its tip a single very long bristle. Spiracles moderately large,
located at base of lateral lobes. Head capsule very long and narrow, the four plates very

Mandible very small. Maxillary palpi short. Esophageal region surroundedby chitinized plates which are conspicuously obliquely ridged. Body of larva covered withnumerous long, appretsed hairs, producing a satinj^ appearance. Coloration saturated
elongated.

golden yellow.

Pupa.— Form slender.

A small crest on vertex above eyes.

narrowly separated on frontal and vertical regions.
recurved against it.
tubercles.

Eyes very large, globular,

Rostral sheath very elongated; palpi

Antennal sheaths lying across eye.

Head and thorax with setiferous

Abdomen with rudimentary lateral spiracles.

Elephantomyia is a small genus of crane-flies including about a dozen
These are found in widely separated regions

described living species.

of the world, there being about four in the East Indies, five in Africa,

and two in North America, one of which has been recorded also from
Europe. The genus is found fossil in the Baltic amber. The adultflies are remarkable for their very elongated rostra, and undoubtedly
they feed on the nectar of tubular flowers as in the related genus Toxor-

The Crane-Flies of New York— Part953

IIwhat species ofIhedecaying wood

No records are available, however, to indicate

hina

The

plants are thus frequented.

larvae live in

Elephanto-

known is the genotype
only species whose immature stages are
immature stages of this group
theon
myia westwoodi. The literaturerecord being the unknown Limnobnneof flies is very limited, the only
a good description ot
No 1 of Malloch (1915-17 b:235-236), who gives
described and figured for the first time.
the larva.

The pupa is here

Elephantomyia westwoodi 0.
1869

S.

Elephantomyia westwoodi 0. S.

Elephantomyia westwoodi

isMon. Dipt. N. Amer., part 4, p. 109.

a rathercommonfly

ineasternNorthwoods and may
are usually found in cool, shaded

The adults
America
The larva lives m deca>ang
localities.
be swept from vegetation in these
avaUable to the writer.
wood as is shown by the three records
in a very wet, rotten, willow log
Shannonbyfound
The larvae were
brewery at Rosslyn, Virgmia, on May
lying near the tidal flat above thethru the log.
About eight larvae were found, well scattered
21 1913and were very beautiful m
They were very active in their movements
placed myellow color. These larvae werelife being of a deep golden
indicating
27 and 29 and June 5 and 7,
rearing, and adults issued on May
Larvae were found m thxs
less.
slightlyorweek
a pupal duration of aand at that time they were ahnost grown,
log also on November 23, 1912,
This shows that the species spends
being about one-half inch in length.
larvae.
the winter as almost fully grown
Edge Hill, Pennsylvania, on JunenearlarvaeJohnson found these
Larvaejust beneath the bark.
25 1899 and on May 25, 1905, in a log,
preparationmaterial studied in theand pupae obtained by him were in the^of this paper.

j„+m a much decayed log at
Malloch found a single larva of this species(Malloch, 1915-17 b: 236).
White Heath, Illinois, on April 30, 1916,

i

Larva.— Length, 10-13.4 mm.

Diameter, 1.2-1.3 mm.

golden yellow thruout.
Coloration of body, a deep saturated
end
slender, tapermg gradually toward either
Body moderately elongated, terete, relatively432) the three horac.
(Plateend
anterior
towardabruptly
but more noticeabfy andsegmen^from the prothorac.c backward. Abdommal
segments gradually increasing in size
segments gradually shorter. Surremaiuing
longest,
the
fifth
the
2 short, the third to
1

LXXX

Ld

;

Charles Paul Alexander954

face of body densely covered with long, satiny, appressed hairs; thoracic and abdominal
segments with a dorsal and a ventral transverse ridge of short, stiff hairs just before caudal

margins of segments, these being longest on lateral parts of ridge; dorsal segments 6 and7 with incomplete transverse rows at about midlength; on ventral surface of abdominal seg-ments 1 to 3, near base, two transverse rows of tiny spines with a depressed area between;
segment 4 without this distinct double ridge; segments 5 to 7 with a very conspicuous
mouthlike depression at base of each, with a liplike margin on either side, the anterior margin
narrow with stiff hairs, the posterior margin swollen and densely set with tiny spines; on seg-ment 8 the transverse mouth lying near end of segment, its anterior lip with a dense fringe
Spiracular disk (Plate LXXX, 434) surrounded by fourof long hairs directed backward.
lobes; laterallobes the shortest, blunt at tips, inner face of each lobe slightly expanded

at tip, a few short silky hairs on outer face of lobe, inner face slightly chitinized; ventral lobes
longer, broad, tapering gradually to obtuse tips, notch between lobes deep, V-shaped, anarrow fringe of short, dense, golden, recurved hairs along outer face; at tip of each ventral

lobe a single.

powerfullong,

Spiracular disk unmarked.

bristle.

Spiracles

large,

very

widely separated, situated at base of lateral lobes.

Head capsule (Plate LXXX, 433) very small and narrow,being greatly elongated.

the plates, four in number,(The exact details are difficult to see in the material available for

Labrum broad, anterior margin evenly rounded, with a few long hairs.

EpipharynMandible very small, basenarrowed, tip produced into an acute point with smaller teeth at about midlength. Mental
region feebly chitinized.
Hypopharynx semicircular, rounded, anterior margin delicately
grooved. Esophageal region elongated, inclosed by chitinized plates provided with parallelridges running outward on one side and inward on opposite face, upon focusing producing a latticed appearance.
Maxilla densely hairy. Antenna two-segmented, conspicuous;basal segment short, apical segment larger, suboval. Sides of capsule on swelling behindstudy.)

region provided

geal

the maxilla with aandchitinizedatwith longsetae

directed backward.

brush of very long hairs.
tips

Plates of capsule elongated, expanded
and along margins; ventral plate near esophageal region withsetae.

Pwpa.

— Length, 8.4 mm.
Width, d.-s., 1.1 mm.
Depth, d.-v., 1.3 mm.

Coloration light yellowish brown; abdomen darker; thoracic dorsum, sheaths of legs,
and ovipositor more yellowsh. (In younger pupae the coloration is more uniformly
pale.)

Form slender, body narrow.

Eyes very large, rounded (Plate LXXXI, 441). Antennal
Vertex with a small but prominent crest lying
transversely above eyes; cephalad of this crest and just proximad of the antennal bases, a
setiferous tubercle.
Front between eyes moderately broad, produced caudad into the very
elongate rostral sheath, which is transversely wrinkled.
Sheaths of maxillary palpi recurved,

sheaths lying directly across face of eye.

lying alongside rostrum.

Sheaths of labium tiny, bilobed, lying at tip of rostrum.

Twolong setae on front between eyes.

Eyes very large, narrowly separated on dorsum of head,
the hinder part sunken under pronotum. Two conspicuous black tubercles behind antennalbases, each bearing a long, stout seta.

Pronotal breathing horns (Plate

LXXX, 435) small,The Crane-Flies of New York— PartII

955

short, clavate, yellow, constricted at bases, which are blackened; a prominent seta just above

each breathing horn; two setae on each side of pronotum beneath eye.

Mesonotal prescutumstrongly convex, with six strong setae on each side, one just cephalad of base of wing, anotherat joint of wing,

and two semitransverse groups in front of and behind level of wing base.

Wing sheaths short, ending some distance beyond tips of hind tibiae and just beyond baseof third

abdominal segment.

Leg sheaths (PlateLXXXI, 442) with fore femora strongly

swollen, lying alongside rostral sheath; tips of middle tibiae ending just beyond tips of fore

ending just before tip of fifth abdominal segment.
Abdominal segments densely and microscopically punctulate. Segments 2 to 7 with two
narrow basal rings and a broad posterior ring. Setae of dorsal abdominal segments (Platetibiae; legs very long,

LXXX, 437) en the posterior ring consisting of a caudal series of two strong outer setae andtwo smaller inner ones, the outer one of the inner series close to the proximal one of the outer
series; lateral series of setae

powerful; basal series in alineraent with anterior lateral seta,consisting of two powerful outer setae and a delicate inner one; pleural segments with a rudi-

mentary spiracle on segments 2 to 7, each spiracle with a stout seta above it; setae of ventral
segments (Plate LXXX, 436) with two strong bristles on each side near posterior lateral margin, the outermost with a tiny seta above it; a single basal seta located on a level with vestigial spiracles.
Female cauda (Plate LXXX, 439) with sternum of eighth segment having
two sharp, curved, widely separated spines, and just laterad of these a stout seta; pleuralregion with another seta on same level; sternal valves of ovipositor only a little shorter thantergal valves; dorsum of segment 8 with two long teeth, above which are two slender, divergent tubercles; tergal valves broad basally, narrowed suddenly at tip, on either side with asmall, subapical seta.
Male cauda (Plate LXXX, 438 and 440) with eighth sternite having a strong median tubercle that is two-toothed; laterad of this a strong chitinized tooth
bearing a powerful seta on outer face; a similar strong lateral seta; eighth tergite with fourstrong tubercles arranged to form a square; ninth sternite blunt, each lobe ending in two
small tubercles; ninth tergite ending in two divergent lobes bearing at tip a large and a small
seta and on dorsal face at about midlength another strong seta.
Nepionotype.

— Rosslyn, Virginia, May 21, 1913.

Neanotype.— Edge Hill, Pennsylvania, May 25, 1905.
Paratypes.
Larvae with type larva (two) and with type pupa (two).

—

Genus Toxorhina Loew (Gr. boiv + nose)
1835
1851
1869
1910

Limnobiorhynchus Westw. Ann. Soc. Ent. France, p. 683 (spurious name).
Toxorhina Loew. Linnaea Entomol., vol. 5, p. 400.
Toxorrhinn O. S. Mon. Dipt. N. Amer., part 4, p. 109-114.
Neoceraiocheiius Wesche.
Journ. Linn. Soc, Zool., vol. 30, p. 358.

Toxorhinaalmostallofa small genus including about nine described species,
which are from the New World. Two species occur inisAfrica and one in India.
Toxorhina madagascariensis Meun. is describedfrom African copal (Pleistocene). Nothing has been published concerning the immatui'e stages of any member of this genus.

Charles Paul Alexander

956

Toxorhina muliebris (O.
1865

S.)

Toxorrhina muliebris 0. S.

Proc. Ent. Soc. Phila., p. 233.

Toxorhina muliebris is the commonest species of the genus in the
United States, with a rather wide range thruout the Northeastern States.

The adult flies suck nectar from various flowers, such as the following:
Rhamnaceae, Ceanothus americanus Linn.

(Banks); Ericaceae,

Clethra(McAtee); Apocynaceae, Apocynum medium Greene(McAtee); Compositae, Solidago canadensis Linn. (Knab).
The immature stages are unknown, but from Mrs. Tothill's tent-trap
observations they are presumably spent in mud, since adult flies were
found in her traps set over wet, sedgy spots near Ithaca, New York. It
may be, however, that the insects live in fragments of decaying wood
which might be buried in this mud, since such a habitat conforms moreclosely to that of Elephantomyia, which is apparently closely related toalnifolia

Linn.

Toxorhina.

Eriopterine No. 1

A very curious larva, which has not been reared, has been found invarious places near Ithaca during the past few years.

It isa small, palelarva, very delicate and almost diaphanous in appearance, at the posterior

end with five flattened black plates with serrated margins, and with its
thoracic segments capable of considerable lateral extension.
The larva
but it introduces a type of spiracularis undoubtedly 'an eriopterine,
disk that has not been found elsewhere in the tribe.
The writer finds
it difl[icult to believe that this curious larva can belong to any of theeriopterine genera discussed in this paper, and yet there are very few
possibilities remaining; and one of these (Cryptolabis) does not occur
The genus Atarba, whosein the habitat frequented by this larva.
immature stages are still wholly unknown, is a possibility. Empeda, which
the writer considers to be a subgenus of Erioptera, has not been
If this is the larva
reared and must also be considered as a possibility.
of Empeda, the group at once assumes full generic rank as given it byOsten Sacken, but occupying an isolated position and no closer to Gonomyia than to Erioptera. The larvae of this species were found commonly
on Bool's hillside, at Ithaca, where they occurred in association withnumerous other crane-fly larvae discussed elsewhere (page 781). The

The Crane-Flies of New York— Part II

957

larva is described here in the hope that it will be reared and its identity
ascertained.
Larva.

— Length, 7-8.2 mm.
Diameter, 0.4-0.5 mm.

Coloration pale yellowish white; skin very delicate, almost diaphanous.

Body moderately elongated, terete; raeso- and metathoracic segments (Plate LXXV, 404)
and eighth abdominal segment capable of considerable expansion laterally, and, in death,
usually greatly swollen; last segment of body narrowed, cyhndrical, with a number of longsetae, including a group of five near base of lateral lobes.
inal

segments.

A few scattered setae along abdom-

Spiracular disk (Plate LXX\\405) surrounded by five equal elongate spatu-

late blades, these blades flattened, jet-black in color,margins

finely toothed; paired lobes

near base with a subhyaline median spot; margins of lobes (Plate LXXV, 406) with twentyfive to thirty hooks, recurved ones alternating with others laterally directed; when blades areclosed, these margins hooking closely together; at ends of blades and sparsely scattered along
margin, long, delicate setae; at apex of blades, two bristles; no spiracles found at base oflobes.

Anal gills four, lateral pair elongated, telescopic, inner pair shorter.

Head capsule much as in other eriopterine genera, especially Ormosia and Gonomyia,
dorsal plates slender, ventral bars a little longer.
Labrum as in the tribe; epipharyngeal
two smaller pads nearer base. IMentum notformed of ventral bars of capsule as in Molophilus. Hypopharynx a semicircular cushion
region with a large apical setiferous pad andcovered with long, dense setae, their tips a little recurved.

Antenna withapical papillavery long for this tribe, about equal in length to basal segment, cylindrical, with tip rounded.

Mandible moderately large, apical tooth not prominent, lateral teeth rather conspicuous,basal ones smaller but not so excessively reduced as in other members of the tribe; prosthecalarge.

(Described from larvae, Ithaca, New York, May 11, 1917.

No. 29-1917.)

Tribe Styringomyiini

The Styringomyiini comprise a small group of very peculiar crane-flies
witha tropicopolitan

There

only

the

singlegenus,

Styringomyia, with about twenty-five described species.

Most

of

distribution.

is

the

species are from tropical Africa and Asia, tho a few range into Australia

and the Hawaiian Islands, and one, Styringomyia americana
found in tropical South America.

Alex.,

is

Genus Styringomyia Loew (Gr. a kind of tree-gum -\- fly)

L

1845

1903
1912
1917

,'^tijringomyia Loew.

Dipt. Beitr., vol. 1, p. 6.
(Correctly Syringomjia
roth in litt.)
Idiophlebia Griinb.
Zool. Anzeig., vol. 26, p. 524-528.
Zool. Jahrb., vol. 32, part 1, p. 65.
Pycnocrepis Enderl.
Mesomyiles Ckll. Proc. U. S. Nat. Mas., vol. 52, p. 377.

— Berg-Charles Paul Alexander958

The history of the genus Styringomyia is remarkable. It was erected
by Loew in 1845, being based on the fossil species Styringomyia venusta
Locw, from African copal. Many years later it was found to be still
The earliest fossil records
living in the Tropics of both hemispheres.
pertain to the Oligocene of northern Europe (page 765).

The adult flies have such a curious structure that it seems best to
remove them from the former tribe Antochini, where they have long been
Concerning the first living species to be described, the Hawaiian
placed.
S. didy ma Grimsh., Perkins (1913:clxxxii) says:
It sometimes swarms at night aroimd the electric lights, sitting quietly on the walls andceilings, with the body pressed closely to the surface, and the front and middle lege extended

straight forward in front of the head in a characteristic manner.

Annandale has made similar observations on the resting positions of S.
He says (cited by Edwards, 1914-15 207)*'
This species rests on walls with the two anterior pairs of legs stretchedout straight in front and the posterior pair behind, resembling a stray
Jacobson has recorded much the same for S. jacobsoni
piece of cobweb."
Edw. (De Meijere, 1911:41^2, as S. didyma) in Java. The flies are
attracted to lamps and are almost always to be found in copulation, thehead of one directed away from the other; while thus engaged, sometimes one, sometimes the other, will run forward for a short distance,

ceylonica Edw., taken in India.

producing a peculiar appearance.
of S. vittata:

Munro {in litt) reports that the habits

Edw. as observed in eastern Transvaal in late April,

are very similar.

He writes: "Two specimens taken 'in cop.'

1920,

Settledon under side of a twig, heads in opposite directions, front legs of eachstretched out in front along twig, wings laid flat along abdomen."
Styringomyia didyma Grimsh.
1901

Styringomyia didyma Grimah.

Fauna Hawaiiensis, p. 10.

In Fauna Hawaiiensis, Perkins (1913:clxxxii) mentions the breeding

didyma by F. W. Terry. Before the reference could beby the writer, Mr. Terry died. The following letters from
0. H. Swezey in regard to the matter were then received.
In response to a letter sent on February 21, 1915, Mr. Swezey replied
on March 26 that " no information on the rearing of the species is available
in Mr. Terry's notes."of Styringomyia

investigatedThe Crane-Flies of New York— Part959

II

In a letter dated April 10 of the same year, however, the following
notes were enclosed:
About forty-five eggs deposited in tube December 5, 1910. Chorion jet black, shining andby 0.15 mm., very finely parallel-striate.
One hatched December 10. The batch was placed with rotten apples and cow manure,
Larva long, head small, mandibles distinct and well chitinized.
hatching December 15.
An adult male emerged about January 21, 1911.
thick, resisting dryness, 0.3 mm.

Mr. Swezey, thru whose kindness the above notes are available, adds:''You see from the notes of Terry's that he did not breed

didyma in its natural habitat.

Styringomyia

That is yet unknown, I guess."

The striking feature of this life history is its brevity, the entire egg,
and pupal stages being passed in about a month and a half. The
writer knows of no other crane-flies in which this is equaled, its nearest
larval,approach presumably being in the smaller Eriopterini.

SUBFAMILY Cylindrotominae

The subfamily Cylindrotominae constitutes a small, isolated group of
crane-flies,

with twenty described species arranged in seven recent genera.

All the species are Holarctic in their distribution with the exception offive speciesof theOriental genera

Brun., and Agastomyia de Meij.

Stibadocera Enderl.,

Stibadocerella

The group is a decadentone, having

been much better developed in the early and middle Tertiaries than atpresent (page 764).

The adult flies are sluggish in their habits, occurring on vegetation
shaded spots. The species of Cylindrotoma are brightly colored,

in cool,yellow and black, but the other forms are somber in appearance andblack or dark in color, the body being in some cases highly polished or
metallic.

The immature stages of the Cylindrotominae differ from thoseis known to the writer, in being spentof all other Tipulidae, so far as

on various bryophytic and spermatophytic plants, on the leaves of which

The larvae are usually bright green in color and suggest
Most of them simulate their hostplants to an astonishing degree.
The immature stages of Cylindrotoma
and Liograa are terrestrial, while those of Triogma and Phalacroccra are
the larvae feed.

a caterpillar in their general form.

aquatic or nearly so.

The genera
followsof the subfamily

Cylindrotominae

may be separated asCharles Paul Alexander

960

Larvae1.

2.

3.

Body appendages very long and filiform; aquatic

Phalacrocera Schin. (p. 961)
Bod}' appendages short, leaflike or tuberculate2
terminal
Dorsal appendages all simple, on theabdominal segments in a single row; terrestrial on spermatophytic plants
Cylindroloma Macq. (p. 966)
Dorsal appendages with teeth on anterior convex side3
Some of the dorsal appendages with three or four teeth on anterior face; aquatic on
Triogma Schin. (p. 973)
mosses
Dorsal appendages with one (L. nodicornis) or two (L. glabrata) teeth.
lAogma O. S. (p. 969)

Pupae
1.

2.

3.

Basal abdominal tergites without spines
2
Basal abdominal tergites with acute spines
3
Mesonotum unarmed; segments 6 and 8 each with two powerful dorsal hooks; segment
7 with a pair of strong ventral spines; pronotal breathing horns elongate, directedbackward
Phalacrocera Schin. (p. 961)
Mesonotum with two flattened erect lobei; sagmsnts G, 7, and 8 naked; pronotal breathinghorns small, directed slightly forward
Cylindrotoma Macq. (p. 966)

Abdominal spines branched

Liogma (glabrata) (p. r-69)
Triogma (irisidcata) (p. 974)
Liogma (nodicornis) (p. 971)

Abdominal spines not branched

The most important literature on the Cylindrotominae is as follows:
General account of subfamily. Osten Sacken, 1897; Alexander, 1914:105-106; Malloch,
1915-17 b: 210-211; Lenz, 1920 b: 113-115.
Larva, pupa, general.
Phalacrocera replicaia
De Geer, 1773; 1776: 135-141,r351.
.'
Larva
Grube, 1868.
Phalacrocera replicaia
Larva
Engel, 1884.
Phalacrocera replicaia
GeneralPhalacrocera replicaia
Giard, 1895 b.
Larva, general
Phalacrocera replicaia
Bengtdson, 1897. (Morphology of
.

.

.

.

.

larva.)

Larva(MorMiall and Shelford, 1897.
phology of larva and pupa.)
Bengtsson, 1899. (Morphology of

Larva

Holmgren, 1908.

Larva, pupa, general

Phalacrocera replicaia
Phalacrocera replicaia>

heart.)

Phalacrocera replicaia

Larva, pupa
Larva, pupa, general.
Larva, pupa, general.
Larva, general
Larva, pupa, general...
Larva, pupa, general
GeneralLarva, pupa, general.

Phalacrocera replicaia
Phalacrocera replicaia
Phalacrocera replicaia
Cij'indro'oma distinclissima
Cylindrotoma distinclissima..
Cylindrotoma disiinctissima ....
Cylindrotoma disiinctissima.
Cylindrotoma distinclissima
.

.

.

.

Cylindrotoma distinclissima.

.

.

.

.

.

.

.

Cylindrotoma disiinctissima ...
Cylindrotoma splendens

Liogma glabrata
Liogma glabrata

.

.

.

.

.

.

.

.

.

.

.

.

.

.

.

Larva
Larva, pupa, general ..
Larva, pupa, general.
Larva, general
General.

.

.

(Morphology ofhead.)
Griinberg, 1910:32-35.

Wesenberg-Lund, 1915:343-347.
Lenz, 1920b: 127-129.
Schellenberg, 1803:22-23.
Boie, 1838:234.
Zeller, 1842.
Schiner, 1864:563.

Kaltenbach, 1874:7.
Wesenberg-Lund, 1915:335 (as Triogma).
Lenz, 1920 b: 115-117.

Cameron, 1918.

De Rossi, 1876.
Osten Sacken, 1878 a.

The Crane-Flies of New YorkI^o,n.,Mrata961

l:™'^^r"':: ^f^S^S^^'1914 10r,-107.

Liogma glabrata
Liogma glahrata
Liogma glabrata..

Triogma tnsulcata

Generkl
general ..
Larva, pupa, gn
Larva, P^P^.
Larva generaf".^
Larva, P^Pf ^ g^'^^'"^
Larva, general.

Triogma

Larva, pupa, general..

Wvlpupa

Lwgma nod^corn^s
Tnpgma tnsulcata
Tnogma tnsulcnU^
trisulcata

Genus Phalacrocera Schiner
1863— Part II

Phalacrocera ScWn.

(Gr. hald

Alexande;,Lenz, 1920b:ll7-121.
Alexander, 1914: 107-115.
Steinmann. 1907-08.
:

'

.

im~m.
Wesenberg-Lund, 1915:347-348 (as^^^^^ glabrata).
Lenz, 1920b: 121-127.

+ horn)Wien. Ent. Monatschr., vol. 7, p. 224.
trachea-bearing filaments, the posterior
Spiracular d,sk w.th dorsal pa.r
others simple.

Larvae Body covered with numerous elongate,

thepair on dorsal segments deeply forked,
abdommalbranch of branched filaments of eighth
lobes formed bv rudimentary posteriorrfsegment.

Head capsule compact.

P^pa.- Cephalic

crest

low,

Mentum with about fifteen teeth.

not setiferous.

Pronotalbreathmg horns

almo tlong,

segments
tubercles, those of sixth and eighth
Dorsal abdominal segments withstraight.
stermte.
seventhon
tubercles
pointedenlarged into spinous hooks; twoofmedium-sized to large,

Phalacrocera is a small genus (four species)
Phalacrocera rephcata, is i^urodull-colored flies, of which the genotype,
the two remammg species are
P. mikado Alex, is Japanese, andpean,

North American.

u u+best-known,
common, thesphagnum bogs inbeing most frequently found in or neari

The adult flies of the American species are not

P tipuUna O. S.,

mountainous localities.

Needham (1908a:209) found the wings of an indi-

Sarracenia purpurea Unn., in thevidual of this species in the pitcher plant,of four specimens of LleAdirondack Mountains, together with the wings
Most of the specimens
insects.
phantomyia westwoodi and numerous otherwere taken in close proximity to bogs.
that have been found l^y the writer
have long been known,
The immature stages of Phalacrocera repUcata
workers on insect
described by De Geer and other early

having been
biology

More recently the hfe history, anatomy, andmorphology haveseveral other writers (page 960)^

been discussed in commendable detail byliving among submerged
Both the larvae and the pupae are aquatic,plants in quiet, but non-stagnant, water.
,
to be desired, as
The immature stages of P. tipuUna &re very muchorganization not found in the otherthe adult shows some features in its,

species of the genus.^
6

The larva of this species was discovered by J.

Speed Rogers in 1920.

•.

Charles Paul Alexander

962

Bengtsson (1897) erected for this genus the group Erucaeformia, which
he considered as the primitive form from which the Ncmatocera and theBrachycera have been derived.

This group, of course, has no standing

whatsoever,Phalacrocera replicata (Linn.)
Tipula replicata Linn. Fauna Suecica, 2d ed., p. 500-502.
Phalacrocera replicata Schin. Wien. Ent. Monatschr., vol. 7, p. 224.

1761
1863

The larvae and the pupae of Phalacrocera replicata have been discussed
by Miall and Shelf ord, by Bengtsson, by Holmgren, andby Wesenberg-Lund, that they are considered here only in general terms.
The habits of the immature stages have been discussed by many writers
since the time of De Geer.
They are oftentimes rather numerous amongin such detailaquatic plants such as Ranunculus fiuitans Lam., Fontinalis antipyretica
Linn., Hypnum elodes Schp.,

H. exannulatum Guenbel, and other species,
These moss fragments give a green tinge to young larvae when seen thru the nearly transparent body wall. Older larvae are more opaque and are brownish green
in color, indistinctly striped with pale and darker.
The larva is extremely
sluggish, remaining almost motionless for hours.
It clings to moss stems
by its large anal hooks, and, thus secured, it often sways its body fromfeeding on these mosses and probably on other plants.

side to side asif

to accelerate respiration.

periods of time without fresh
alive fortwo weeksair.

The larvae can go for long

Miall and Shelford kept specimens

in bottles completely filled with water,long time in water that had been boiled.

and for a

The larvae can live for a longtime out of water.

Progression thru the mats of submerged vegetationaccomphshed by grasping with the mandibles and the anal hooks,
alternately.
When alarmed the larvae curl into a rounded ball, after
the manner of many caterpillars. The skin, and more especially the long
body processes, are often covered with ectoparasitic organisms, suchas algae and infusoria, on which small fresh-water mollusks, Planorbis, have been observed feeding and creeping about over the body ofthe larva.
This coating of organisms, the body outgrowths, and the generaliscoloration of the larva, give it a striking resemblance to the mosses among

which

it

hves.

excessive cold.
winter,De Geer (1776:355) shows that the larva can endure
He placed four larvae in a vessel at the beginning ofand examined them in the following May.

During the winterr

The Crane-Flies of New York— Part II

963

had frozen into a soUd mass, yet,the water in which the larvae lived
Gcer found two of the larvae
on investigating the jar in the spring, De
both had pupated.
month
within astill ahve and able to feed, andvegetation flourishes
moss
a
which
habitat is in ponds in

The larval

A femalewhich currents keep the water in constant motion.
her eggs
depositing(1897:360)
Shelford
fly was observed by Miall andnuml^er,
in
sixty
about
eggs,
The
in the leaf axils of a submerged moss.
dark
opaque,
are
they
moss;
the
are laid singly and adhere slightly toandinirregularly

and spindle-shaped, with the surface of the chorion
one end. Bengtsson, Miiggenpitted, and with a rosette-hke micropyle athas but a single brood m aberg, and others believe that Phalacrocera
eleven months; Miall andabout
existence occupyingin color,

the larval
second brood. The egg
Shelford, however, admit the possibility of a
pupal duration is seventheandstage requires from eight to twelve days
to Miall and Shelford,
Accordingor eight days, according to Bengtsson.
the pupal period is considerably longer.
moltings, at leastDuring the larval development there are numerous
dorsal split
simple
a
by
cast
is
skin
eight and possibly ten; the old larval
emergmgafter
Just
segment.
extending from the first to the fourth
about
and
lengthinmillimeters
2.25
from the egg the larva is from 2 to
first
The
projections.
bodythe
0.5 millimeter in diameter, excluding
bodyThe
molting.
secondtheuntillarval stage (Bengtsson, 1897) lasts
delicate, threadlike, lateral projections,is provided with ten pairs of long,segments and arewhich are located on the second to the eleventh bodyolder larvae are
theofprojections
The otherhalf as long as the body.
yeari

placed immediately
merely indicated. The attachment apparatus ischitinized hooks,
twelvetoeight
before the anus, and consists of fromThe color ofcrossrow.
arcuated
directed forward and arranged in an
the mandishow
parts
mouth
The
almost transparent.

the body is white,
The second larval
horizontally.
bles without a prostheca and moving
body appendages
The
molting.
fourthstage lasts from the second until thein appearance andhave appeared and are clearly developed, resemblingapparatus
attachment
The
stage.
length those of the definitive

relativeispost-anal.

The body takes on a distinctly striped appearance.

The

which
mouth parts have the prostheca well developed on the mandibles,
are

The

placed and therefore have an up-and-down movement.
from the fourth
third larval stage represents the fully grown larva

verticallyCharles Paul Alexander

964

molt up to the time of pupation. Here the dorsum of the body is a dirty
brownish green with more or less distinct brighter spots, and the venter
The mouth parts and the attachment apparatus
is bright green in color.
are as in the second stage.

The pupa is comparatively active, moving about by flexion of theabdomen. Its usual position is vertical, with the tips of the breathinghorns just reaching the surface of the water.

This vertical position the

pupa maintains by grasping the vegetation with its caudal abdominal hooks.
At times the pupa descends beneath the water by clinging to the vegetation,
but a submergence of six hours causes asphyxiation. The pupal existence
was determmed by Miall and Shelford as eleven days. When the adultemerges, the cast pupal skin is left attached to a moss leaf by the dorsal
abdominal projections at the posterior end of the body.
Larva.

— Length, about 25 mm.

Young larvae distinctly greenish, especially on ventral side, this coloration caused, atleast in part,by contents of alimentary canal showing thru body wall; older larvae moreopaque, brownish green in color; dorsum with an indistinct striping of brown and whitish;
ventral surface whitish.

Head entirely retractile within prothorax and usually so retracted except when larva is
Prothorax, viewed from above, roughly rounded, anterior
margin convex; on ventral surface traversed by a weak suture. Meso- and metathorax
narrow. Abdominal segment 1 indistinctly divided into two annuli, the more basal one
very narrow; abdominal segments 2 to 7 divided into a narrow basal ring and a much broader
feeding; opening transverse.

posterior ring, each of these annuli

stillfurther subdivided into

two annulets^

Body pro-vided with numerous elongate trachea-bearing filaments, both simple and branched, giving
larva a very bristly appearance (Plate LXXXTII, 448), these spines arranged as follows: tergites

with both simple and bifurcated filaments; on posterior part of pronotum two short, simple

on meso- and metanotum, two pairs of simple filaments; on abdominal segment
an anterior pair of simple, and a posterior pair of deeply branched, filaments; segments

filaments;
1,

2 to 7 with basal ring unarmed, posterior ring with an anterior pair of simple, and a caudal
pair of deeply branched, filaments; segment 8 with only a branched pair, anterior branchlong, slender, posterior branch very small, its outer face heavily chitinized

and forming

dorsal lobes of spiracular disk; pleurites with all the filaments simple; one on posterior partof prothorax,

and on anterior part of each of the other two thoracic segments; posterior

filament on these latter nearly vestigial; two unequal filaments on first abdominal segment;

segments 2 to 7 with one filament on basal ring and two on posterior ring, the anterior one
the longest; segment 8 with a single rudimentary filament; stenu'tes with all the filaments
simple; prosternum without filaments; meso- and metasternum with a strong filament nearlateral margins; abdominal segment 1 with two pairs of filaments, posterior pair the longer
and more widely separated; segment 2 with three pairs of filaments, anterior pair very short;segments 3 to 7 with four pairs of filaments and an additional median one, arranged as follows:

The Crane-Flies of New York — Part II965

two pairs of small filaments on basal ring, the anterior pair more widely separated, twopairs of much longer filaments on posterior ring, the last pair more widely separated, longer,

and tipped with blackish, between them a tiny median filament. Chaetotaxy as follows:
tergites with four solitary setae along anterior margin of prothorax, and two setae justlaterad of each dorsal filament of prothorax and posterior filaments of meso- and meta thorax;
abdominal segments with a seta laterad of each simple and branched filament; plourites
with a stiff seta at base, and out toward apices, of middle lateral filaments; sternites with agroup of about four stiff setae on either side of posterior ring of presternum, and a single
stiff seta on either side of median line; meso- and metasternum with a lateral group of setae;
a seta laterad, and another at about midlength, of each of the posterior pair of ventral
filaments.
iSpiracular disk with ventral lobes elongate, slightly recurved, the posterior face intensely

blackened, chitinized, each lobe with two acute spines attip,with two setae near them;a stiff seta on side of base of each ventral lobe; dorsal lobes as already described, the rudi-

mentary posterior branch of last furcate dorsal filament much smaller than in ventral lobes,
the posterior face heavily blackened; spiracular disk rhombic in form, white,almost the appearance of porcelain.

and having

Spiracles situated between bases of dorsal lobes.

Skinabout spiracles capable of retraction so as to form a deep recess.

Head capsule rather short, almost conical, formed of two large lateral plates and a somewhat smaller and shorter prefrontal plate. Mentum with an outer (ectolabial) part andan inner (endolabial) part; mentum a triangular or somewhat pentagonal plate, strongly
chitinized, fore margin with about fifteen teeth; mandibles working against teeth of bothendo- and ectolabia.
Antenna of a single segment, bearing on its truncated apical end a
few sensory papillae. Mandibles small but strong, curved inward at tip and furnished witha fringe of setae, which assist in closing the mouth opening. Maxilla expanded into flattened,
shovel-like structures, inserted high on side of head; palpus with a number of sensory papillae

at apex, and with a porous plate on outer side which seems to be an organ of hearing.

Pupa.

— Length, 16-18 mm.

(Miall gives length up to 20 mm.)•Width, d.-s., 2.9-3.3 mm.
Depth, d.-v., 2.7-2.9 mm.

Coloration greenish brown, in alcohol a paler yellowish brown; a very broad dorso-mediandark brown stripe which is narrowly margined laterally with yellowish; dorsum of abdomen
suffused sublaterally with dusky, extreme lateral margins of body yellowish; ventral surface

with two broader sublateral stripes and a very narrow ventro-median stripe.
Anterior end of body very deep and thick, as is usual in this group of crane-flies.
greatly depressed, with lateral margins very thin and flattened.

Abdomen

Cephahc crest low, non-between antennal bases. Labrum broad basally, narrowed toward ape.x,
which is broadly rounded; two setae at base of labrum. Labial lobes subcircular in outline.

setiferous, locatedMaxillary palpi elongate, bent strongly backward so as to lie along flattened cheek.
rather elongated, extending far beyond origin of wing pad (Plate LXXXIH, -119).

very deep, flattened above.

AntennaThorax

Pronotal breathing horns, which are broken in the writer's speci-

mens, seen from other descriptions to be rather elongate, almost straight, and slightly
divergent; two small setae between bases of breathing horns; two groups of setae on anterior
part of pronotura before breathing horns.

Mesonotum (Plate LXXXIII, 449) withsixCharles Paul Alexander

966

small, setiferous tubercles, one above each wing axil, the other four arranged in a trapezoidon dorsum with the anterior pair closer together; metanotum with six setiferous tubercles,

four of which are median in position, the other pair at antero-lateral margin, near base ofWing sheaths ending about opposite apex of second abdominal segment.

halteres.

Legsheaths ending before apex of third abdominal segment.
First

abdominal segment similar to metanotum; segments 2 to 7 indistinctly subdivided

into a narrow basal ring and a much broader posterior ring; basal ring unarmed except fora seta on pleural margin and a trapezoid of tubercles on sternum; posterior annulus armed

and located in the dark
median stripe, posterior tubercles with a stiff seta just laterad of each; a lateral seta on aas follows: tergites with four tubercles arranged in a quadrangleline with anterior tubercles; two lateral setae on a level with posterior tubercles, the proximal

one considerably the larger; these setae located in the yellow sublateral stripe; pleural margin
with two setiferous tubercles with an acute subappressed spine between them; a seta immediately ventrad of anterior tubercle; on segments 6

and 7, spine closer to caudal margin ofsegment, and tubercle beyond it lacking; sternites with a trapezoid of naked tubercles onbasal ring, posterior pair the closer together; a trapezoid of larger setiferous tubercles onposterior ring, anterior pair the closer together; posterior punctures each having two setae,

with an additional slender seta laterad of these in the lateral dark stripe; dorsum of segment
6 (Plate LXXXIII, 450) with posterior pair of tubercles replaced by two powerful lobes
directed caudad and laterad; segment 7 unarmed on dorsum, sternum with two acute spines

near caudal margin; segment 8 with lateral angles produced dorsad into slender lobes which
are spinous on all the faces; caudal angles directed caudad into slender lobes, acute at tips

and with anterior inner face spinous and bearing a few setae. Male cauda with sternalvalves very blunt and rounded, on either side of ventro-median hne produced caudad into anacute spine. Female cauda with dorsal acidothecae a little longer than the more slendersternal valves, both pairs much exceeded by caudal angles of eighth segment.

———

Nepionotype.
Larva, Denmark.
Neanotype.
Pupa, Denmark.
Paratypes.
One larva and one pupa.

(The writer is indebted to Dr. C. Wesenberg-Lund for this material,
as well as for several other interesting European crane-fly

life

histories.)

Genus Cylindrotoma Macquart (Gr. cylinder + 1 cut)1834
Larva.

row.

Cylindrotoma Macq.

Suit, a Buff., vol 1, Hist. Nat. Ins., Dipt., p. 107.

— Body covered with simple tubercles, a median dorsal row and a double ventral

Spiracular disk large, surrounded by small lobes.

Head capsule compact.

Menturawith about fifteen teeth.
Pupa.-— Pronotal breathing horns short, cylindrical, directed strongly ventrad.

notum with two flattened lobes directed cephalad.

Meso-

Abdomen unarmed with spines or lobes.

Cylindrotoma is a small genus including six species distributed thruout
the North Temperate Zone, three of these species occurring in North
America.

The Crane-Flies of New York— PartII

967

The adult flies of Cylindrotoma tarsalis Johns., the only species thatthe writer has ever found in nature, arecool,

boggy, and swampy woods.

common on rank vegetation in

The immature stages of the genotype,

C. distinctissima (Meig.), have long been

known, having been discussed

by Schellenberg (1803), Boie (1838), Zeller (1842), Kaltenbach (1874),
and others. The larva shows some resemblance to that of Phalacrocera,
but is almost entirely

terrestrial in its habits, feedingon the leaves ofvarious spermatophytic plants such as Caltha palustris, Anemone nemorosa,

Ranunculus repens, Chrysosplenium, Stellaria nemorum, Sanicula europaea,
Viola biflora, Valeriana officinalis, Allium, and others.
It attains a lengthof nearly 25 millimeters, and is narrow, depressed, tapering to either end,and of a grass-green color. There is a slight dorsal ridge from which arow of short, fleshy spines projects, these spines being directed backward
and one spine on each segment being longer than the others. There is abroad lateral margin bearing very short processes, and there are also
eight pairs of ventral ridges without hooks and a pair of longer backwarddirected processes near the anus.
The pupa affixes itself to stalks or leaves
by the caudal end, to which the remains of the last larval skin adhere.
The larvae generally remain on the lower surface of the leaves, on which
they feed, gnawing holes in them. When about to pupate they generally
leave their food plants and fasten themselves on grass blades and leaves.

near by, usually pupating the following day.

From the foregoing obser-

vations it would seem that there are in the various locahties two generations a year, one in the spring and the other in the autmnn.

The life history of C. splendens has recently been worked out in conby Dr. A. E. Cameron, thru whose kindness the writersiderable detailhas received specimens for study.

Cylindrotoma splendens Doane
1900
1900
1918

Cylindrotoma splendens Doane. Journ. N. Y. Ent. Soc, vol. 8, p. 197.
Cylindrotoma juncta Coq.
Proc. Wash. Acad. Sci., vol. 2, p. 401.
Cylindrotoma splendens Cameron. Ann. Ent. Soc. Amer., vol. 11, p. 67-89.

Cylindrotoma splendens is an interesting crane-fly occurring from British
Columbia northward to Alaska. Dr. Cameron's excellent notes on thelife history of this species are abstracted in detail on pages 708 to 710
of this paper.
The following descriptions were made from material sentCharles Paul Alexander968

to the writer by Dr. Cameron, a few details being added from Dr. Cameron's

published notes.
Larva.

— Length, 15-17 mm.
Width, d.-s., 2.4-2.5 mm.
Depth, d.-v., 1.5 mm.

Coloration light chlorophyll green, with two narrow, pale brown lines on dorsum, extending
from posterior end, above spiracles, anteriorly, becoming more expanded and diffused onfore part of body.

Body very depressed, both dorsal and ventral surfaces being flattened, lateral margins Head completely retractile within prothorax. Skin delicately

sharp (Plate LXXXIV, 453).
reticulated

and roughened.

Thorax, viewed from above, semicircular in outline, marginedwith about four tubercles on either side, anterior pair the largest, separated by a V-shaped

Pronotum with an anterior pair of small tubercles and a larger median tubercle
and 3 each with lateral margins two-toothed, theanterior tooth the larger and more pointed; a blunt dorsal tubercle on anterior margin anda larger one behind. Abdominal segments indistinctly divided into four annuh which arepoorly delimited; the two basal annuli narrow and corresponding to basal annulus of mostcrane-fly larvae, the third annulus the largest and bearing a slender lateral tooth; othersegments less distinctly divided serrately on lateral margin; dorsum of abdominal segmentswith a row of blunt median tubercles, there being four on each segment, the first very small
notch.

behind, directed backward; segments 2

to vestigial, the third slender, the last the stoutest; on venter of abdominal segments
7,

1

to

near posterior margin, a fleshy conical lobe (pseudopodium) on either side of median line,

these being smaller on anterior segments (1 to 4),

segments (5 to 7).

and much larger and paler on posterior

Spiracular disk very large and flattened, obliquely truncated, surroundedby six small lobes, the dorso-lateral pair small, widely separated, the ventro-lateral pairmuch larger, the ventral pair very small, slender.

Spiracles very small, circular, widely

separated, the distance between them about equal to six times diameter of one.

Mouth parts with labrum linguliform, terminating in four teeth, external pair smaller

Mentum with seven teeth on either side of small median one, the first
and the third on either side being the largest. Hypopharynx with two rows of small teeth.
Antenna two-segmented; basal segment elongate, pyriform; apical segment thimble-shaped.
Mandible (Plate LXXXIV, 454) powerful; cutting edge with about five marginal teeth andtwo dorsal teeth; a large basal prostheca. Maxilla large; outer lobe fringed with stiff hairs;
a few acute sensory bristles situated on elevated papillae, two on cardo, one on outer lobeof maxilla; maxillary palpi short, cylindrical, apex obUquely truncated and provided withnumerous sensory knobs, on side near tip a circular porous plate (which as it appears inPhalacrocera is suggested by Bengtsson as probably being an organ of hearing).
than internal pair.

Pupa.

— Length, 11.7-14 ram.
Width, d.-s., 2-2.8 mm.
Depth, d.-v., 1.4-2 mm.

Color of live pupa, leaf green; preserved specimens much paler, yellowish; pronotal
breathing horns grayish white; margins of abdomen nearly translucent.

I

The Crane-Flies of New York
Labrum narrow, apex evenly rounded.

— Part II

969

Labial lobes stout, separated by labrum, at tip

narrowed and somewhat pointed. Maxillary palpi long and stout, just beyond base bent
Antennae rather short, bases not widely separated (Plate LXXXIV,
strongly backward.
456).

Thorax very deep, much flattened anteriorly(PlateLXXXIV, 455).

breathing horns short, cylindrical, slightly divergent, apex of each a Uttle expanded.

notum at declivity with two flattenedplates,PronotalMeso-

which are blunt and directed cephalad andWing sheaths ending just before apex
abdominal segment. Leg sheaths ending just before apex of third abdominal
segment; tarsi ending about on a level.
Abdomen strongly depressed, lateral margins carinate; segments distinct, unarmed, subdivided into a narrow basal ring and a much broader posterior ring; segments 2 to 5 a little
slightly laterad; above wing axil a smaller tubercle.

of secondand 7. Male cauda with valves divided into two rounded lobes
by a small median notch. Female cauda with dorsal valves straight, a little longer andmuch stouter than the slender sternal valves.
longer than segments 6——

Westhclme, Vancouver Island, B. C,
Nepionotype.
Female pupa with type larva.
Neanotype.

May 15, 1917.

Genus Liogma Osten Sacken (Gr. smooth + furrow)
1869

Liogma 0. S.

Mon. Dipt. N. Amer., part 4, p. 298.

— Body covered with elongate, leaflike projections, dorsal ones with one or two

Larva.-

teeth on anterior convex face.

compact.

Spiracular disk surrounded by four lobes.

Head capsule

Mentum with about fourteen teeth.

— Metanotum and abdominal tergites with elongate spines, branched in L. glahrala,

Pupa.-

simple in L. nodicornis.

The genus Liogma includes three known species, of which the genotype,
Liogma nodicornis (0. S.), is American. The adult flies of this species
are sluggish, and are found resting on vegetation growing in and aboutshaded ponds and similar situations.
In Europe the hfe history of L. glahrata (Meig.) has been worked out
by De Rossi (1876), by Wesenberg-Lund (1915:347-348), and in considerable detail by Mliggenberg (1901). The larvae were found in thewoods near Berlin, in wet, grassy spots where the moss Hyimum squarrosum
Brch. & Schp. occurs. The complete metamorphosis of the insect takes
one year, the larval life requiring the greater part of this period. The
egg stage lasts from eight to ten days and the pupal stage from eleven to

The duration of adult hfe is not known, but it is certainly
occupying but a few weeks at the most.

twelve days.
short,

Near Berlin the flies emerge during the first half of July. The males
appear first, the females later, and the latter are always seized in copulation

Charles Paul Alexander970

by the males just after they have forsaken the pupal skin and while still
Each female lays about sixty eggs, and theseteneral and undeveloped.
are deposited singly on the leaves or branches, or attached Hghtly to the

Hypnum squarrosum. Egg deposition begins fromday after copulation, and may be extended, with manyaxils of the leaves, of

one-half to one

interruptions of greater or less extent, thru a whole day.

The eggs arefrom 1 to 1.2 millimeters long, spindle-shaped, yellowish green in color,the chorion with a reticulate sculpturing. After the accomplishment ofoviposition the exhausted female soon dies.

The larvae when first hatchedare 2 millimeters in length and do not yet possess the beautiful moss-green

With the exception of the chitinized head capsule,
The numerous thornlike projections are to be
noted already in the same positions as those of the developed larva, but
do not show the branching of the later stages. In the autumn the young
larva grows very slowly, and dm-ing the winter it is still very small andcolor of the later stage.

they appear ashy gray.

difficult to detect.

In the spring the growth is greatly accelerated, andthe larva becomes fully grown during the latter half of June.

While
growth the larva molts several times, probably at leasteight
the number determined for Phalacrocera by Bengtsson.
Pupation
occurs in the moss where the larva happens to be. In its green color,
with brown blotches, the larva remarkably simulates its host plant andthe effect of the shadows cast by the plant stems and leaves.
The larvae
attaining—its

are extremely sluggish in their habits.

The American species Liogma nodicornis has been found in various mosses
Hypnum (Alexander, 1914). The immature stages of thisof the genus

species are discussed below.

The following keys separate the species of Liogma:
LarvaeProthoracic segment bearing foxir conspicuous dorsal projections about on a line; meso- andmetathorax with two pairs of dorsal appendages, each bearing two lateral teeth in front;
second abdominal segment with four dorsal appendages, the last two bearing two teeth/in front (Palaearctic)
glabrata (Meig.) (p. 969)
Prothoracic segment bearing four inconspicuous dorsal tubercles; meso- and metathorax withtwo pairs of dorsal appendages, the anterior pair small, both pairs simple; second abdominal
segment with four dorsal appendages, the last two bearing a single small tooth in front(Nearctic)
nodicornis (O. S.) (p. 971)

The Crane-Flies of New York— Part II

971

Pupae
Pronotal broathing horns directed cephalad and dorsad; roesonotum bearing two pairs ofspines, the more anterior being the smaller, situated just behind breathing horns, posterior
pair the larger; metanotum with two pairs of spines; abdomen with first tergite bearing
two pairs of spines, the first having two lateral branches, the second simple; second tergite
bearing two pairs of spines, the first with two lateral branches, the second with one branch;
third tergite bearing three pairs of spines, the first very short and simple, the second with
two lateral branches, the third with one branch; tergites 4 and 5 with three pairs of branches,the first two similar to those of third segment, the last possessing two lateralglabrata (Meig ) (p. 969)
branches
Pronotal breathing horns directed cephalad and ventrad; mesonotum spineless; metanotumwith one pair of spines; abdominal tergites bearing but a single pair of appendages, which'are unbranched and correspond in position to the last or more posterior of those of thenodicornis (O. S.) (p. 971)
European species

Liogma nodicornis (0. S.)
1865
1869
1887

Triogma nodicornis 0. S.

Proc. Ent. Soc. Phila., vol. 4, p. 239.
Mon. Dipt. N. Amer., part 4, p. 301.
Berl. Ent. Ztschr., vol. 31, p. 226.

Cylindroloma nodicornis 0. S.

Liogma nodicornis 0. S.

Liogma nodicornis, the only American species of the genus, is widely

The larvae are found inspecies.
They are thecrane-flies
known
the
writer,movingonly withofany
tosluggish
most
of
time
appearing
be
They
and
much
the
to
quite
dead.
great slowness
probably
of
their
host
plant
and
never
leave
among
the
stems
crawl about
durationindoors
is
not
than
six
The
pupal
more
it, even to pupate.
distributed thruout the Northeastern States.

moss — Hypnum cupressiforme Linn, and related

days.

Further details of the life history are given by the writer in an

earUer paper (Alexander, 1914).
Larva.

— Length, 14.5-15 mm.
Width, 3 mm.
Depth, 2.5 mm.

Color of live larva light green, the numerous spines covering the body darker; sides withseven black marks, thefirston first abdominal segment, thelaston seventh abdominal

segment; the marks on ends the smallest and least distinct, the five intermediate marks
large and conspicuous; these marks all lying parallel to one another; posterior face of ventral

lobes surrounding stigmal field intensely black.

Prothorax in front (Plate LXXXV, 462) sloping from anterior end, on ventral slope provided
with liplike lobes, with a transverse slit from which head capsule is exserted; upper lip the

few small, scattered bristles, these more
numerous on sides of lobe; lower lip not so high, with small, scattered bristles not arranged
in a row as in L. glabrata: at angle of slit a small rounded lobe bearing a small bristle.
Dorsalbody appendages reduced to a pair of lobes in front, separated by a space a little greaterthan diameter of one, and a pair of smaller lobes behind, very widely separated; lateral body
appendages long, conspicuous; ventral body appendages not apparent. Meso- and metathorax swollen and arched ventrally like prothorax; dorsal appendages two, a small conicalhigher, not strongly chit^nized, provided with a972Charles Paul Alexander

one in front and a much larger one behind bearing a small tooth in front
and with its tip
directed backward; lateral appendages, viewed from above, two in
number, anterior one thelarger, directed sharply backward, the second smaller, conical;
ventral appendages, viewed

from side, four, anterior pair the larger, posterior pair small, slightly behind
Dorsal appendages of abdominal segments (Plate

the others.

LXXX V, 457) as follows first segment with:two pairs of appendages; anterior pair the shorter, conical, tips strongly
recurved and bearing
a tiny tooth on anterior face at about midlength; posterior pairmuch longer, with tips
bent strongly backward, a small tooth on anterior face at about
one-third length' from base;segments 2 to 7 with four pairs of appendages, the first pair very small,conical, the second
exactly similar but larger, the third and fourth pairs similar toappendages of first abdominal
segment; the tiny anterior appendage largest on second segment, gradually

becoming smaller
Lateral abdominal appendages as follows: first segmentwith threepairs of appendages, the first directed laterad, the posterior
two more recurved and directedcaudad; segments 2 to 7 with four pairs of appendages, the first very small,situated at anterolateral angle of segment, the other three subequal anddirected caudad.
Ventraltoward end of body.

abdominal

appendages as follows: first segment with three pairs of appendages, which

are successively

larger from the short anterior one to the large posterior one;segments 2 to 7 with five pairsof appendages, the first three small, the fourth intermediate
in size between them and theenlargedfield

fifth pair.
Eighth segment bearing spiracular disk and its lobes; dorsalside ofwith a pair of long, slender lobes bent conspicuously cephalad.

Spiracular disk (PlateLXXXV, 461) small, oval, the two rounded-oval spiracles situated side by side and close
other, inclined

toward each other and capable of being closely appressed;

to eachventral lobes of

disk directed ventrad, inner faces of lobes with a conspicuous

jet-black line, tip of each lobe
(Miiggenberg regards these lobes as representmg theninth abdominal segment.) Ventral surface of terminal
segments with small protuberances.
Head completely retractile into first thoracic segment. Mentum (PlateLXXXV, 45S) with
about seven teeth on either side, terminal pair the larger, separated
by a V-shaped notch
Antenna (Plate LXXXV, 460) two-segmented, basal segment
elongate-cylindrical, tip very
short, thimble-shaped, with a diameter less than that ofelongate basal segment.
Mandible(Plate LXXXV, 459 and 460) many-toothed on inner face,with a prominent basal prosthecamandibles working vertically against teeth of mentum. IMa.xilla(Plate LXXXV, 458) with
palpi very short, broad, basal segment chitinized, tip
narrow, pale.

endmg in a sharp recurved hook.

Pupa.— Length: male, 10.4-n.4 mm.; female, 10-13 mm.
Width, d.-s.; male, 2-2.2 mm.; female, 2.2-2.6 mm.
Depth, d.-v male, 1.9-2.2 mm.; female, 2.1-2.5 mm.
;

Living pupae with pronotal breathing horns light yellow,

the terminal half a little morebrownish; a brownish black mark on prescutum; abdomen
greenish, more yellow behinddorsal spines clear light green thruout or with tips insome specimens a little infuscated'(In pupae preserved in alcohol, the greenish colors are
lost, the dark brownish blackmarkof the mesonotal prescutum is irregularly U-shaped,with the arms of the U directed backward, and the dark color is produced caudad and cephaladalong the dorso-median linethere is a triangular or rounded black spot on either sideof the scutellar lobe- on the metanotum IS a large blackish median blotch, which is continued
cephalad onto the mesonotal
postnotum; the abdomen has an interrupted brownish black
longitudinal line along eitherside of the midline of the dorsum; the posterior
margin of each tergite is suffused with

— Part II

The Crane-Flies of New York973

In old and fully colored pupae, bases of dorsal spines brown, with the tips paler;brown.)

head and thorax with appendages brown, in some specimens very dark; abdomen yellowish.
Cephalic part of head very flat and broad, without spines but with a small, blunt tubercle

between antennal bases.
apex.

Labrum transversely wrinkled, narrowed to the bluntly rounded
Sheaths*

Labial lobes large, divergent, each lobe rectangular with angles rounded.

of maxillary palpi rather long, curved strongly backward.

Antennal sheaths rather enlarged,

directed cephalad, bending around anterior margin of eye

ending just beyond knee joint of fore legs.

segments of the adult show thru the sheath.)464) 'large,

conspicuous,

directed

dorsadand thence directed caudad,(In older pupae, the peculiar nodose antennal

andPronotal breathing horns (Plate
laterad,

LXXXV,

the terminal half bent

ratherRlesonotum transversely wrinkled. Metanotum (Plate LXXXV, 4(58)with two long, slender spines, arising beyond midlength of segment, directed caudad andsuddenly cephalad.
slightly dorsad.

ment.

Wing sheaths broad, reaching posterior margin of second abdominal seg-

Leg sheath ending just before posterior margin of third abdominal segment; fore legs

the shortest, hind legs the longest.

Abdominal segments with a narrow basal ring and a broader posterior ring; segment 1
about half as long as segment 2; tergites 1 to 7 with a pair of long, slender, spinous projections, shortest on anterior segment, longest on seventh segment, these projections arisingfrom near caudal margin of segment, directed dorsad and caudad, those of anterior segment
almost parallel, those of posterior segments more divergent; segments 2 to 7 having lateral

margins produced into three sharp spines, one on basal ring and two on posterior ring ofeach segment, these spines directed laterad and caudad, the terminal spines more sharply

caudad than the other two; abdominal sternites armed as follows: segment 3 with a small,eubapical spine on either side, these spines very widely separated, segment 4 with similarspines but larger and more prominent, segments 5 to 7 similarly

armed but with anotherpair of small spines about midlength of segment and much nearer midline of body, segments

2 to 7 with a subbasal triangular pit on either side, these pits widely separated; eighth
tergite with caudal margin rounded, concave, the lateral angles produced backward, upward,

and slightly outward as strong spines; suture on ventral surface incomplete; two small spines
on either side of middle line of body; posterior margin of segment produced caudad as twostrong spinous projections.
Male cauda with sternal valves rather long, tipped with two
to four acute spines, in some specimens with two spines on one of the lobes and only one
on the other; tergal valves a little rounded at tips, sHghtly longer than sternal valves. Female

Cauda (Plate LXXXV, 465 and 466) with sternal valves slender, feebly notched at
tergal valves broader, rounded at tips, and with a deep median split.

tips;

—
—
—

Ncpionntype.
Coy Glen, Ithaca, New York, May 8, 1913.
Neanotype.
Ithaca, May 30, 1913.
Paratypes.
Larvae and pupae with types; others from Orono, Maine, June 17 and 19,
1913.

Genus Triogma Schiner (Gr. three + furrow)
1863
Larva.

Triogma Schin.

Wien. Ent. Monatschr., vol. 7, p. 223.

— Body covered with elongate

leaflike projections,as many as four teeth on anterior convex face.

Pupa.

some of the dorsal ones with

Spiracular disk surrounded by four lobes.

— Dorsum of abdomen with elongate branched spines.

Charles Paul Alexander974

The genus Triogma includes but three known species, the genotype
Triogma trisulcata (Schum.) of Europe, T. kuwanai (Alex.) of Japan,
and T, exculpta O. S. of the eastern United States. The last-named
species is very rare and its habits are entirely unknown.
The first reference* to the immature stages of T. trisulaita is by Steinmann(1907-08), who discusses the larva as that of an unknown species of
Phalacrocera.
The true identity of this insect was made known by
Miiller (1908-09). Steinmann found the larvae at Sackingen at the endof April, fourteen days after the melting of the snow.
They were foundin mountain streams, attached to and living among the stems of theaquatic moss Fonlinalis antipyretim Linn., to which they clung firmlyl)y means of two strong chitinized hooks at the caudal end of the l^ody.
The larva resembles to a startling degree the moss on which it lives.
Along the dorsal surface are two rows of leaflike appendages, each of theabdominal segments having three such appendages, of which the mostanterior one is the shortest and the posterior one is the longest.
The
anterior appendage is untoothed, the others have as many as four teethon the anterior face. The pleural region likewise bears a row of leaflike
structures, while the ventral surface shows a double row of small, knoblike
Thru the strong accentuation of the dorsal and the two lateral
leaflets.
rows, there is produced a copy of the ternate condition of the leaf arrange-Iment in FontinaUs.
The larva of T. trisulcata (Plate LXXXIII, 452) is of a light green color
marked with darker blotches, and measures 19 millimeters in length. The
longest body appendage is 1.5 millimeters in length.
Miiller found larvae
and also pupae. The pupa is characterized by the possession of somewhat
similar appendages to those of the larva, the dorsal row showing thepecuUar branched condition found in the larva.

SUBFAMILY TipuHnac

The subfamily Tipulinae comprises a remarkably homogeneous assemIt is made up of a relatively few but inblage of usually large species.

some cases very extensive genera, which are found thruout the world.
Efforts

have been madeinthe pasttomaintain

Dolichopezini, the Ctenophorini, and the Tipulini.

threetribes— the

It has become increas-

ingly difficult, however, to define these groups on the constant accessionof exotic generaand species.

A study of the immature stages likewise

IThe Crane-Flies of New York
I-

fails

--

.i

— Part975

III

to substantiate the validity of these groups, and, for the present

at least, or until otherand better characters may be found, it is betterand the Ctenophorini, founded, as they

to eliminate the Dolichopeziniare,on insufficient or sexual characters.

The species of the Tipulinae include the largest crane-flies known, someexotic species of Ctenacroscelis attaining aof the genera Longurio and Holorusia.
in thiswing expanse of nearly 10

In North America the largest species are representatives

centimeters.

The great majority of the species

subfamily are well above the average in size, this feature alone

being sufficient to eliminate
Tipulidae.

allbut a very few of the other groups of

The smallest member of the subfamily Tipulinae known to

the writer is Microtipula aniazonica Alex., of Brazil, in which the wing of
the male measures but 7.2 milhmeters in length andnarrow.

isvery long andPractically all of the common local species of this group belong

to the two genera Tipula and Nephrotoma.

The immature stages of members of the subfamily Tipulidae are foundwide range of habitats. Some species of Tipula are almost entirely
The majority of the known species live inaquatic in the larval state.

in amoist earth near water, or beneath damp cushions of moss.
as Oropeza, live inmuch drier mosses on exposed rocks.

Some, such

A rather con-siderable number of species (Brachypremna, Ctenophora, Dictenidia,

andwood or beneath the bark ofwood which is relatively sound,several species of Tipula) live in decaying

prostrate trunks.

and

this

Tanyptera

lives in

represents the extreme development of this tendency in thefamily.

The larvae are never very slender, and are usually very plump andIn a few cases only is the body decidedly depressed. There is

terete.

a definite arrangement of setae on the body, there being none on theanterior annulus of the abdominal segments except a single seta

pleura of either side.

The

spiracular diskissurrounded byon thesix lobes,

a number not found in the Limnobiinae.
the normal

In DoUchopeza the number
number in the Eriopterini, but

isdescribed as beingallother features of the genus are essentially tipuline.

the numl)er of lobes

is

five,

increased to eight.

In a few species

In the genus Tanyptera thelobes are exceedingly reduced in size so that the caudal end appears almost

The spiracles are always present and in some cases
The anal gills are almost always present and are variously

naked and exposed.
are very large.

Charles Paul Alexander

976

ideveloped according to the habitat. The head capsule is remarkably
iinifoT-m thruout the group, being l)road, compact, and massive, with theposterior incisions shallow and the prefrontal sclcrite very large andfringes of hairs.

The labrum is usually conspicuous, transverse, with
The mentum has from seven to nine teeth along theanterior margin,

and is deeply split behind but not completely divided.

usually distinct.

The hypopharynx consists of a narrow, flattened plate, with the basal
produced into strong arms and the anterior margin having
more numerous aad in some
The antennae are cylindrical, and are stoutest in thecases obsolete.
wood-inhabiting species; in many species of Tipula and Prionocera they
are long and slender, the length being about four times the diameter;
lateral angles

usually five teeth, the teeth being rarely

the apical papilla, in some cases obsolete,flattened.

The mandibles

isare not large inthe capsule; they have few teeth, in

usually very small andproportion

to theissize ofsome species only a dorsal and aventral tooth in addition to the apical point; the prosthecal appendage
isvariously developed.

The

maxillaearesimpleand generalizedinstructure.

The pupae are fairly uniform in structure thruout the subfamily.

The

tips of the sheaths of the maxillary palpi are strongly curved, or, in the

majority of species, actually recurved. The pronotal breathing horns arevariously formed, but in practically all species they are short, cylindrical,

and with the tips but little expanded. In some genera (Longurio,
Prionocera, and Tipulodina) the breathing horns are greatly elongated,
and, in some cases at least, are slightly unequal in length, the longer
measuring nearly half the length of the body. In some genera, suchas Prionocera and Holorusia, the horns are spht at their tips into two
divergent

flaps.

The only

short,

clavate horns in the subfamily are

The mesonotum is often provided with
The abdominal segments are
four or six variously developed tubercles.
almost always armed with transverse posterior rows of spines, theseranging in number from about four to twenty.
The large size of the larvae and the pupae will, as a rule, easily separate
The few
this subfamily from almost all members of the Limno])iinac.
large members of the latter group are readily separated by the characters
those of the genus Tanyptera.

outlined above.

The Crane-Flies of New York— Part977

II

The following keys separate the genera of the subfamily Tipulinae:
LarvaeDnlichopeza Curt. (p. 981)
Spiracular disk surrounded by five lobes; living in moss
2
Spiracular disk surrounded by four or six lobes, or with lobes indistinct
2. Spiracular disk with lobes indistinct; living in nearly solid or semi-decayed wood.
Tanyplera Latr. (p. 988)
3
Spiracular disk with lobes distinct
Tipula {selene Meig.) (p. 1016)
3. Spiracular disk with four slender, hornlike lobes
4
Spiracular disk with six lobes
Longurio Loew (p. 990)
4. Anal gills pinnately branched
Aeshnasoma Johns, (p. 993)
5
Anal gills not pinnately branched
5. Antepenultimate segment of abdomen with a strong lateral tubercle.
1.

6.

Oropeza Needm. (p. 982)
6
Antepenultimate segment of abdomen without such a tubercle
Dorsum of head behind antemia with a slender, fle.xible spine; living in wood.
Clenophora (apicata O. S.)

Dorsum of head without such a spine
7.

(p.

986)

7

Lobes surrounding spiracular disk elongate, digitiform, fringed with long hairs.
Prionocera Loew (p- 995)

8.

8Lobes surrounding spiracular disk less elongate
Size large (over 50 mm.); spiracular disk with the six moderately long lobes fringed withlong hairs; mandible small, with a dorsal and a ventral tooth; found in western North
Holorusia Loew (^p. 993)
AmericaSize smaller; if large {T. abdominalis) the lobes surrounding disk bifid; mandible withTipula Linn. (p. 998)
two or three ventral teeth
Nephrotoma Meig. (p. 1016),

Pupae
Pronotal breathing horns very long, slender, the longest one nearly if not quite half length2of body3
Pronotal breathing horns short, cyhndrical or flattened, subequal in size
2. Length 40 mm.; kmgest breathing horn 18 mm.; ma.xillary palpi not recurved at tips;
Longurio Loew (p. 990)
venation with petiole of cell Mi very shortLength 20 mm.; longest breathing horn 9 to 10 mm.; maxillary palpi recurved at tips;
Prionocera Loew (p. 995)
venation with petiole of cell Mi longer
3. Pronotal breathing horns short, compressed, deeply bicrenulated; living in wood.
Tanyplera Latr. (p. 998)
4
Pronotal breathing horns cylindrical
5
4. Maxillary palpi not recurved at tips
6
Maxillary palpi recurved at tipy
5. Pronotal breathing horns with apices deeply split; mesonotum with two tubercles;
abdominal segments with fourteen to thirty-four spines; found in Western States;
Holorusia Loew (p. 993)
living in mud
Pronotal breathing horns short, slender, apices not split; mesonotum with eight tubercles;
abdominal segments with four powerful spines near posterior margin; found in Southern
Brackypremna O. S. (p. 984)
States; living in wood
6. Mesonotum with two spines; ventral abdominal spines six to eight; fore and middle tarsi
Clenophora Meig. (p. 986)
subequal, shorter than hind tarsi; living in wood
7
Characters not as above
7. Mesonotum with a large, roughly triangular, reticulated area on either side of median
Oropeza Needm. (p. 982)
line; dorsum of cauda with four lobes
Mesonotum unarmed or with four or six lobes; dorsum of cauda with six, or rarely four,
Tipula Linn. (p. 998)
lobes
Nephrotoma Meig. (p. 1016)
1.

10

'

I
978

Charles Paul Alexander

The most important literature on the Tipulinae is as follows:
Dolichopeza albipes

Pupa

Dolichopeza alhipes

Larva, pupa, general

Tanyptera airata

GeneralGeneralGeneralLarva, pupa, general.
GeneralBeling, 1879:44-45 (as sylvicola).
.

.

.

Beling, 1886: 189-191 (as sylvicola).

Tanypkra alrnta
Tanyptera atrata
Tanyptera atrata
Tanyptera atrata
Tanyptera atrata ruficornis

1840:92; 1849:333.
Nordlinger, 1848.

Ferris,De Rossi, 1882.
.

.

Gercke, 1884, PI. I, figs. 12-19.

Hermann, 1880.

Larva

Gerbig, 1913: 150-158 (asCtenophora flamcornis)
Malloch, 1915-17 b: 194-195;

Tanyptera fumipennis

Larva, pupa, general.

Dictenidia bimaculata
Dictenidia bimaculata
Dictenidia bimaculata
Dictenidia bimaculata
Dictenidia bimaculata

Larva

Bouch^, 1834 32.

GeneralLarva, pupa, general
GeneralGeneralLarva, pupa, general
GeneralLarva, pupa, general
Larva, pupa
GeneralLarva, pupa
GeneralGeneralZetterstedt, 1851 4023,
Weyenbergh, 1872.
Beling, 1873 b 575.
Czizek, 1913: 102.
Reaumur, 1740, PI. I, fig. 9.
Zetterstedt, 1851 4016.
Weyenbergh, 1872.

.

.

1919.

Ctenophora flaveolata
Ctenophora flaveolata
Ctenophora flaveolata
Ctenophora pectlnicornis
Ctenophora pectinicornis
Ctenophora pectinicornis
Ctenophora pectinicornis
Ctenophora pectinicornis
Ctenophora jestiva
Ctetiophora festiva
Ctenophora nigricoxa
Ctenophora apicata
Ctenophora angustipennis

Ctenophora angustipennis

Aeshnasoma rivertonensis
Aeshnasoma rivertonensis

:

:

.

.

.

:

.

.

.

.

.

.

:

Bouche, 1834:29-31.
Fischer von Waldheim, 1838.
Weyenbergh, 1872.
Zetterstedt, 1851 4014.
:

Kaltenbach, 1874:202.
Kaltenbach, 1874:631.

Larva
Larva

Czizek, 1911: 48.
Lundstrom, 1906:7.

Pupa

Larva, pupa
Johannsen, 1910:32-33.
Larva, pupa, general.
Anthon, 1908.
Egg, larva, pupa, injury. Lovett, 1915.
Larva
Johnson, 1906: 1-2.
Johnson, 1907-12 [1909]: 115General
.

.

116.

Tipulodina pedata
Holorusia rubiginosa
Holorusia rubiginoia

Pupa

Prionocera fuscipennis.
Prionocera parri, supp

Larva, pupa, general.

(Falaearctic species)
Tipula caesia

Tipula dilatata
Tipula dilalata
Tipula flavolineata
Tipula flavolineata
Tipula flavolineata
Tipula fulvipennis

Larva, pupa, general

De Mei jere, 191 1 64.
:

.

.

.

.

.

Larva

Kellogg, 1901, a and b.
Comstock and Kellogg, 1904:
54-62.
Malloch, 1915-17b: 199-200.

Larva

Alexander, 1919 c.

GeneralLarva, pupa
Larva, pupa
GeneralLarva, pupa
Larva, pupa
Larva, pupa

Schiner, 1864 516.
Beling, 1886: 176.
Czizek, 1913: 169.
Staeger, 1840: 23.
Beling, 1873 b 581-582.
Czizek, 1913: 146.
Beling, 1879: 22-24 (as lutes-

Larva, pupa
Larva, pupa

Czizek, 1913: 74.
Beling, 1873 b: 578-579,
Gerbig, 1913: 154-156.

::

ce7is).

Tipula fulvipennis
Tipula hortensis
Tipula hortensis

Larva

The Crane-Flies of New York

979

Beling, 1879:25;

LarvA, pupa, general

Tipuld horlulana— Part II

1886:178-

179.

Tipui 'a horlulana
Tipui a imbecilln
Tipui
irrorata''
Tipui 'a irrorata
Tipuilateralis
Tipu, a lateralis

Larva, pupa, general.
GeneralLarva, pupa
Larva, pupa
Larva, pupa

Tipu 'a lateralis
Tipu lateralis
Tipu lunata
Tipu a luteipennis
Tipu a luteipennis
Tipu a marmorata
Tipu a maxima

Larva, pupa
GeneralLarva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa

Brocher, 1909.
Beling, 1886: 181-182.
Czizek, 1913: 143.
Beling, 1886: 182-183.
Beling, 1886: 177-178 (as gigan-

Tipu a maxima

Larva

Gerbig, 1913 152-153 (as gigan-

Tipu a maxima

Larva, pupa
GeneralCzizek, 1913 70-71.

Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
GeneralLarva, pupa
Larva, pupa
Larva, pupa

Beling, 1886: 183-184.
Beling, 1879: 28-29.
Czizek, 1913: 132.
Beling, 1873 b 575-577.

.

.

Czizek, 1913:90.

Loew, 1869 9.
:

Beling, 187.3 b: 586-587.
Czizek, 1913:98-99.
Beling, 1879 26-28.
Gerbig, 1913: 153-154.
Czizek, 1913: 128.
Cameron, 1917: 61.
/
:

Larva

'

iea).
:

tea).

TipUi a maxima:

Wesenberg-Lund, 1915:335 (asgigantea).

Tipu,
micans''Tipui a nigra
Tipui a nigra
Tipu, a nubeculosa
Tipu. a nubeculosa
Tipu, a ochracea
Tipu, a ochracea
Tipu, 'a oleracea
Tipu, oleracea.

Tipu 'a pabulina
Tipu a pabulina
Tipu a pagana
Tipu.

Tipu a pruinosa
Tipu, 3 rufina
Tipu, a scripta
Tipu a scripta
Tipu, a selene
Tipu, a selene
Tipu, a signata
Tipu, a signata

Tipu a simplex
Tipu, a subnodicornis

Tipula truncorum.

.

Larva pupa
Larva
Larva, pupa
Larva, pupa

paludosa

Tipu paludosa
Tipu a paludosa
Tipu, a paludosa
Tipu, a parva, supp
Tipu, a peliostigma
Tipu, 2 peliostigma
Tipu, a pruinosa

.

*.

^

GeneralLarva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
GeneralLarva, pupa
Larva, pupa.

:

'.
.

Czizek, 1913: 113.
Beling, 1873 b 582-583.
Czizek, 1913:157.
Del Guercio, 1914.
Patterson, 1908.
Beling, 1873 b: 579-580.
Czizek, 1913:86.
Beling, 1879:29-31.
Beling, 1873 b: 583-585.
Gerbig, 1913:136-151.
Czizek, 1913: 136.
Rennie, 1916; 1917.
:

Onuki, 1905.
Beling, 1879: 33-34.
Czizek, 1913: 159.
Beling, 1879:31-32; 1886:184.
Czizek, 1913: 148.

Mik, 1882 a.
Beling, 1873 b: 577-578.
Czizek, 1913: 110.
Beling, 1879: 34-35.
Czizek, 1913: 160-161.
Beling, 1879: 32-33.
Czizek, 1913: 104.
Doane, 1908.
Beling, 1886: 185-186.
Beling, 1879:24-25.

' Tipula mieann is oonsid-red by Kertrsz and others as a synonymoftions indicate that ho had two distinct species before him.

T. irroratti, but Beling's descrip-

980Charles Paul Alexander

Tijmla truncorum
Tipula ujica

Larva, pupa
Larva, pupa, general

Tipula unca

Larva, pupa

Czizek, 1913: 101-102 (as longi-

Tipula variicornis

Larva, pupa

Beling, 1886: 173-174 (as annu-

Tipula variicornis
Tipula variipennis
Tipula variipennis
Tipula variipennis
Tipula variipennis
Tipula variipennis
Tipula vernalis
Tipula vernalis
Tipula vittata
Tipula vittata
Tipula winnertzii^

Larva, pupa

Tipula winnertzii

Larva, pupa

Czizek, 1913: 122-123.
Beling, 1873 b 580-581.
Beling, 1886: 186.
Westhoff 1879.
Gerbig, 1913: 131-136.
Czizek, 1913:91-92.
Beling, 1879:25-26.
Czizek, 1913: 120.
Beling, 1886: 186-188.
Czizek, 1913: 78.
'..... Beling, 1873 b: 585-586; 1886:
188-189.
Czizek, 1913:84-85.

(Nearctic species)
Tipula abdominalis

Larva

.

.

Czizek, 1913:82.
Beling, 1886: 179-181 (as longi-.

cornis).
cornis).
licornis).

Pupa
Larva
General

Larva
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa

Tipula arctica
Tipula arctica
Tipula bicornis
Tipula caloptera

Larva, pupa
Larva, pupa
Larva, pupa

Tipula cunctans
Tipula cunctans
Tipula eluta
Tipula eluta
Tipula serta (?)
Tipula trivittata
Tipula trivittata
Tipula ultima

Larva, pupa, general
Larva, pupa
Larva, pupa, general.
Larva, pupa:,

Malloch, 1915-17 b 200-201 (asTipula sp. 2).
Nielsen, 1910:57-59.
Alexander, 1919 c: 18c, 19c.
:

Forbes, 1890.

Needham and Betten, 1901

Larva

575-576 (as abdominalis).
.

.

.

Ilyslop, 1910 (as infuscala).
Malloch, 1915-17 b: 204.

Pupa, generalHart, 1898 [1895]: 212-214.
Malloch, 1915-17 b: 203.
Malloch, 1915-17 b:205.
Greene, 1909.
Malloch, 1915-17b: 204-205.
Needham, 1903: 280-281 (asNephrotoma annlis
Nephrotoma analis
Nephrotoma cornicina

GeneralGeneralLarva, pupa
Larva, pupa
Larva, pupa

Caudell, 1913 (as flavicans).
Skuse, 1890: 95.
Beling, 1886: 172-173.
Czizek, 1911: 70-71.
Beling, 1879 39-40 (as iridi-

Nephrotoma cornicina
Nephrotoma crocata
Nephrotoma a-ocata
Nephrotoma lineata
Nephrotoma lineata
Nephrotoma lunulicornis
Nephrotoma lunulicornis
Nephrotoma maculata
Nephrotoma maculata
Nephrotoma pratensis

Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa

Czizek, 1911 :76.
Beling, 1879:40-41.
Czizek, 1911:87.
Beling, 1879:42—43 (as histrio).
Czizek, 1911 83-84.

Pupa
Larva, pupa

Pupa

.

.

flavicans).

Tipula ultima
Habromastix cinerascens:color):

Beling, 1879:41-42.
Czizek, 1911:61.
Beling, 1879: 36-37.
Czizek, 191 1 80-81.
Beling, 1886: 175-176.
:

sRiedel (191.3:2.5) considers Tipula winnertzii as a synonym of T. truncorum, but Beling's descriptions
indicate that the larvae he had before him represented two distinct species.

— Part II

The Crane-Flies of New York
Nephrotoma pralensis
Nephrotoma quadrifaria
Nephrotoma quadrifaria
Nephrotoma ferruginea
Nephrotoma ferruginea

Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa
Larva, pupa

981

Czizek, 1911: 8.5-80.
Beling, 1879:37-30.
Czizek, 191 1 66.
:

Hart, 1898 [1895]: 218-219.
Malloch, 1915-17 b:206.

Tribe Tipulini

Subtribe Dolichopezaria

Genus Dolichopeza Curtis (Gr.
1825
1830
1846

long

+ feet)Dolichopeza Curt. Brit. Ent., p. 62.
Leptina Meig. Syst. Beschr. Zweifl. Ins., vol. 6, pi. 65, fig.
Apeilesis Macq.
Dipt. Exot., Suppl. 1, p. 8.

10.

The genus Dolichopeza is a small group of flies including about a score
most numerous in the Oriental and Australian regions. Theof species,

immature stages of the European Dolichopeza alhipes Strom, a species
is very close to the American species D. americana Needm., havebeen described by Beling (1879, 1886). He found larvae in and beneathmoss cushions covering the piles of waste copper slag in the Harz MounOn June 11, 1878, larval and pupal material was taken from atains.
thattussock of the Jungermanniales liverwort Alicularia scalaris Corda.

The

pupal period was found to be six days.

Females were noted depositing theireggs in these hummocks, and small swarms of males were observed dancing
near by. It appears that the larvae feed on the upper side of the moss
cushion at night, withdrawing into the interior at other times. A larva wasfound in earth, which shows that these larvae are not dependent on mosses.
The larva is described by Beling (1886:189-191) as being about 12
millimeters long and 2.3 millimeters in diameter.
The body is almost
terete.
The color is a rather bright green, the dorsum being marked
with two zigzag dark brown stripes. The spiracular disk has but five
lobes; the three dorsal lobes are long and narrow, the median tooth being
formed by the apparent fusion of two teeth; the ventral lobes are very
short, are tuberculate, and have a small, dark brown, triangular markThere is a small dark cross-stripe at the base of eachat the inner tip.
lobe.
The
spiracles are small, are circular, and are separated bylateraldistanceequala
to about one and one-half the diameter of one. Beneath

The anal
The pupa measures about 14.5 millimeters
The pronotal breathing horns
in length and 2.5 millimeters in diameter.
each spiracle is a rather large, irregular, blackish brown spot.
gills

are strongly protuberant.

Charles Paul Alexander982

are directed straight outward and finally downward.

has acirck^ of four dorsal, four ventral,

are finely l)ifid at their tips.

The eighth segment
and two pleural spines, which

The pupa is green, as is the larva, with similar

angular dorsal stripes.

This is the only tipuline larva known to the writer with five lobes surrounding the spiracular disk. This feature is the more remarkable whenis compared with that of Oropeza, apparently closely related
but with a normal tipuline appearance. The immature stages of otherthis larva

species of Dolichopcza and related genera will be of interest.

The resting positions of Dolichopeza arc described on page 713 of thispaper, and the striking dissimilarities to Oropeza noted.

It

may be that

Oropeza is not so close to Dolichopeza as has been believed. Osten Sacken(1886:157) describes the mating of a European Dolichopeza, presumably
D. albipes, as follows:
I had occasion to observe the copula (in Heidelberg, July 26) the female was hanging downfrom some support to which it held on by its front legs; it bore the whole weight of the male,
who was fastened to it merely by the forceps, hanging head dov/nwards, with his legs stretchedout.
I have seen Bittacomorpha copulate in the same manner.
;

Genus Oropeza Needham (Gr, mountain + feet)
1908
Larva.
disk.

Oropeza Needm.

23d Rept. N. Y. State Ent. (1907), p. 211.

— Form somewhat depressed. A strong tubercle on sidesof body before spiracularby six lobes, the four dorsal ones slender, the ventral
Head capsule of the tipuline type. Mandible blunt, with about

Spiracular disk surroundedpair blunt.

seven teeth.

Pupa.

Anal gills blunt.

Hypopharynx five-toothed.

Mentum seven-toothed.

Coloration dark green.

— Pronotal breathing horns elongate-cyUndrical, slender. Mesonotum with a

Leg sheaths ending about
on a level, or fore legs a little longer. Abdominal segments with two stout pleural spines;
posterior armuii before margin with a transverse row cf twelve or fewer spines.

slightly elevated triangular area on either side of mid-dorsal line.

Oropeza is a genus of crane-flies including ten described species, allwhich occur in eastern North America with the exception of three
Japanese forms. The flies are common beneath dark culverts and bridges,
in outhouses, in crannies and crevices of rocky cliffs, beneath overhanging
boulders along mountain streams, and in similar situations. The resting
positions of ihv, adult flies are discussed on page 712.
Many species showa notable predilection for resting on spiders' webs. The immature stages
Otherof Oropeza ohscura are spent in dry moss, as discussed below.
species have been reared in sandy soil and in wet moss.
ofThe Crane-Flies of New York— Part983

II

Oropeza obscura Johns.
1909

Oropeza obscura Johns.

Proc. Boston Soc. Nat. Hist., vol. 34, p. 122.

Mr. Hyslop has sent to the writer, at various times during the past
two or three years, larvae of Oropeza obscura.
Numerous larvae andone pupal skin were found under dry moss (Hedwigia albicans [Web.]
Lindb.) on rocks in the South Mountains, Maryland, on November 4,
1916, by H. L. Parker.
They were associated with the larvae of a dascillid
beetle, Eunjpogon niger (Melsh.).
Additional specimens were found ina decaying log, but the preferred habitat seems to be beneath moss.

Thedark green larvae are very sluggish.
Larva.

— Length, 11-12 mm.
Diameter, 1.7-1.8 mm.

Color dark brownish green, ventral surface clearer green.

Form depressed; body short and stout.

Dorsum covered with small, blackened points,
Segments much wrinkled, especially

producing the dark color of this region of the body.

on basal ring.

On each side of penultimate segment a stout tubercle which simulates dorsal

Spiracular disk (Plate LXXXVl, 469) moderate in size, surroundedby six fleshy lobes, dorsal pair short and slender, lateral pair long and slender, ventral pairvery short and blimt; ventral lobes sparsely fringed with short, black hairs, and with a palelobes of spiracular disk.

apical area bearing a sensory bristle; inner face broadly suffused withbrown; at base ofother lobes similar but smaller triangular brown marks; a dusky area ventrad of spiracles.
Spiracles large, placed obliquely.

Anal gills four, large and blunt, pale yellow.

Labrum broad, with a short, blunt lobe on
median epipharyngeal region with dense rows of stout setae and two small
papillae on margin; lateral lobes with about four or five sensory bristles or papillae of various
Mentum (Plate LXXX^'I, 467) seven-toothed, median point long and slender, a prosizes.
longation of outer mental plate.
Hypopharynx (Plate LXXXVl, 468) five-toothed, thethree intermediate teeth the largest, lateral tooth on either side small.
Antenna slender,
cylindrical, with apical segment very reduced.
Mandible rather slender, with about seven
Maxilla with cardo large, triangular, with two setifero us punctures,
blunt teeth on margin.
the outermost very large and hyaline and with two long, powerful setae; lobes of maxilla
fringed with dense hairs; palpus short, disklike, with a large group of tiny hyaUne papillae
Head capsule of the massive tipuline type.

either side;

at apex.

Pupa.

— Lengthof cast skin,about 12 mm.

Coloration brown; pronotal breathing horns dark blackish brown.

(It ia possible thatthe pupae when alive are dark green in color, like the larvae.)

Labrum triangular.
palpi short

Labial lobes large, separated by apex of labrum. Sheaths of maxillaryand stout, apex recurved to beyond midlength. Antennal sheaths moderately

elongated.

Pronotal breathing horns elongate, slender, cylindrical, tips alittleexpanded.

Mesonotum on either side of median line with a large, roughly triangular, reticulated area
which is sUghtly elevated, with apex of triangle directed toward, mid-dorsal line. Mesonotum

Charles Paul Alexander

984
transversely wrinkled.

Metanotum (Plate LXXX VI, 470) near anterior margin with a trans-verse row of setiierous punctures, there being four on either side, the intermediate ones with

Wing siieaths reaching base of third abdominal segment. Leg sheaths reaching
beyond midlength of fourth abdominal segment, fore legs a little the longest, hind legs
two setae.

a little the shortest.

abdominal segment on dorsum somewhat similar to metanotum, with a transverse
four punctures at about midlength, the lateral ones with two setae; remaining
abdominal segments (Plate LXXXVI, 471) subdivided into approximately equal basal and
First

row

of

posterior rings; on tergites, posterior ring with a transverse row of spines, basal ring unarmed;
on sixth tergite, where best developed, spines on posterior ring numbering about twelve, witha seta at each end of the row and two others interspersed; on anterior tergites, spines very
email; on seventh tergite spines large, elongated; on eighth tergite four powerful spines intransverse alinement; on pleurites, a strong spine on each ring, that of basal ring deeply
bifid and with a strong seta in notch thus formed; posterior ring with a single spine bearing

a long seta on its face; on sternites, condition generally similar to that on tergites, the spines
a little more prominent but of about the same number, these spines slender at their sHghtly

Female cauda (Plate LXXXVI, 472 and 473)
on either side of tergites at base with a powerfulcurved tips and lacking on sternites 2 to 4.
with acidothecae elongate, contiguous,incurved hook; posterior margin of eighth tergite with four spines, lateral pair alarger; ventral side at base with a medianlittletheprotuberance bearing two powerful lateral lobes

and two smaller chitinized spines which are directed caudad and ventrad; base of segment
8 with a crossrow of eight spines; posterior ring of segment 8 with a powerful spine atlateral end.

———

Nepionotype.
South Mountains, near Myersville, Maryland, March 31, 1916.
Neanohjpe.
Hagerstown, Maryland, May 9, 1916; cast pupal skin in collection ofUnited States National Museum.
Paratypes.
Larvae from type locality.
'

Genus Brachypremna Ostcn Sacken (Gr. short + trunk)
1886

Brachypremna 0. S.

Berl. Ent. Ztschr., vol. 30, p. 161.

— Unknown.
Pupa. — Antennal segments enlarged at base. Antenna short. Sheaths

Larva.

palpi short, not recurved at tips.

ofmaxillary

Pronotal breathing horns short, slender, finely annulated.

Mesonotum with eight prominent tubercles.

Leg sheaths reaching beyond midlength ofAbdominal
segments each with four slender spines on posterior ring of both sternites and tergites, justbefore posterior margin; two pleural spines; eighth segment of male with four prominent,
fourth abdominal segment, fore tarsi very short, other tarsi subequal in length.

spinous lobes.

Brachypremna is a small genus, including but nine recent species foundand Tropical regions of the New World. Brachypremna
eocenica Meunier is described from the Baltic amber.
The flies of thein the Austral

best-known

species,B. dispellens, are

Imown in parts of the Southern

IThe Crane-Flies of New York— Part II

985

shady places and have"weavers." They frequent ratherof!t Lkable dlnce over three or four" feetgiven them by Johnson. This
name "king of the dancing tipuUds
immature stages arc at all known.
species is the only one whose
Statesas^^^^^^^^^^^^^

Brachypremna dispellens (Walk.)
333-334.
Ent. Sec. London, n. ser vol. 5 p.
Tipula dispellens Walk. Trans.
Ztschr., vol. 30, p. 162.
Brachypremna dispellens 0. S. Berl. Ent.
widely distributed species of theBrachypremna dispellens is the most

1860
1886

North An.er.a
from New Jersey southward thru-^^
by R. C- Sha-^ofoundlarva
Argentma. A?hru South America as far as1913
near Washington, D C., on April 23,
in a rotten log by a stream
les.
spe
this
of
male
adult
an
May aswas placed in rearing and emerged in
described.
here
is
and
preserved
badly mutilated pupal skin was
genus.

It ranges

The

No part of the larva was preserved.
p.inn— Leneth of cast pupal skin, about 18 mm.
brown sublateral stnpe on both ventral
Soltion browmsh yell; abdomen with a broad
spot.
brown
a
in
set
spinespleuraland dorsal segments; each of
crowded at base, soon passmg mto the short,
and
large
very
spinesAnt^malHead small
Labial obes
(Plate LXXXVII. 474) blunt.
^^^^^y^^^^^^^
slender flagellum. Labrum

lobe atlarge, transversely rectangularso as to appear as a single^^^^^^^^^^^..^f(Plate11, 4/o)horn
breathmg
Pronotal
tip.
atrecurvedof maxillary palpi short, notmargin
apex;
small
the
to
gradually
annuli, taperingsLdl slender curved, ringed with fineconspicuous, blunt, naked tubercles;
eight
with
Mesonotum
e wUh b^^^^^^^^^ pores.
of a trapezoid; anterior mediantubercles larger, arranged in the form

LXXX\

^^

o7al

the'our Lrmediate

Wing sheaths reaching end ofclose to mid-dorsal hne.
pa r high conical, located rather
LXXXVII, 476) extending beyond nudsend abdominal segment. Leg sheaths (Plateof third
short, endmg opposite base
very
legs
fore
length of fourth abdonnnal segment;
tarsal segment of other legs.

.

.

spines before posterior margin;

segn.ent^2

Abdominal tergite 1 with a pair of long, slendertransverse row of four long
and a posterior ring, the latter with ato 7 subdivided into a basal

six such spines; stermtes

seventh tergite with about
L der spines before posterior margin, the pleurites
posteriorwith a slender spine on basal andt^Zl with four spines on posterior ring;protuberant spiracle^
between spines, an indistinct, slightly
at base of posterior rLg
near basesmall, valves blunt; on dorsa side
narrowed,
477)
LXXXVII,Mat Cauda (Platechitinized spine; a small acute spme on

Tgin a slender,
four conspicuous lobes, each terminatingbase.
sides of ninth segment at

Neanotype.- est pupal skin, Washington,
States National

Museum).

UnitedD. C, May, 1913 (in coUection of

986Charles Paul Alexander
Subtribe Ctenophoraria

Genus Ctenophora Meigcn (Gr. comb + to bear)^/'S^["^'"f"''Ctenophora Meig.
Phoroctenia Coq.

?om
lsU.5
1910

Nouv. Class. Mouch., p. 13 (nomen nudum).
Ilhger's Mag., vol. 2, p. 263.

Proc. U. S. Nat. Mus'., vol. 37, p. 589.

Larva.— Body opaque, integument rather thick.

Pupa.— Sheaths of maxillary palpi recurved.

Two spines on mesonotum.
tarsi.

Spiracular disk surrounded by si.x lobes
Pronotal breathing horns long and slender

Fore and middle tarsi subequal, a
Ventral abdominal segments with six to eight spines.

littleshorter than hindCtenophora is a small genus including about sixteen
nominal species
found thruout the Holarctic region. The larvae andthe pupae occur
m decaying wood. The early stages have long been known, having been
described by Reaumur and De Geer.

In Europe, Ctenophora flaveolata (Fabr.) is
described and figured by(1740) and "by Weyenbergh (1872).
C. pectinicornis

Reaumur(Linn.)

described or mentioned

by Bouche (1834), by Fischer von Waldheim(1838), by Zetterstedt (1851:4014),
by Weyenbergh (1872), and by
Kaltenbach (1874). C. festiva Meig. was reared by Kaltenbach(1874:631)
ISfrom larvae in decayed beech stems. C. nigricoxa
Lundst. (Molpighia
vittata Meig., auct. Frey) was reared by
Lundstrom (190G:7) from pupae
in rotten birch stumps.
The immature stages of the various European
species of Ctenophora are described as livingin the wood oftrees such as willow(Salix), birch(Betula),variouscherry (Prunus), and otherhardwood species.
In North America, C. apicata is described by
Johannsen (1910) fromelm (Ulmus), and C. angustipennis Loew by Anthon(1908) in alder (Alnus)
and in poplar (Populus). The latter species is recorded
also as injuring

prune trees (Prunus) in Oregon, by Lovett(1915), who gives an excellent
account of all stages and the type of injury done.
The female lays from
200 to 400 eggs, which hatch in from nine to seventeen
days and thelarvae tunnel into the surrounding dead wood.
Here they feed and growreachmg maturity the following spring. Pupation
takes place in theburrows, the pupal stage requiring about
ten days.
Osten Sacken
1877:211) supposed that the larvae live in thestumps of redwood(Sequoia), but this has never been confirmed.
Ctenophora apicata O. S.

ISW

Ctenophora apicata 0. S.

Proc. Ent. Soc. Phila., vol. 3, p. 46.

The Crane-Flies of New York— Part987

II

The larvae and the pupae of Ctenophora apicata that were describedby Johannsen (1910) have been studied by the writer in the collection
They were collected atof the Maine Agricultural Experiment Station.
In 1913
Orono, Maine, on June 23, 1909, by Dr. William C. Woods.
the writer examined the stump from which they were taken, and founda few cast pupal skins.
The notes here given are taken partly from Dr. Johannsen's description

and partly from the original material.
Larva.

— Length, about 30 mm.

Color white.

Body

stout,

cyhndrical.

Antemiacylindrical,

On dorsum of head behind antennae, a slender, flexible spine.

with anapicalpaoilla.

Spiracular disk surroundedby six lobes, dorsal and lateral pairs slender, finger-like; ventral pair blunt.
Pupa.

— Length, 25-27 mm.

^

Length of breathing horns, 3 mm. additional.
Width, d.-s., 5.2 mm.
Depth, d.-v., 5 mm.

Pupa somewhat similar to that of Tanyptera, differing as follows:a little depressed.

Form stout; abdomen

Sheaths of maxillary palpi strongly recurved at tips.

horns long and slender.

Mesonotum with a prominent

Pronotal breatliing

tuberculate spine on either side of

median line. Legs shcrt, ending before tip of third abdominal segment; tarsal sheaths ending
about on a level, or those of hind legs a very httle longer. Abdominal segments 5 to 7 with
Female cauda with six powerful ventral spines or tubercles and two
six to eight spines.
dorsal tubercles on either side. Valves of ovipositor short, tergal valves a Httle longer thansternal valves.

———

Nepionotype.
Orono, Maine, July, 1909.
Nennotype.
Orono, Maine, July, 1909.
Paratypes.
Numerous pupae in collection ofMaine Agricultural Experiment Station.

Genus Dictenidia Brulle (Gr. double + comb)
1833
1856
1863

Dictenidia Brulle.
Ann. Soc. Ent. France, vol. 2, p. 401-402.
Ceroctena Rond.
Dipt. Ital. Prodr., vol. 1, p. 186.
Dicera Lioy. Atti dell' Institut Veneto, ser. 3, vol. 9, p. 216.

Dictenidia is a genus of Palaearctic crane-flies including three species,

which one is European and the others are Japanese. The genotype,The immature stages
is very well known.
are described or mentioned by Bouche (1834), by Zetterstedt (1851), by
Weyenbergh (1872), by Beling (1873 b), by Czizek (1913), and by otherof

Dictenidia bimaculata Brulle,

investigators.

Beling found the larvae in decaying birch (Betula).

He

describes the larvae as being grayish yellow, with four spiracular lobes.

The pupal duration is seven days.

Osten Sacken (1886:173-175) statesthat he has often found larvae in the wet detritus underneath the bark

Charles Paul Alexander

988

The larvae are much closer to Ctenophora than toof decaying trees.
Tanyptera, the skin being tough and opaque, with a fine pubescence, andthe spiracular disk consisting of small but distinct lobes. The pupae likewise are similar to those of Ctenophora, having the pronotal breathinghorns elongate, five spines on abdominal sternites 3 to 6, and four spines on
tergites 2 to 7.

Genus Tanyptfera Latreille (Gr. extend + iving)
1805
1832
Larva.

Tam/ptern Latr.
Xiphura BruU^.

— Integument very

Hiat. Nat. Crust, et Ins., vol. 14, p. 286.
Ann. Soc. Eut. France, vol. 1, p. 206.
thin,

with numerous setae, those on dorsum very small.

Spiracular disk with lobes practically lacking.
last segment.

tooth.

Anal gills bluntly rounded.

AntennaSpiracles large, lying exposed on the face ofMandible small, with one dorsal and one ventral

cyUndrical, capped with an apical cone.

Mentum with seven to nineteeth.

Pupa.— Cephalic crest lacking.

Sheaths of maxillary palpi not recurved at tips.

breathing horns large, broadly flattened, margin deeply crenulated.
blunt tubercles.

Pronotal

Mesonotum with two

Abdominal segments with six to ten spines on tergites, three to five spines

on sternites.

Tanyptera is a small genus including about twelve nominal species
whose limits and relationships are still but little understood. The immature stages are spent in the decayed or partly decayed wood of various
deciduous trees.
In Europe, Tanyptera atrata (Linn.), the genotype, was found by Ferrisand by De Rossi (1882) in decaying alder (Alnus) stems. Nord(1848) found the same species in linden (Tilia) and in poplar(Populus).
It has also been taken in oak (Quercus), beech (Fagus),birch (Betula), and other hardwood species.
Gerbig (1913) discussed thevariety ruficornis Meig. under the name Ctenophora fiavicornis.
In America, Malloch (1915-17 b: 194-195) describes T. fumipennis(0. S.) from a much-decayed chestnut log (Castanea), and later (1919)
in basswood (Tilia), where the species was associated with larvae of
Xylota fraudulosa Loew and Chalcomyia aerea (Loew), of the family
Tanyptera frontalis, discussed below, was found in red mapleS)a-phidae.
(1840)

linger(Acer).

Tanyptera frontalis (O. S.)
1864

Ctenophora frontalis 0. S.

Proc. Ent. Soc. Phila, vol. 3, p. 48.

The Crane-Flies of New York— Part989

II

The writer found numerous larvae of Tanyptera frontalis in a fallen
maple {Acer riibrum Linn.) near Beebc Lake, Ithaca, New
Larvae of several sizes were found. They
which
was well preserved, not entirely sound but
inwood
worldng
were
The larvae pupatedstill so hard that it had to be cut with a hatchet.
log of red

York, on IMarch 22, 1913.

A small male emerged on May 1.
Other larvae of Tanyptera were found in a hickory log (Carya sp.)at Sandy Landing, Virginia (opposite Plummers Island), on September
Detailed drawings of the larval structure made by Dr. Boving
9, 1913.
are in the collection of the United States National Museum.
in April.

Larva.

— Length, 30-35 mm.
Diameter, 7-7.2 mm.

Coloration, pale yellowish white.

Form terete, very stout. Integument thin. Numerous long black setae on segments
LXXXVIII, 482), arranged in transverse rows before posterior margin; setae of
dorsum (Plate LXXXVIII, 481) very tiny, one on either side of median line; a pair of setae(Platelaterad of these and in alinement; setae on pleural region very long and delicate; on thoracic

segments, setae at about midlength; on abdominal segments, setae closer to posterior
margin; mid-ventral setae very tinj^ four in number, on thoracic segments at about midlength, forming a

stiff

pencilon sides of venter, with two small setae between; laterad ofthese four intermediate setae, a group of three setae, two long and one very short; ventral setae
lying at a level posterior to that of pleural setae.

483) with lobes practically lacking, thetwo

LXXXVIII,

Spiracular disk (Platelarge, oval

spiracles lying

exposed on trun-cated end of last segment; above and laterad of each spiracle, a small, blunt lobe with ablackened mark and three long setae; below spiracles, two narrow black lines representing
the two ventral lobes; a small pencil of setae below each of these marks, and each markhaving a sensory bristle; three or four long setae on sides of spiracular disk. Spiracles withsmall middle piece black, ring yellowish brown; spiracles separated by a distance a little
bluntly rounded, very protuberant

andgreater than diameter of one.

Anal

evidently formed for propulsion.

(In older specimens the anal gills are minutely roughened

gills

four,

and are darker in color.)

Head capsule of the massive tipuline type.

Labrum rather broad; median epipharyngeal

region with a small brush of hairs surrounding two sensory setae; lateral lobes large, on theirand a flattened hyaline peg.
JMentum (Plate LXXXVIII, 478) rather small, broadly rounded, anterior margin withseven or nine teeth, in the latter case the outermost tooth on either side very blunt andAntenna (Plate LXXXVIII, 479) shortreduced, the median tooth long and flattened.
ventral face densely hairy, surrounding four sensory setae

cylindrical; apical papilla very small, hyaline, conical,*

with surface sculptured; in addition

to this cone, three or four small hyaline sense pegs; the usual auditory organ located atabout midlength of segment.

Mandible (Plate LXXXVIII, 480) small, with one dorsal andone ventral tooth in addition to the apical point, ventral tooth flattened and with margin
crenulated; a stout seta at heel of mandible; prostheca an elongated cone, situated at baseCharles Paul Alexander990
of mandible.

IMaxilla rather small, simple; palpus large, antenniform, with apex bluntly
rounded; inner lobe densely covered with short, stout setae surrounding a powerful bristle
and a small brown sensory organ.

Pupa.— Length, 30-33 mm.
Width, d.-s., 5-6 mm.
Depth, d.-v., 5.5 mm.
Coloration pale yellow; pronotal breathing horns liver-colored.
(In older specimens>
the thorax and appendages are dark-colored, and the abdomen has broad brown sublateral
stripes.)Head rather small, cephalic crest lacking.
beyond ends o palpi, segments angulated.
Labial lobes slender, divergent.

but not recurved (Plate

Antenna stout, rather elongate, extending far

Clypeus and labrum tumid, transversely wrinkled.

Sheaths of maxillary palpi rather slender, tips curved
Pronotal breathing horn (Plate LXXXVIH,LXXXVIII, 485).

486) large, broadly flattened, slightly incurved, broader at apex than at base, deeply furrowed

up middleof outer face, the broad margin thus formed deeply wrinkled to crenulate.
JMesonotum large, transversely wrinkled, with two blunt tubercles provided with short setae(Plate LXXXVin, 484).

Wing sheaths small, reaching end of second abdominal segment.

Leg sheaths ending before tip
tarsi the longest, those of

of

third

abdominal segment; foretarsi

very short, hind

middle legs intermediate.

First abdominal tergite with two spines; segments 2 to 7 broad, divided into the usual basal
and posterior rings; second segment on posterior ring with four spines; segments 3 to 6 with
six to ten spines, the intermediate ones usually smaller; segment 7 with four spines; pleurites

with a spine on each ring; sternites with similar arrangement to that of tergites, but spines
usually fewer in number, segments 3 and 4 with only a single widely separated spine on eachs.de,segments 5 and 6 with four or five spines, segment 7 with three spines; segment 8 has

Male cauda blunt, dorsal
Female cauda (Plate LXXXVIII,

four small spines between the large lateral ones described below.
lobes very divergent, ending in sharp, chitinized points.

487) with tergal valves elongate, narrowed to the moderately acute tips; sternal valvessimilar in shape but smaller; lateral lobes of ninth

at tips running out into chitinized points;

segment directed caudad and laterad,

two small setae before tips; segment 8 with apowerful lateral lobe on either side, each terminating in a chitinized point; posterior
eral angles oflat-segment produced into slender, blunt points.

———

Nepionolype.
Ithaca, New York, March 22, 1913.
eanoly pe.
M&y 1, 1913. No. 11-1913.
Four larvae and two pupae with types.
Paratypes.

N

Subtribe Tipularia

Genus Longurio Loew (Lat. a tall man)
1869
Larva

Longurio Loew.
(supposition).-Berl. Ent. Ztschr., vol. 13, p. 2.

— Body^massive.

Integument

semi-transparent.

Formclearly

by six lobes, dorsal pair very small, ventral pairvery long; ventral and lateral lobes provided with but few setae at and near tips; spiraculardepressed.

Spiracular disk surrounded

The Crane-Flies of New Yorkdisk

and lobes unmarked.

Spiracles small.

Anal

gills

— Part II

branched.

but a single dorsal and ventral tooth in addition to apical point.

991

Mandible small, with

Mentum with nine teeth.

Hypopharynx five-toothed.
Sheaths of maxillary
Antenna short, ending opposite tips of maxillary palpi.
Pupa.
Mesonotum unarmed. Pronotal breathing horns very long andpalpi not recurved at tips.
Wings showing clearly the characteristic
slender, one, at least, about half l.ength of body.

—

Abdomen elongate, posterior ring of individual segments with short,

venation of Longurio.

stout spines, including a few on pleurites; dorsum of eighth segment with four powerful lobes.

Longurio is a small genus including al)Out ten described species, ofwhich two occur in eastern North America. The genotype, Longurio
The immature stages of thistestaceus, is the best-known species locally.
They are spent in sand
first
time.
species are here discussed for theor sandy earth.

The branched anal gills of the larva, and the excessively

elongate breathing horns of the pupa, are notable features.

Longurio testaceus Loew
1869

Longurio testaceus Loew.

Berl. Ent. Ztschr., vol. 13, p. 2.

Longurio testaceus is probably the largest

ci*ane-fly inNorth America,

the females exceeding the lietter-known Holorusia of the Western States.

The adult flies are difficult to capture, being very wary. When the insect
at rest the body generally hangs perpendicularly, with the wings foldedisincumbent over the abdomen.
On November 9, 1916, Mr. Hyslop sent the writer two living larvae
which are referred with little doubt to this species. They were foundin wet sand in a bog on the top of South Mountains, near Myersville,
Maryland. The larvae were very restless, the head capsule being conWaves of contraction start from thestantly exserted and withdrawn.
A large pupa taken
posterior end of the body and pass toward the head.
by Dr. J. C. Bradley at Tallulah Falls, Georgia, on June 17, 1910,
undoubtedly belongs to this species, the venation being clearly apparent
on the wing pad. An additional cast pupal skin is in the collectionof the United States National
Larva (supposition).

Museum.

— Length, 31 mm. contracted, 58 mm. extended.
Diameter, 11-12.5 mm.

Color whitish, subhyaline; thoracic segments more yellowish; in life the brown food contents

showing clearly thru abdomen.

Form very depressed, lateral folds prominent; body very stout and fleshy. Skin very thinand semitransparent, showing internal organs within, practically destitute of pubescence;

Charles Paul Alexander992

a few short setae on body, especially on thoracic segments. Spiracular disk (Plate LXXXIX,
490) large, flattened, surrounded by six lobes; dorsal pair very small, represented by two short,conical protuberances; lateraland ventral lobes slender, the latter a little the longer, eachwith three or four long, delicate setae at tip and two or three others before tip on outer face;a long seta on margin of disk between dorsal and lateral lobes; spiracular disk and lobes
Spiracles small, circular, stigmal rings very narrow; spirentirely unmarked with darker.
acles separated

by a distance equal to about three times diameter of one.

large, pinnately branched,

Anal gills four,

each gill with six lateral branches.

Head capsule massive, of the tipuline type.

Labrum with a distinct, densely hairy lobe

on either side. Mentum (Plate LXXXIX, 488) large; anterior margin with nine slender
teeth, median one the longest, outermost teeth on either side blunt, flattened, evidently
formed by fusion of two teeth. Hypopharynx (Plate LXXXIX, 489) five-toothed; teeth
Antennablunt, the three middle ones larger, the lateral tooth on either side a little smaller.
Mandiblelong and slender, cylindrical; apex blunt and without distinct sense pegs or setae.
small, with a large conical dorsal tooth and a single flattened ventral tooth in addition
to apical point; prostheca distinct.

Maxilla small; palpi large, cylindrical, truncated atapex and with a circular auditory plate near end; inner lobe of maxilla with abundant elongate setae.

Pupa.

— Length, 42-45 mm.
Length of longest breathing horn, 18-19 ram. additional.
Width, d.-s., 3 mm.
Depth, d.-v., 4.2 mm.

Color dark brown; pronotal breathing horns paler attips;

mesonotum chestnut brownj

abdominal segments indistinctly ringed with pale and darker.
Front between antennal bases swollen, finely tuberculate but without distinct

Antenna very short, ending opposite tips of maxillary palpi.
maxillary palpi not recurved at tips.

Labrum large.

setae.

Sheaths of

Mesonotum unarmed, with fine transverse wrinkles.

Pronotal breathing horns (Plate LXXXIX,491) very long and slender, the right one, at least,
exceedingly elongate, with tip expanded.

(The left breathing horn was broken before theapex in both the pupae studied; it was almost as long as the right horn, and may, of course,
have been longer.) Wing pads reaching end of second abdominal segment; characteristic
venation of genus showing clearly on sheath.

Leg sheaths long, extending to beyond mid-length of fourth abdominal segment; fore legs shorter than the others.

Abdomen elongate. Abdominal segments 2 to 7 near posterior margin with a transverse
row of short, stout spines which are interrupted only near pleura; pleural area with fouror five spines; ventral and dorsal segments with numerous spines; dorsal row of spines moredistant from posterior margin of segment than the other areas; dorsum of eighth segmentwith four powerful, chitinized lobes directed dorsad and caudad; posterior margin of theselobes with about five or six small teeth; two spines near base of pleural region on segment 8.
Male Cauda consisting of two blunt sheaths, lying between posterior pair of lobes describedabove.

—South Mountains, near Myersville, Maryland, November
Neanotype.
Tallulah Falls, Georgia, June 17, 1910.
Paratypc.
Cabin John Bridge, Maryland, May 31, 1900.

Larva.

—
—

6,

1916.

The Crane-Flies of New York— Part II

993

Genus Aeshnasoma Johnson (Gr. a dragon fly + body)
Aeshnasoma Johns.

1909

115-116.
Proc. Boat. Soc. Nat. Hist., vol. 34, p.

Aeshnasoma is a monotypic genus found in northeastern North America.
and may be congenerous with it.
It is unquestionably close to Longurio

The
its

Aeshnasoma

type,

rivertonensis

Johns.,

isapparently very local in

distribution.

undoubtJohnson (1906 1-2) described an unknown tipuline larva whichJuneon10, 1900,
found
was
larva
edly pertains to this species. The
laboratheintobroughtwasIt
a cold spring at Riverton, New Jersey.
:

inthe cold spring (about
tory but could not be reared, the change fromwhen fully extended
larva
The
fatal.
60° F.) to warmer waters being
yellowish white inwasIt
length.
inabout 45 millimeters

measuredwith its contents beingcolor and was translucent, the alimentary canaland figures thedescribes
Johnsonclearly visible thru the thin skin.

LXXXIX, 492) of this genus. The
the adults of Aeshnasoma
Longurio,
doubtfully referred topeculiar branched anal

larva

was

gills

(Platebeing undescribed at that time.
the taking
In a later paper (1907-12 [1909] 115-116) Johnson mentions
adult flies
theofcapture
the
July
20,
of several more larvae in 1902, and, on:

on which the genus and species are based.
The only larva that was preserved was kmdiy sent to the writer forLongurio, bothstudy by Mr. Johnson. It is undoubtedly very close tobranched anal
the
and
disk
spiracularshowing the same peculiar
generaa condition that is found nowhere else in the Tipulidae
known to the writer.

gills,

Genus Holorusia Loew
1863

Larva

—

Holorusia Loew.

so far as is(derivation obscure)Berl. Ent. Ztschr., vol. 7, p.

1.

fringed with long
Spiracular disk surrounded by six moderately elongate lobes
between spiracles
disk
Hnes;
black
capillary
with
lobes
ventralandof lateral

hairs; inner facedusky.

Anal gills six.

to apical point.

Mandible small, with a single dorsal and ventral tooth in addition

Antenna with a conical apical papilla.

Mentum seven-toothed.

Hypo-pharynx six-toothed.
breathing ho^ns rather
Pupa. - Sheaths of maxillary palpi recurved at tips. Pronotal
Armature of abdominal segments almost as inshort and stout, the long apices flattened.
numerous (fourteen to twenty-four);Prionocera, but the posterior rows of spines more
one behind the other. Cauda
pleurites with three spines, the two on posterior ring situatedwith six stout dorsal lobes.

Charles Paul Alexander

994
Holorusia
flies,

isa small gonus (about ten species) of

only one of which

New World crane-— the genotype, Holorusia ruhiginosa — Nearctic.
isThis species and Longurio are the largest Nearctic crane-flies. The
anatomy of th(^ "giant crane-fly" has been described in some detailby Kellogg (1901, a and b) and by Comstock and Kellogg (1904). The
immature stages are spent in moist earth. The genus is undoubtedly
closely related to Prionocera, and, presumably, to the Old World genus
Ctenacroscelis Enderlein.

Holorusia ruhiginosa Loew
1863
1888

Holorusia ruhiginosa Loew. Berl. Ent. Ztschr., vol. 7, p. 1.
Tipida (Holorusia) grandis Bergr. Ent. Tidskr., vol. 9, p. 140.

Holorusia ruhiginosa is widely distributed thruout the western UnitedStates and Canada.

A number of larvae were taken by H. Morrison

near Stanford University, California, on February 22, 1915.

They wereshipped to the writer at Ithaca, New York, where the species was reared.

The massive larva is used for purposes of dissection in the entomologicalcourses at some of the western universities.
Larva.

— Length, 50-00 mm.
Diameter, 6.2-6.4 mm.

Coloration, dark greenish brown.

Form stout, subterete.

Integument covered with dense,

short,

erect,

black hairs.

A

few weak and delicate setae, two on dorsum and on venter of each abdominal segment; twolong setae on lateral margins of posterior rings.
large,

Spiracular disk (Plate XC, 496) moderately

surrounded by six stout, elongate lobes which are similar to those of Prionocera but

are stouter and less digitiform; ventral lobes a little the longest, dorsal lobes a little the shortest;allthe lobes capable of close approximation, completely protecting spiracles; lobes

fringed with long black hairs which are longest near apices, shorter

between lobes; ventral

and lateral lobes with a delicate black line down inner face, these Hues barely indicatedon dorsal lobes; remainder of disk and lobes dusky. Spiracles very large, circular, separated
by a distance a Uttle less than diameter of one. Anal gills six, short, slender, the twoanterior

gills

of either side united basally, posterior pair simple.

Head capsule of the usual massive tipuline type, prefrons running caudad as a narrow
point, lateral plates broad.
Labrum broad, with a densely hairy lobe on either side. Mentum (Plate XC, 493) with a prominent median point; behind it on either side three flattened
middle tooth more acute, the outermost formed by
Hypopharynx (Plate XC, 494) about six-toothed, the intermediate
teeth with a large notch between.
Antenna (Plate XC, 495) with basal segment very longand slender; principal apical papilla conical; a number of small hyaline sense pegs. Manditeeth, the innermost the broadest, thefusion of two small teeth.

The Crane-Flies of New Yorkble slender, with only two teeth, a stout dorsal tooth— Part II

995

and a single flattened ventral tooth.

Maxilla small, lobes covered with short, dense hairs.

Pupa.

— Length, 32 mm.
mm.
Width, d.-s.,
Depth, d.-v., 5 mm.
.'5Color brown; flattened lateral margins of abdomen broadly yellowish.
depressed, with lateral margins flattened, carinate.

Cepharc

crest represented by two low, parallel ridges, provided with one or two tiny .setae.

LabrumThoraxlarge,

shapedterete;abdomentumid, transversely wrinkled, the blunt apex completely separating the diamondlabial

lobes.

Maxillary palpi stout, extreme tip recurved.

elongated, extending some distance beyond maxillary palpi.

Antenna moderately

Pronotal breathing horns rather

short and stout, finely ringed, the rather long tips flattened, about equal to one-fifth lengthof

entireorgan.

Mesonotum convex(PlateXC,

497),

with transverse anastomosing

Wing sheaths reaching end
Leg sheaths extending just beyond base of fourth abdom-

wrinkles; on either side of median line behind, a blunt tubercle.
of second
inal

abdominal segment.

segment; fore tarsi short, middle tarsi a little longer than hind tarsi.

Abdominal segments with the usual basal and posterior rings; armature almost as inPrionocera; tergites with posterior row of spines numbering between twenty and twentyfour on intermediate segments, near anterior lateral angle two spines, basal ring unarmed;
pleurites with one setiferous spine on basal ring and two similar spines on posterior ring,
one placed considerably behind the other; sternites armed similarly to tergites, but the posterior row of spines larger and somewhat fewer in number (fourteen to eighteen) posterior ringon either side median line near base with two spines, the innermost very large and powerful.
Male Cauda with ventral lobes blunt, each armed with a slender black spine near posterior margin; dorsal surface of cauda almost as in Prionocera, armed with six stout lobes,
which here are shorter and stouter, with tips abruptly narrowed; lateral margin of segment
8 with a stout lobe on either side, each terminating in a cylindrical spine. Female cauda
similar to male cauda, dorsum with the same six lobes; acidothecae short, tergal valvesslightly exceeding the more blunt sternal valves.
;

—
—
—

Ncpionotype.
Stanford University, California, February 27, 1915.
April 5, 191.5.
Neanotype.
Paralypes.
Larvae and pupae with typed.

Genus. Prionocera Loew (Gr. saw + horn)
1S44
1863

Prionocera Loew. Stett. Ent. Zeit., vol. 5, p. 170.
Stygeropis Loew. Berl. Ent. Ztschr., vol. 7, p. 298.

— Spiracular disk surrounded bysix long, finger-like lobes fringed

with long, deh-cate hairs; each lobe with a capillary black hne down middle of inner face.

Spiracles large.

Larva.'

Anal gills unbranched. Mentum seven- to nine-toothed.
Mandible with about two dorsal and three ventral teeth.
Pupa.-— ]\Iaxillary palpi recurved at

tips.

Hypopharynx

five-toothed.

Pronotal breathing horns very elongated,

unequal, the longer one about half length of body; horns at tips split into long flaps.
inal tergites

Abdom-

with a posterior transverse row of fifteen or fewer spines, and two small spines

Charles Paul Alexander

996

near anterior lateral margin of posterior ring; pleurites with a spine on basal ring and two
transverse spines on posterior ring.

Cauda with six strong dorsal lobes.

Prionoccra is a small genus (about a dozen species) of usually far northern
of

flies

Americasomberiscoloration.

The only

Prionocera fuscipennis,

species founddiscussed

stages arc somewhat similar to those of Holorusia.

easterninbelow.

North

The immature

The apparent similarityof the pupa to that of Longurio is probably not indicative of a very close
relationship.

The immature stages have been discussed but little in the literature.
The 'Tipula sp. No. 1" of Malloch (1915-17 b: 199-200) refers to P. fusciThe immature stages of a species supposed to be P. parri (Kirby)
pennis.
have been discussed and figured by the writer in his report on the CanadianArctic Tipulidae (Alexander, 1919 c: 19c-20c).

The name Stygeropis has been in use for many years under the belief
name Prionocera was preoccupied in the Coleoptera.

that the earlier

Dr. Bergroth states that this is not so and that Prionocera should be
used.

Prionocera fuscipennis (Loew)
1865

Stygeropis fuscipennis Loew.

Berl. Ent. Ztschr., vol. 9, p. 129.

Kennedy found two cast pupal skins among Sparganium stems
New York, on November 20, 1916. Several
larvae had been found here in the preceding July, and some others were
C. H.

in Ringwood Hollow, Ithaca,found on Junehillside,4,

1917 (No. 106-1917), in a cat-tail swamp near Bool's

Ithaca, where they were associated with the characteristic helo-

phytic crane-fly fauna (Bittacomorpha, Rham.phidia flavipes, Pseudolimnophila luteipennis, Pilaria recondita, Tipula

tricolor,

Malloch's material was taken in Wisconsin inMay.

and other species).
Dr.

Needham hasreared the species near Lake Forest, Illinois.
Larva.

— Length, 18-22 mm.
Diameter, 2-2.2 mm.

Coloration dark brown, in some cases with a pale dorso-median stripe.

Form terete, tapering gradually to anterior end of body.
elongate setae.

Spiracular disk (Plate

Segments with several scattered

XCI, 502) surrounded by six long, finger-like lobes

which are delicately fringed with long hairs; ventral lobes considerably the longest; laterallobes a little larger and stouter than dorsal lobes; all the lobes broadly margined with darkbrown, these marks expanding at inner ends; on ventral lobes, lateral margin expanded at inner

The Crane-Flies of New York— Part II

997

end and continued across disk, meeting its fellow of the opposite side between spiracles;
dorsal margin of lateral lobes touching spiracles; each of the lobes marked with a capillarydark brown hne down center of inner face, this beginning near ends of lobes and extending
almost to base; lobes fringed with long hairs, these very tiny near base, longer near tips oflobes,

but scarcely,

if

rated by a distance a

at

all,

littleinterrupted between lobes.

greater than diameter of one.

Spiracles large,

circular,

Anal gills six,

long,

sepa-

slender,

unbranched.

Head capsule of the usual tipuline type.
on lateral lobes longer.

Labrum covered with dense, short hairs, those

Mentum (Plate XCI, 498) u.sualiy seven-toothed, in some casess nine-

median tooth slender, lateral teeth flattened, subacute. Hypopharynx (Plate
XCI, 499) narrow, five-toothed. Antenna (Plate XCI, 500) long and slender; first segment
toothed;a little enlarged near base; at apex several tiny sensory papillae;

three ventral teeth; prostheca large.

Pupa.

auditory plate near baseMandible (Plate XCI, 501) moderately large, with two large dorsal and aboutof segment.

Maxilla small; palpus large, cylindrical, apex truncated.

— Length, 15-22 mm.
Width, d.-s., 2.4 mm.
Depth, d.-v., 2.6 mm.

Coloration dark brown; lateral and posterior margins of abdominal segments paler.

(In

old specimens the general coloration is very dark brown; in younger specimens the abdomen
ismore or less distinctly lined with brown.)
Thorax subterete; abdomen depressed, lateral margins flattened.

point narrow.

Labial lobes broad, slightly separated on medianLabrum broad, apicalline.

Maxillary palpi

Antenna moderately elongated, extreme tip darkened. Pronotal breathing horns long and slender, unequal in length, the longer about 9 or 10 mm.
in length, the other 6 mm., at tips split into divergent flaps (Plate XCI, 504) almost as in
the hexatomine genera Pseudolimnophila and Pilaria, which live in the same muddy situations. Mesonotum (Plate XCI, 503) transversely wrinkled. Leg sheaths reaching posterior
margin of third abdominal segment; hind legs the longest; middle legs a little shorter thanshort, stout,

apex recurved.

fore legs.

Abdominal segments divided into a basal and a posterior ring; tergites with basal ring
unarmed; posterior ring with a subterminal transverse row of short spines, with a few setae
located on lateral face of some of the spines; on second tergite, four to six spines, on tertwo small spines with setae near anterior lateral angle
and two such spines
on posterior ring located side by side; sternites with the basal ring unarmed, posterior ringarmed similarly to that of tergites; in addition to posterior row of spines, a pale oval area
gites 3 to 7, three to fifteen spines;of posterior ring; pleurites with a small setiferous spine on basal ring,

Male cauda
XCl, 505) with four powerful lobes on dorsum of last segment, directed dorsad andon either side of midventral line, each with two transversely placed spines.
(Platebetween anterior pair
two additional slender lobes, each ending in two acute spines.

slightly caudad, lobes bearing three or four small spines before tips;of lobes,———

Nepionotype.
Ringwood Hollow, Ithaca, New York, July 20, 1916.
Neanotype.
Cast pupal skin, type locaUty, November 20, 1916.
Paratypes.
Larvae and pupal skins, type locality.

Charles Paul Alexander

998

Genus Tipula Linnaeus (Lat. a water-strider)
1758
1842
1864
. 18S7
1894
1894
1894
1916
1916
1916
1916

Syst. Natur., ed. 10, p. 585.
Mag. Zool. Ins., pi. 106.
Atti dell' Institut Veneto, ser. 3, vol. 9, p. 218.
Anmnido'iAcra Lioy.

Tipula Linn.

Pterelachisiis

Rond.

Oreomijza Pokorny.
Wien. Ent. Ztg., vol. 6, p. .50.
Manapsis Scudder. Proc. Amor. Philos. Soc, vol. 32, p. 222.
Rhndinohrochus Scudder. Proc. Amer. Philos. Soc, vol. 32, p. 223.
Tipulidea Scudder.
Proc. Amer. Philos. Soc, vol. 32, p. 238-239.
Nippotipxda Mats. Thous. Ins. Japan, add. 2, p. 457-458.
Plahjtiprda Mats.
Thous. Ins. Japan, add. 2, p. 459.
Yamaiolipida Mats. Thous. Ins. Japan, add. 2, p. 461-462.
Togotipida Mats. Thous. Ins. Japan, add. 2, p. 4G5.

— Form generally stout, terete or nearly

Larva.-

almost invariably with a definite chaetotaxy.

so.

Integument with pubescence andSpiracular disk surrounded by si.x or rarely

eight lobes, simple, or in certain species (as T. abdominalis)more or less split at their tips.

Spiracles small and widely separated (in T. abdominalis), or in other species large and rather

Analclose together.

gills

almost invariably present, with

branches simple, not pinnate.

six

or eight

Head capsule compact and massive.

branches, theseLabrum broadly trans-

Mandible usually small, with few teeth, ventral cutting edge with usually two or
Maxilla rather complicated, of the generalized tipuline structure. Antenna
usually elongated, basal segment two to four times as long as it is thick, stouter in speciesverse.

three teeth.

wood. Mentum with seven to ninemargin usually with five teeth.

living in decaying
plate, anterior

teeth.

Hypopharynx a flattenedsetae.

— Form generally stout. CephaHc crest lacking or very small, with rudimentary
Mouth parts as in the subfamily, sheaths of maxillary palpi strongly recurved

at tips.

Pronotal breathing horns subequal in length, short, stout, usually straight, tips

Pupa.-

but little expanded.
(asT.

triviltata)

Mesonotum transversely wrinkled, in some wood-inhabiting species
Wing sheaths and leg sheaths

with about four conspicuous tubercles.

moderate in length. Abdominal armature usually strong, each segment with a posterior
row of four to twenty spines; in some species a basal ventral row of spines on posterior ringof segments.

Cauda with dorsal armature of four powerful lobes; eighth segment adding,as a rule, ten spines, of which six are ventral and lateral in position, and two or four are dorsal;

dorso-median pair lying between anterior pair of lobes of cauda, as discussed above, andlacking or very reduced in some wood-inhabiting species {T.

Iriviitala).

Lateral abdominal

spiracles lacking or merely vestigial.

Tipula is the largest genus of crane-flies, comprising a vast assemblage

and seven hundred described forms) which arefound on all the continental areas of the world but are few in the Australof species (between six

asian region and apparently lacking on many of the lesser oceanic islands.

The genus is one of extreme interest, and its study will require many
years of conscientious application.

Subaptcrous species are not rare inmany are far northern forms, others are coastal species, while a few live inland and under influences that make it difficult tothis group, of which

explain their subapterous condition.

The Crane- Flies of New York— Part II

999

The immature stages of the various species are diverse in their habits,
ranging from species that are nearly if not quite aquatic, thru the majorityof the known forms which Hve in generally moist earth or mud along themargins of water bodies, to still others that live in the semi-decayedwood of prostrate tree trunks.

Mellor (1919:64) has recorded Tipula
So far as is known, the larvae are herbivorous, tho they will eat animal food under stress (as described by Patterson [1908] for Tipula oleracea, which feeds in considerable numbers onearthworms).
In Europe, a great number of life histories in this genus have been
worked out in commendable detail, mainly thru the efforts of Beling,
who discusses no fewer than thirty species. His descriptions give a clear
idea of the range in structure and habitat to bo expected in the genus.
The number of lobes surrounding the spiracular disk varies from four
larvae as breeding in manure.

(apparently) in T. selene and related forms, to as
suhnodicornis.
tipuline

Practicallyallof theknown

many as eight inspeciesT.

show the normal

number of lobes, six.

A summary of the larval habitats of the Palacarctic species .s as follows
1. Species living in saturated earth along watercourses or in debris at the water's edge,or species that are aquatic
Tipula fulvipennis de Geer, lateralis Meig., lunata Linn., maxima
Poda, variicornis Schum., variipennis Meig., viltata Meig.
2. Species living in earth, usually in woods, underneath a mold of leaves or coniferous
needles
Tipula caesia Schum., dilatata Schum., fulvipennis de Geer, hortensis Meig.,
hortulana Meig., nigra Linn., nubeculosa Meig., ochracea Meig., pabulina Meig., paludosa
Meig., pruinosa Wied., scripta Meig., selene Meig., truncorum Meig., unca Wied., variipennis
Meig., viltata Meig.
3. Species living in earth in gardens, pastures, or meadows, usually beneath turf
Tipulairrorata Macq., luteipennis Meig., nigra Luin., ochracea Meig., oleracea Linn., paludosa Meig.,
pruirwsa Wied., suhnodicornis Zett., truncorum Meig., vernalis Meig.
4. Species living in or beneath cushions of moss or in earth overgrown with a mossy
covering
Tipula dilatata Schum., hortulana Meig., marmorata Meig., pagana Meig.,
peliostigma Schum., pruinosa Wied., rufina Meig., signata Staeg., truncorum Meig., uma

—

—

—

—

Wied.
5.
6.

—

Species living underneath moss on logs
Tipula irrorata Macq.
Species living in decaying wood
Tipula flavolineata Meig., irrorata Macq., truncorum

—

Meig.

Bouche describes T. lunata and T. ochracea as living in decaying willow
wood, and Sopotzko records T. flavolineata as injuring clover; but theserecords are presumably based on mistaken identifications.
Comparatively few of the eastern American species have been reared,
and it is not considered advisal)le to attempt a key to the larvae or thepupae atthis stage ofknowledge of the subject.

Such a key would

Charles Paul Alexander

1000

include but a fraction of the possible species and would be of little value.
It will require the careful rearing of species for many years before aworkable key to the immature stages of the eastern species of the genus

The characters that will prove of greatest value incan be produced.

the separation of the larvae and the pupae of the species of Tipula are as
follows:

Larvae(These are rarely lacking, and the number and arrangement of the branches,
their form, and their function, are of primary importance.)
2. Spiracular disk.
Number of lobes surrounding disk and whether they are simple orbranched; character and nature of fringe of hairs around disk, if such is present; size, shape,and distance apart of spiracles; markings on inner face of disk and lobes.
3. Chaetotaxy.
Arrangement, length, and number of setae on segments.
4. Body form.
Terete, subdepressed, or flattened ventrally only; clothing of pubescence,and pattern formed on dorsum.
5. Head capside.
(The head is remarkably uniform thruout the group, a condition to
be expected in a group so compact as Tipula.) Shape of mentum and hypopharynx, andnumber, size, and shape of teeth along their anterior margins; shape of antenna, and other1.

A7ial gills.

details of head.

Pupae
1.

General form, whether terete or depressed.

2.

Mouth parts.

•

size, and form.
Armature of mesonotum.
5. Wing sheaths and leg sheaths.
6. Spines on abdominal segments, their size and number; whether lacking or present atbase of posterior ring of sternites; arrangement and number of pleural spines.
7. Cauda, shape of genital sheaths, armature of dorsum, and ventral margin of eighth
3.

Pronotal breathing horns, their relative length,

4.

eternite.

Descriptions are given in the following pages of about ten life histories
which are entirely new or have been insufficiently considered elsewhere.
A few notes on certain other species that have been observed in the past
few years may be added here:
Tipula cayuga Alex.
in association(page 781).

A conspicuous yellow larva, living in organic earth beneath leaves,with Bittacomorphella jonesi and other forms which are discussed elsewhere

The pupal duration is slightly over seven days.
Found living in rather dry earth beneath leaves in shaded woods

T. angustipennis Loew.

(Lawrence, Kansas, Mrs. C. P. Alexander).
T.

umbrosa Loew.

Occurs in garden soil in company with the larvae of Tipula hicornis

Forbes.
T. fuliginosa Say.

Reared from larvae living in debris under the nest of a turkey vulture(Jackson Island, Maryland,

May 23, 1913, R. C. Shannon).

and T. tricolor Fabr. In saturated mud in marshy or swampy situations.
2'. tephrocephcUa Loew.
A large larva, nearly if not quite aquatic in its habits.
T. sayi Alex,

The Crane-Flies of New York— Part II

1001

The life histories of other Nearctic species are recorded in the summary of hterature on page 980. These are as follows: Tipula arctica
Curt. (Nielsen, Alexander), T. eluta LoeW (Hart, Malloch), T. cunctans
Say (Hyslop, Malloch), T. bicornis Forbes (Forbes), and T. ultima Alex.
(Needham, Caudell). T. arctica, according to Nielsen (1910:57-59), wasfound commonly in eastern Greenland. The immature stages were dis-covered in circular holes from two to three centimeters deep in the ground,
The pupae
especially beneath tufts of Casdope tetragona (L.) D. Don.
as early
found
were
cases
pupa
empty
and
June,
of
end
the
were found at

According to Nielsen, the larvae requireas the 25th of the same month.

two years to attain their growth.
Tipula (Trichotipula) oropezoides Johns.
1909

Tiyula oropezoides Johns.

Proc. Boston Soc. Nat. Hist., vol. 34, p. 131-132.

Larvae of Tipula oropezoides were first found on March 30, 1917, living
beneath saturated moss in Needham's Glen, Ithaca, New York, where
they were associated with larvae of Dicranomyia badia, Penthoptera
Tipula collaris, and other species. Numerous additional larvae
were found in the same locality on April 18, 1917. Some of these wereplaced in rearing and emerged on May 6. The larvae are nocturnal in
albitarsis,

becoming
their habits, being very sluggish and retiring during the day but
active after sunset.

The adult flies bear a strong resemblance to species of the genus Oropeza,
They may often bewith which they arc sometimes found associated.
swept from rank herbage in cool Canadian woods.
Length, 16.5-17 mm.
Larva.
Diameter, 1.8-2 mm.

—

Coloration above, a deep velvety brown with mottlings of paler; on basal ring
ventertransverse pale spots, posterior ring less regularly marked; pleura andof tergitea

medianhairs,(The dark markings on the dorsum are produced by patches of dark-coloredpale.
which cover the body densely in places.)
Form subterete. Integument with an abundant pubescence, longest on dorsum. Chaetosetae, three on
taxy as follows: dorsum (Plate XCIII, 516) on po.sterior ring with six stout(Plate XCIII,
segments
ventral
inner
seta;
the
closertolittleside, the middle seta asixeitherhttle more separated.
517) with four setae, two anterior and two posterior, the latter asubequal lobes, theirapproximately
sixbysurrounded
Spiracular disk (Plate XCIII, 518)
rounded spot, largest on
inner faces heavily lined with dark brown; at tip of each lobe a palespiracles; lateral
ventral lobes and here with a sensory bristle; lateral mark not reachingbrown
and dorsal lobes slightly paler medially; above and below each spiracle a transverse

Charles Paul Alexander

1002
line;

on disk, between spiracles, two indistinct dusky spots; lobes fringed with rather short,pale hairs which are narrowlyinterrupted between lobes.
Spiracles irregular, roughly
Anal gills four, slender, posterior pair the larger (Plate XCII, 506).

triangular.

Head capsule as in genus.

Labrum and maxilla very densely fringed with long golden-

Mentum (Plate XCIII, 513) with two flattened lateral teeth, the medianpoint elongated; mentum very deeply split behind.
Hypopharynx (Plate XCIII, 514) withyellow hairs.

but three evident teeth, the lateral teeth very broad, flattened.

(In some specimens thesehypopharynx appears merely
Mandible (Plate XCIII, 515) withteeth are all very blunt, so that the anterior margin of the

Antenna with apical disk very flattened.

crenulate.)a dorsal tooth and a powerful ventral tooth.

Pupa.

— Length: male, 12 mm.; female, 12.5-13 mm.
Width, d.-3. male, 1.6-1.7 mm.; female, 1.<S-1.9 mm.
Depth, d.-v.: male, 1.6 mm.; female, 1.7-1.8 mm.

':

Coloration dark brown; dorsum of thorax and abdomen, and face, more reddish brown.

General features as in Tipida

andslender,dark-colored,

Form slender.

collaris.

divergent at

tips.

opposite apex of second abdominal segment.

Pronotal breathing horns long

Antenna elongate. Wing sheaths ending
Leg sheaths long, extending to beyond mid-length of fourth abdominal segment.

INIale cauda with dorsal lobes of genitalia short, blunt;caudad into slender, blunt lobes which are transversely wrinkled,separated by a U-shaped notch, at base on outside with a prominent spine.
Female ovipositor elongate; dorsal valves narrowed to the blunt tip; ventral lobes stout, a little shorter
than dorsal valves, tips strongly divergent; the six dorsal lobes of cauda spinous-tipped,

ventral lobes produced

sharply pointed.

Nepionotype.~Uha.c&, New York, April 18, 1917.
With type. No. 7 1917.

—— Numerous larvae and pupae.

No. 6-1917.

Neanotype.
Paratypes.

Tipidacollaris

1823

Say

Tipula collaris Say.

Journ. Acad. Nat. Sci. Phila., vol. 3, p. 23.

of Tipula collaris, a common vernal crane-flj^
beneath saturated moss {Amblystegium irrignwn[Wils.] B. & S.) in Needham's Glen, Ithaca, New York, on April 17,
1917.
Their associates are noted under the account of T. oropczoides(page 1001). In the same moss areas occurred numerous small red-backed
salamanders (Plethedon cinereus), which probably fed on the insect deni-

Larvae and pupae

occurredfrequently

zens of the place.

Specimens emerged in the writer's breeding jars as late

as May 10.

The adults are on the wing during April and May, some persisting into
The life history undoubtedly requires

early June in cool northern woods.

a year for its conii)I(^tion.
Larva.

— Length, 21.5-25 mm.
Diameter, 2.5-3.5 mm.

The Crane-Flies of New York— PartII

1003

Coloration pale dusky, darker above; dorsum behind with two broken lines which arepleuradivergent on each annulus, those of anterior annulus made up of three circular spots;
lines;
withindistinct
pale,
uniformly
almostventerstripe;brown
darkwith a conspicuous
anterior part of thoracic segments darker.

(In life the color is rich reddish brown, and thepleural stripe is not evident.)

Body covered with a delicate, pale pubescence, in addition to the usual
Chaetotaxy as follows: tergites (Plate XCIV, 521) with a transverse row of six setae

Form terete.
setae.

on posterior ring before margin, the two innermost solitary, each lateral pair closely
approximated; a solitary seta on extreme lateral margins of tergite, at margin of dark pleuralbasal ring with a single seta, posstripe, and at about midlength of posterior ring; pleura on
the others; sternites
terior ring with a group of about three or four setae, one larger thanwith four widely separated setae on posterior ring, the lateral pair a little nearer posterior
margin than the median pair. Spiracular disk (Plate XCIV, 522) surrounded by six lobes

which are moderately slender; ventral pair a little longer, dor.sal pair a little shorter; all theblack capillary
lobes broadly margined with pale brown; ventral lobes having in addition aextending from tips backward to beyond midlength of lobes, this line broadest at tip,
gradually narrowing, and becoming paler toward base of lobes; below each spiracle, twolineconspicuous black dots; lobes fringed with numerous rather long hairs. Spiracles large,
separated by a distance a little greater than diameter of one. Anal gills six, one pair muchgill.
shorter than the others, consisting of a basal branch of the anterior
XCIV, 519) very
(PlateMentumtype.
tipulineusual
thesmall,
ofrathercapsule
Head

Hypo-

broad, anterior margin almost transverse, seven-toothed, median point the longest.
pharynx (Plate XCIV, 520) with five teeth, which are very short and blunt giving anterior

margin a deeply crenulated appearance; before hypopharynx a rounded lobe which is densely
covered with six short, blunt, chitinized points, this being probably the prementum. Antenna
Mandible small, withelongate-cylindrical; apex with apical disk very small, button-like.
about one dorsal and two ventral teeth.

Maxilla slender, densely hairy; palpus subglobular,

with several small, hyaline papillae.

Pupa.

— Length, 17-19 mm.
Width, d.-s., 2.S-3.2 mm.
Depth, d.-v., 3-3.3 mm.

Coloration brown; wing sheaths, except in older individuals, pale; pleural region of abdomen

Ught yellow; abdominal incisures often pale.
Head rather small. Cephalic crest low and indistinct, with tiny setae.

Labrum broad,

Labial lobes oval, contiguous at inner end. Ma.\illary palpi strongly recurved
Antenna slender, moderately elongated, extending some distance beyond wing

apex pointed.
at tips.
root.

tened,

Pronotal breathing horns equal in length, rather short, the moderately long tips flatsmooth; two small, approximated setae on either side of median hne. Wing sheathsegment; venation dis(Plate XCIV, 523) extending just beyond end of second abdominal
fore tarsi consegment;
abdominal
fourthofbasebeyondextendingsheaths
Leg
tinct.
siderably shorter than the others.

Abdominal segments with armature of posterior ring weak, the spines very short and stout,with a few setae; maximum number of spines on the tergites about twenty; lateral anterior
spine, andangle of posterior ring of tergites with two small spines: pleurites with one basal

Charles Paul Alexander1004

three setiferous spines on posterior ring, arranged transversely;,

spiracles very rudimentary,opposite base of posterior ring; sternites with spines slightly more numerous and stouter;near base of posterior ring two transverse spines on either side median line, the outermost

Male cauda on dorsum with six lobes; the four posterior ones stout, with tips

setiferous.

spinous or those of ventral pair slightly bifid; the two anterior median lobes shorter andmore

slender;eighth sternite with four large spines about equally spaced; eighth pleurite

with a large, powerful, acutely tipped lobe, and two or three smaller dorso-lateralabove base of middle pair of dorsal

lobes.

.spines

Female cauda almost the same as male cauda,

due to the blunt terebral sheaths of this species.

———

Nepionotype.
Ithaca, New York, March 30, 1917.
With type larva.
Neanotype.
Numerous larvae and pupae, March 30 to April 18, 1917.
Paratypes.

Tipula nohilis (Loew)
1864

Pachyrrhina nobilis Loew.

Berl. Ent. Ztschr., vol. 8, p. 62.

Larvae of Tipula nohilis wore found in wet moss and beneath decaying
An adult emergedon July 1. A fully grown pupa found on July 11 attempted to transform,
but died after two hours without being able to extricate itself from thewjtch-hazel leaves at Orono, Maine, on June 17, 1913.

pupal case.

The adults, which strikingly resemble some species of Nephrotoma, flysomewhatlater

than does T. collaris, but both species

may

be taken

together in early June.
T. nobilis
Larva.

isvery similar in all respects to T.

collaj'is.

— Length, 20 mm.
Diameter, 2.9-3 mm.

Coloration, reddish brown.

Spiracular disk as in T. collaris, the brown lateral margin to the lobes a little paler.
gills

as

Anal

shown in Plate XCII, 507.

Head capsule almost as in T. collaris. Mentum with apical point elongate, with three
Hypopharynx with five moderately acute teeth. Antenna withblunt teeth on either side.

a blunt conical papilla, larger and more conspicuous than in T collaris.
very blunt.

— Length of cast pupal skin, about 18.5 mm.

Pupa.'

Pupa very similar to that of T. collaris.
Nepionotype.— Orono, Maine, June 19, 1913.
Neanotype.

— With type.

No. 40-1913.

Tipula hella Loew
1863

Tipula hella Loew.

Berl. Ent. Ztschr., vol. 7, p. 291-292.

Mandible with teeth

The Crane-Flies of New York — Part II

1005

Tipida bella is a common species, flying thruoiit the summer.
Larvaeand pupae are not rare in sandy or loamy soil along streams. A larva
taken on April 28, 1917, in gravel at Ithaca, New York, where it was
It (mierged
associatt^d with larvae of Hexatoina, was placed in rearing.
On May 27, 1913, three fully colored pupae
as an adult male on May 15.
were found along the sandy banks of Fall Creek, Ithaca, in association
with Eriocera spinosa. They emerged as females on May 29 and 30.
Larva.

— (The descriptionisfrom fleld notes on the larva mentioned above.)

Length, 20-25 mm.
Coloration light grayish brown, with a slight reddish cast most noticeable on venter;dorsum with two narrow, almost continuous, dark brown hnes, these lines subparallel atanterior part of each segment, then strongly bellied out, and then parallel but finallydivergent.

Spiracular disk surrounded by six rather short lobes; ventral lobes with a linear, rather'pale,

brown mark, and a few sensory bristles at tips; lateral lobes with ventral margin linedwith brown; dorsal lobes with both margins feebly bordered with brown; two small brown
dots below each spiracle.

Anal gills six, very long and slender.

Pupa.

— Length of cast pupal skin, about 24 mm.

Pupa

similar to

pupae of other species of genus. Pronotal breathing horns short,narrowed to tips. Spines on abdominal segments rather large, especially onsternites; spines on base of posterior ring of sternites small but evident.
Female cauda
with dorsal valves long, pointed; sternal valves shorter. Cauda with the usual six lobes
on dorsum, the four posterior stout, divergent, spinous-tipped, the anterior median pairmuch smaller; venter of segment 8 with three strong spines on either side, gradually smallerfrom lateral spine toward mnermost spine.
cylindrical,— Ithaca, New York, reared May 25, 1917.
— Pupal skins, type locaUty, May 1914 (No. 42-1914); August, 1911; May

Neanotype.
Paratypes.
25, 1917; etc.

Tipula caloptera
1863

5,

Loew

Tipula caloptera Loew.

Berl. Ent. Ztschr., vol. 7, p. 292.

The vigorous larva of Tipula caloptera is one of the largest and mostThe larvae live in rapid- or slow-flowing streams
either in the water among debris and under stones, or in the sand, gravel,
striking in the family.

or

mud in very close proximity to the water.

Here they are associatedwith the larvae of various species of Eriocera, Erioptera annata, Tabanus,
Atherix,

and other forms.

exceedingly lotic situations.

Oftentimes they are found in deep water in
A larva placed in rearing on April 19, 1917,

emerged as an adult female on May 13

Charles Paul Alexander

1006

This is evidently the larva taken by Dr. Needham in the Adirondacks
and referred by him with some dou])t (Needham and Betten, 1901:57557G) to T. ahdominalis.

This dubious reference has creattnl considerable

confusion ever since the species was figured on the cover of EntomologicalNews under the facetious name " Quisnam sexcaudatusf " Malloch (191517 b: 200-201) mentions the same larva under thename T. ahdominalis.

As stated elsewhere, the larva of ahdominalis is very different.
Larva.

— Length, 45-55 mm.
Diameter, 4.8-6.8 mm.

Coloration above, dark brown or brownish green; segments beautifully marked with small

white spots, especially anterior segments; a broad, dark brown, median stripe, and a moreor less distinct pale lateral stripe (in preserved specimens the pleura is usually dark brown,

more distinct behind, with numerous pale white dots); sternum dark greenish.

(In olderspecimens the color is very dark and the pattern is more or less obliterated.)

Form stout, terete.

Body smooth, segments with indistinct posterior tubercles.

Chaeto-taxy very weak, a fevv weak pleural setae on posterior ring, sternal and tergal setae minute.
Spiracular disk rather small, surroundedby six subequal, moderately narrow, lobes which

are fringed with short hairs; margins of disk and lobes somewhat as in T. bella, each lobe witha delicate capillary brown line; two brown spots beneath each spiracle.
separated by a distance about equal to twice diameter of one.

Spiracles small,

Anal gills six, very long andslender (Plate XCII, 508).

Head capsule as in genus.

IMentum broad; anterior margin nearly transverse, with threesubacute teeth on either side, median point not conspicuously elongated.

— Length of cast skin, about 32-35 mm.

Pupa.-

Characters almost as in T.

bella.

Pronotal breathing horns short, cylindrical, tips not

Abdominal spines prominent, projecting, few in number, on intermediate tergites 9 or 10; pleurites with a single strong spine on each ring; sternites with a posterior row
of seven or eight strong spines; on segments 5 to 7 two strong spines at base of posterior
expanded.

ring, those of seventh segment the largest.

Cauda almost as in T. bella.

Nepionotype. — Ithaca, New York, April 26, 1917.

No. 16-1917.

Neanolype.
Paralypes.

locality.

— Cast pupal skin, reared May 1917.
— Larvae and cast pupal skins from type
•

13,

^

Tipula dejecta Walk.
1856
1901

Tipula dejecta Walk. Ins. Saunders, vol. 1, Dipt., p. 442.
Tipula fumosa Doane. Journ. N. Y. Ent. Soc, vol. 9, p. 99.

Tipula dejecta is a characteristic vernal species flying in April and May.
The flies are notable inhal)itants of swamps, especially alder swamps.
On April 20, 1917, the writer found larvae of this fly in Larch Meadows,
near Ithaca, New York, in association with larvae of Rhamphidia mainensis,

Pseudolimnophila luteipennis, and other swamp inhabitants.

The

The Crane-Flies of New York— Part II

1007

more fully under the account of Rhamphidia
The larvae are reddish brown in color, and rather sluggish.

conditions are discussed(page 831).

One of the larvae found on April 20 pupated on the 22d and emerged as
an adult female on the 30th, a pupal duration of eight days. An additional
pupa was taken, associated with larva of Pseudolimnophila luteipennis,
P. inornata, Tricyphona inconstans, and other species.
Larva.

— Length, 20 mm.
Diameter, 1.8 mm.

Coloration brown; dorsum marked with Hght and dark brown; a narrow, indistinct, darkbrown, median Hne, with a broader zigzag brown Hne on either side; ventral surface a httle
paler.

Body covered with a short, dark pubescence at sides of segments, at margins longer andmore conspicuous.

Chaetotaxy as follows:

tergiteswithstrong setae in transversesix

alinement, the outermost in pairs; two strong setae on each pleural annulus; posterior ringof sternites with eight strong setae, arranged in four pairs.

52G) pale, surroundedbrown; dorsal and

by

six

Spiracular disk (Plate XCV,

approximately equal lobes which are heavily marked withlateral pairs pointed, ventral pair blunt; ventral lobes with apical half

shiny black, on ventral inner margin continued dorsad, almost contiguous on midline; innerface of dorsal

and lateral lobes suffused with dark brown, proximal margin of dorsal lobe

produced inward so that the marks are almost contiguous on median line; beneath eachspiracle a transversely rectangular,

dark brown mark.

Anal gills with four anterior lobes

which are long and slender, and a pair of rudimentary blunt posterior gills (Plate XCII, 509).

Head capsule and mouth parts as in genus. Mentum (Plate XCV, 524) seven-toothed,
Hypopharynx (Plate XCV, 525) bluntly five-toothed.

apical point the longest.

Pupa.

— Length, 15.3 mm.
Width, d.-s., 2.3-2.4 mm.
Depth, d.-v., 2.1-2.2 mm.

Coloration dark brown; abdominal incisures paler.

Form relatively stout.

General features as in genus.

blunt lobes with microscopic setae.
recurved.

Cephalic crest consisting of two

Maxillary palpi strongly curved at tip, but not entirely

Pronotal breathing horns short, tips a little enlarged.

Abdominal tergites with spines weak, on median area of each row weak or lacking; onintermediate segments about fifteen spines; pleurites with only a single weak spine on basal
ring; on posterior ring a rudimentary anterior spine and a somewhat larger posterior spine;
sternites similarly armed to tergites, but spines fewer in number and larger, on segment
5 about twelve in number; on base of posterior ring a large spine on either side median line
and a small setiferous tubercle laterad of each. Female cauda with tergal valves of ovipositor long and straight, sternal valves a little shorter; cauda with the usual six dorsal lobes,
these terminating in slender spines; at end of eighth sternite six large spines; dorsal spines
reduced to a single small pair, one near each lateral margin.

———

Nepionotype.
Larch Meadows, Ithaca, New York, April 20, 1917.
Neanotype.
With type.
Paratypes.
Two pupae with type pupa.

Charles Paul Alexander

1008
Tipula usitata Doane
1901

Tipula usitata Doane.

Journ. N. Y. Ent. Soc, vol. 9, p. 124.

A large number of fully grown larvae of an unknown species of Tipula
were found beneath the bark of a fallen tree at Stanford University,
They were sent
California, on March 22, 1915, by Harold Morrison.
to the writer at Ithaca,

New York, and emerged as adults on April 15.

The immature stages are very distinct, closest perhaps towhich also lives beneath th'e bark of decaying

trees.

T. trivittata,

Nothing is knownof the habits of the adult flies.
Larva.

— Length, 25-27 mm.
Diameter, 3-3.2 mm.

Coloration pale greenish yellow, darker above.

Form terete.

Body with a very

sparse pubescence.

Chaetotaxy as follows:

tergites(Plate XCV, 527) with a posterior row of eight setae, the middle pair of each side very closely

approximated; a seta near lateral margin at base of posterior ring, on a level with pleuralseta; pleurites, one seta on each ring; sternites with eight setae in closely approximated pairs

on posterior ring.

Spiracular disk (Plate

XCV, 528) surrounded by six lobes; dorsal andlateral pairs slender, tips of former acute; ventral lobes blunt; ventral lobes with tips black-ened, continued down proximal margin of lobes as a paler brown line; lateral lobes with innerface narrowly blackened, this mark not reaching spiracles; dorsal lobes with entire inner facebulging, intensely black, the marks contiguous at their basal inner angle; an indistinct brownspot underneath each spiracle, in some specimens this mark continuous with that of ventral
lobes; lobes not fringed with hairs.

Spiracles large, separated

or a little greater than, diameter of one.

by a distance about equal to,

Anal gills short and blunt, strongly protuberant,

surrounding anus as four fleshy lobes (Plate XCII, 510).

Head capsule as in genus.

Mentum broad, with seven to nine teeth, in the latter casethe outermost pair very small.

Hypopharynx with three or five very blunt teeth.

Antennamuch shorter and stouter than in most species of Tipula, the length only a little greater than
twice the diameter, at apex with a blunt conical papilla and a few small, cylindrical sense
pegs.

Mandible powerful, with two or three flattened teeth on ventral cutting edge.

Pupa.

— Length, 15-16.8 mm.
Width, d.-a., 2 mm.
Depth, d.-v., 2.3-2.4 mm.

Coloration pale brown; posterior margin of abdominal rings pale; lateral marginabdomen conspicuously pale yellowish white.
Form slender. Pronotal breathing horns narrow, a little expanded at tips.

ofLeg sheaths

ending on a level.

Abdominal spines very strong, but few in number; tergites with four to six spines; pleurites
with a single weak spine on each ring; sternal spines very strong, five or six in number, thoseon segment 7 subequal in size to those on segment 8; no sternal spines on base of posterior
ring.

Male cauda with posterior dorsal lobe very strong, pale, tips acute; lateral lobes greatly

The Crane-Flies of New York— Part II

1009

size, the median pair represented only by two blunt brown tubercles; eighth
segment with the usual six strong ventral and lateral lobes. Female ovipositor with valves
small; sternal valves short and broad, much shorter than tergal valves.

reduced in———

Stanford University, California, April 15, 1915.
Nepionotype.
Neanotype.
With type, bred April 15, 1915.
Paratypes.
Numerous larvae and pupae with type.

Tipula trivillata Say
1823

Tipula trivittala Say.

Journ. Acad. Nat. Sci. Phila., vol. 3, p. 26.

The larvae antl the pupae of Tipula trivittata hvc; beneath the bark ofmuch-decayed prostrate trunks or under the layers of moss that often
cover fallen trees. Abundant larvae were found on March 22, 1913, andwere placed in rearing, adults emerging on April 26. On April 16, 1914,
larvae of two distinct sizes
some very small and some nearly fully grownoccurred in abundance beneath moss (Entodon sedudrix [Hedw.] C.
MuclL, Brachythecium acuminatum [Hedw.] Kindb., Hypnum Haldanianum Grev., and Mnium sylvatiaim Lindb.) on prostrate decaying elms,
sycamores, and other trees, at Renwick Park, Ithaca, New York.
The adult flies are among the commonest of the eastern species ofTipula, and fly during a large part of the season.

——

Larva.

— Length, 24-25 mm.
Diameter, 2.7-3 mm.

Coloration pale brownish yellow, a little paler beneath.

Form moderately elongated, terete. Pubescence very short or practically lacking. Setae
on anterior segments strong, on posterior segments shorter. Chaetotaxy as follows: tergites with a posterior

row of six setae, the two middle punctures each with a single seta,

the two lateral punctures each with two setae; pleurites with a single seta on each ring;sternites with

two rows of setae, the anterior row consisting of two closely approximatedgroups of two setae each, the posterior row consisting of a single large seta, laterad of whichisa minute bristle.

Spiracular disk (Plate XCV, 529) surrounded by six lobes; lateral pairlong and slender; dorsal pair a little shorter, slender; ventral lobes blunt; ventral lobes witha jet-black mark on inner face; lateral lobes with dark markings represented only by a verysmall linear dash; dorsal lobes with a small black area.

tance a little less than diameter of one.

Spiracles large, separated by a dis-

Anal gills indistinctly lobed, four in number, two

on either side, very blunt and protuberant.

Head capsule as in genus, the mouth parts almost as in T. usitala. Mentum broad, with
Hypopharynx with five teeth, the three middle ones the longest, subequal

seven teeth.
in size.

Pupa.

Antenna shorter and stouter than is usual in the genus.

— Length, 19-20 mm.
Width, d.-s., 2.5-2.() mm.
Depth, d.-v., 2.S-3 mm.

11Charles Paul Alexander

1010
Coloration dark brown;sternitesandtergites;

abdomen yellow, with a broad sublateral brown stripe on bothabdominal segments beyond posterior row of spines brighter, moreyellowish.

Labrum broad.

Cephalic crest very small.

Characters of head as in genus.

Pronotal

Mesonotal prescutum with fine transverse wrinkles;
two blunt lateral tubercles, and behind these, on either side of median line, two smaller
breathing horns short, slightly curved.

flattened ledges which are often bifid at their tips.

(Similar ledges, but much less prominent,

occur in T. runlata.)

Abdominal tergites with subapical armature weak, spines varying in number from six toeight, those of posterior segments larger; pleural spines long and slender,

sternal spines powerful, four to six in number,

one on each ring;no spines on base of posterior ring.

FemaleCauda with sternal valves long and slender, but little shorter than tergal valves; dorsal lobesof Cauda four in number, posterior pair very powerful, lateral pair small, anterior medianpair lacking.

———Nepionotype.
Ithaca, New York, March 22, 1913.
Neanotype.
With type.
Paratypes.
Numerous larvae and pupae with types, April 26, 1917; March 22, 1913; etc.

Tipula ignobilis Locw
1863[

Tipula ignobilis Loew.

Berl. Ent. Ztschr., vol. 7, p. 2S0.

The adult flies of Tipula ignobilis are not common in collections, clue
The larvae, however, are common inin part to their retiring habits.

their preferred habitat, saturated moss cushions.

At Orono, Maine, numerous larvae were taken
17, 1913, associatedinwet moss on Junewith other larvae, such as those of Rhaphidolabina,

Tricyphona, Pedicia, and Tipula nobilis.

At Ithaca, New York, on April

23, 1917, four very small larvae were found in wet cushions of moss (Ambl}^-They grew very

stegium).

rapidly,emerging as adults on

May 21.

On May 22 this moss was carefully examined and about thirty fully growntoratherinunaturelarvaeofthisspeciesweretaken.

They wereAt
Needham's Glen, the species occurred in the same moss that earlier 'in
On June 14
the season harbored Tipula collaris and T. oropezoides.
two teneral adults of T. ignobilis were captured, in company with
Dicranornyia stulta, Geranomyia canadensis, Dactylolabis montana, andThe species is very common at the Indian Ladder, Helderother species.

associated with equally numerous larvae of Dicranomyia stulta 0. S.

berg Mountains, New York.
Larva.

— Length, 16.5- IS mm.
Diameter, 2.5-2.6 mm.

The Crane-Fltes of New York— Part1011

II

Coloration grayish brown, paler gray beneath; thoracic segments conspicuously reddish

brown.

Dorsum covered with a short, dark pubescence, which gives upper surface
Chaetotaxy as follows: tergites (Plate XCVI, 530) with a posterior transverse row of two setae near base of posterior ring; sternites (Plate XCVI, 531) with about
Body terete.

its

dark color.

eight

anterior

setae,

median row consistingof

two large

setae,

each with a tiny setaj)roxiinad of it, the posterior pair of large setae more widelyseparated.lateradof each of thesetwo closely approximated smaller setae. Spiracular disk (Plate XCVI, 532) surroundedby six moderately long lobes fringed with rather short, pale hairs; tips of ventral lobes witha pale circular area provided with a sensory seta; inner face of spiracular disk with themarkings pale, ventral lobes with an indistinct capillary brown line; two brown spots beneathAnal gills

each spiracle.

Spiracles large.

short, slender,

two-branched lobes (Plate XCII, 511).

eight, consistingofa transverserow

of four

Mentum of head capsule with median point prominent, the three lateral teeth of eitherside small,moderately acute.

Mandible with about four teeth,in addition to the large

basal prosthecal tooth.

Pupa.

— Length: male, 12 mm.; female, 14-15 mm.
Width, d.-s. male, 1.7-1.8 mm.; female, 1.8-2 mm.
Depth, d.-v.: male, 1.9-2 mm.; female, 2.1-2.2 mm.
:

Head and mouth parts as in genus.

Pronotal breathing horns rather long and slender,

Leg sheaths rather short, just exceeding third abdominal segment;
hind legs a little the longest, the other tarsi ending about on a level.
Abdominal tergites with armature weak, spines small; pleurites and sternites with spines
tips

a little expanded.

notably larger and more powerful; sternites with subterminal row consisting of twelve tofifteen spines, those of the sixth and seventh segments larger; no spines at base of posterior

weak, one on basal ring, two on posterior ring. Male
Cauda with ventral lobes widely separated, terminating in slender, curved spines directedcaudad; dorsal lobes slender, approximated on dorso-median line. Female cauda (Platering; pleural spines setiferous, very

XCVI, 533) with tergal valves a little longer than sternal valves, the latter at their tips
terminating in slender points directed ventrad a»nd laterad; dorsum of cauda with the usual
six lobes, posterior pair the largest,

terminating in two distinct points; eighth segment on

pleural and sternal region with six very powerful, curved spines.

———

Nepionotype.
Ithaca, New York, June 3, 1917.
With larva, June 3, 1917.
Neanotype.
Paraiypes.
Numerous larvae and pupae, June 1 to 12, 1917.

,

Tipida abdominalis (Say)
1823

Ctenophora abdominalis Say.

Tipula abdominalis

isJourn. Acad. Nat. Sci. Phila., vol. 3, p. 18.

the largest eastern species of the genus, altho

some specimens of T. caloptera are nearly as large. The adult flies areon the wing in late August and September, and even, in fewer nmnbers,in June and July.

\

Charles Paul Alexander

1012

The largo, fleshy larvae occur beneath or among drift at or near themargins of streams. They are usually abundant under saturated decaying leaves or under tussocks of grass at the edge of the water. They
are entirely herbivorous, feeding on diatoms, decaying plant tissues,and other vegetable matter. The larvae are readily distinguished fromallother species of the genus by the small spiracles and the bifid or split

lobes surrounding the spiracular disk.

or earth at

Pupation takes place in the mud
Tipula sp. 2 (1915-17 b: 200-

Malloch's

the water's edge.

201) seems to refer to T. abdominalis, but the mouth parts of his speci-

mens do not seem to be normal.
Larva.

— Length, 55-65 mm.
Diameter, 7.5-10 mm.

Coloration pale grayish brown; anterior end darker.

Form stout, terete, thin-skinned.

Posterior ring of abdominal segment, both above andbelow, with a naked transverse welt, which bears the setae; this welt very protuberant atends, almost like a pair of prolegs.

its lateral

Skin naked, except for a microscopic dotting

and the rather short, delicate setae. Chaetotaxy as follows: tergites, on welt at about midlength, a transverse row of eight setae, the middle pair of each half closely approximated;
pleurites with a single seta opposite basal ring, and two opposite posterior ring, arranged
one behind the other; sternites with four pairs of setae on welts, anterior middle pair closer
together than posterior lateral pair.

Spiracular disk (Plate XCVII, 537) moderately large,

flattened, truncated, surrounded by six deeply bifid

and irregular lobes; ventral lobes elon-

branch bearing two setae; posterior branch longer, indistinctly
bifid at tip and with a few setae and several hairs; lateral lobes deeply bifid, the ventral

gate, with a blunt basal

dorsal lobes small, simple; inner faces of dorsal and lateralbrown stripe, ventral lobes with two narrow, usually indistinct
Spiracles very small, circular,lines, these markings broadest and darkest on dorsal lobes.
separated by a distance about equal to fouf times diameter of one. Anal gills six, long andone armed withlateralsetae;

lobes with a narrow dark

slender, the middle one on either side a little shorter

than the others.

Pleural region of

penultimate segment of body with a blunt setiferous tubercle.

Head capsule and mouth parts about as in genus.

Mentum (Plate XCVII, 536) broadlytransverse, anterior margin with seven to nine teeth, in the latter case the outermost teeth

very indistinctly separated from the sublateral teeth.

Hypopharynx broad, anterior margin

with teeth very indistinct.

Pupa.

— Lengthof cast pupal skin,about 35 mm.

Pronotal breathing horns short, straight, cylindrical; apex short, and but little if at all

expanded.

Details ofmouth parts as in other species of genus.

before apex of second abdominal segment.
of third

Wing sheaths ending

Leg sheaths extending to about opposite end

abdominal segment; fore tarsi the shortest, hind tarsi the longest, ends of tarsal

sheath thus forming a broad, inverted U-shaped notch.

Abdominal armature generally weak.

Tergites (Plate XCVII, 538) with subterminal row

broken, consisting of an anterior median pair of large spines, laterad of which are two orThe Crane-Flies of New York— Part1013

IIthree smaller spines; close to ends of row a bifid setiferous spine; plcurites with spines long,
slender, directed strongly caudad, tips narrowly bifid, bearing a seta in notch; basal spine

the largest; posterior spines two in number, anterior dorsal one the smaller; sternites (PlateXCVII, 539) with posterior row unbroken, of comparatively few spines,

there being about

eleven excluding the two larger spines at ends of row; the innermost of the large lateral spines

conspicuously bifid, bearing a stout seta in its notch; base of posterior ring on either side of

median hne with a powerful conical spine which is acutely tipped.

Female cauda as in genus,

sternal valves conspicuously shorter than the long tergal valves; the six dorsal lobes powerful,morechitinized,

or less bifid at tips; dorsal lateral lobes at endofeighth segment

spht before tips.

—

Cascadilla Creek, Ithaca, New York, May 31, 1913.
Nepionotype.
Neanotype.Cast pupal skin, reared at Ithaca, September, 1911.
Abundant larvae from type locality.
Paratypes.

——

Tipula taughannock Alex.
1915

Tipula taughannock Alex.

Tipidataughannock

isofProc. Acad. Nat. Sci. Phila., p. 476-479.

exceptional

interestinthestriking

color

dimorphism that it shows, the females being black and yel'ow, the mabs
light yellowish.
is

The following account of the habits of the adult flies

taken from the writer's field notes:

June 12, 1915. Deciduous forest association of the southern Helderberg Mountains, near
This association is an open deciduous
the village of New Salem, Albany County, New York.
forest, with an undergrowth of Cystopteris, Geranium, Caulophyllum, and Impatiens.
It is avery open woods, having an eastern exposure and with the talus slopes so old that an extensive
vegetation has sprung up. The great boulders scattered about thru the woods have comefrom the high Silurian and Devonian cliffs above. The woods are of such a nature that muchsunlight penetrates to the ground beneath.
The forest cover shows a striking lack of coniferous species, but the following deciduous species are common: butternut, hop hornbeam,
hard maple, basswood, white ash. The shrubbery consists of mountain maple, bladdernut,
and a few dogwoods. The dominant herbage consists of jack-in-the-pulpit, wild ginger,
bloodroot, bishop's-cap, false bishop'a-cap, blue cohosh, white baneberry, herb robert, touchme-not, waterleaf, bedstraw, and other characteristic flowering plants in fewer numbers,
as well as several ferns, such as the bulbous bladder fern, maidenhair, and, on the rocks,
The crane-fly under consideration is very common in these woods. The
the walking fern.
proportion of males to females is about one hundred to one, but this is due, in large part
The males are untiring, almost always
at least, to the very secretive habits of the latter.
moving along, silently and relentlessly, in quesj: of their mates. They pass in and cut among
the dense herbage, usually close to the ground, occasionally fluttering up a tree trunk or overa mossy boulder which is covered with various bryophytes and walking ferns. They are
so intent upon their quest that they are readily scooped up by hand.
If tlus is attemptedand fails, however, they become instantly alarmed and fly away with great speed, their flightat this time having a strong undulating motion.
In a position of rest, the male almost
always hangs on the under surface of a leaf, with the body directed straight toward theground. Several specimens of this species, as well as of Tipula trivitlata Say and T. senega
Alex., were found dead in spider's webs.
These small webs, made by species of
Epeiridae and Linyphiidae, are very common on and between the leaves of herbaceous plants
and are presumably intended for smaller game. The large Tipulas are probably taken in

by accident.

Charles Paul Alexander

1014

The females are more active when in flight than are their mates and are capable of movingvery rapidly. Their flight is a rapid, flutlering progression along the ground. Copulation
Sometakes place on either the upper or the lower surface of leaves, usually near the ground.
times copulation is end to end, with the heads directed away from each other; at other times
it is face to face, the bodies being arcuated into a convex loop.
The larva almost certainly lives among or beneath the decaying leaves and d^brie which
cover the talus slope.

Tipula macrolabis Loew
18&i

Tipula macrolabis Loew.

Tipula macrolabis

isBerl. Ent. Ztschr., vol. 8, p. 58.

distinctly northern in its

distribution.

It

is

a

woods in June. The following
notes were made near the village of Indian Castle, Herkimer County,
New York, on June 13, 1915:
characteristic fly of northern deciduous

A email woodland stream with a forest cover of trees such as hemlock, beech, slipperyand basswood, and a ground cover of false Solomon's seal, wood nettle, wild ginger,
herb robert, touch-me-not, waterleaf, sarsaparilla, and the two abundant ferns maidenhair
and the bulbous bladder fern. The males of T. macrolabis were in search of the females,
and fluttered up the tree trunks often to a height of ten or fifteen feet, flying close to theground, around brush heaps, hovering about the leafy ends of branches, and performing
similar actions in their untiring quest for their mates.
They occurred in company withmales of T. fuliginosa and T. valida, which were similarly engaged in searching for the females.
elm,

Tipuline No. 1 (possibly Tipula iroquois Alex.)
Berl. Ent. Ztschr., vol. 7, p. 288-289, not T. cincta Gmel,

1863

Tipula cincta Loew.

1915

Syst. Nat., ed. 13, vol. 1, p. 2820 (1792).
Tipula iroquois Alex. Insec. Inscit. Menst., vol. 3, p. 128.

The larva discussed below has never been
here principally because ofitsreared andinteresting habitat.

It

is
is

mentionedreferredto

Tipula iroquois with considerable doubt.

The larva lives among dense mats of an aquatic moss, a Hypnum(Rhynchostegium) of the dilatatum group, in the most rapid-flowing
streams.
At Coy Glen, Ithaca, New York, these larvae are especiallyfrequent, often living at the brink of falls or rapids in thewaters.

most rushing

A study of the structure of the larva reveals numerous small

but prominent tubercles, which doubtless assist the insect in clinging to
the moss stems. The gills are large, but no better developed than inmany aquatic species of the genus that live in much less lotic conditions.
The green color and the transverse rows of tuljcrcles on the body give thelarva a strong resemblance to its mossy habitat. The larvae are verysluggish and crawl but slowly, often appearing quite dead for long periodsof time.

The Crane-Flies of New York— Part II

1015

Associated with these larvae in Coy Glen, the following characteristic
forms of insect life occur
Plecoptcra: a small species of Perlidae.

Ephcmerida: Ephemeridae, such, as Buetis, Iron fragilis, Ephemerella,
and others.
Trichoptera: Ryacophilidae, Hydroptilidae, and other forms.
Neuroptcra: Chauliodes larvae.
Diptera: Chironomidae, a few; Psychodidae, Psychoda albitarsis
Banks; Stratiomyiidae; Anthomyiidae, Limnophora torreyae Joh.;and other groups.
Coleoptera: Parnidae, larvae of Psephenus lecontei (Lee), and adultbeetles of a species of Elmis in large numbers.
Up to the present time it has been found impossible to rear this larva
to the adult condition, chiefly because of the constant need of well-aerated
water and the difficulty of supplying it. By placing the larvae in the folds
of moistened cheesecloth, it was possible on one occasion to carry the
species to the pupal state, but no further.
It will be of interest to
ascertain the identity of this conspicuous larva.
Larva.

— Length, 24-25 mm.
Diameter, 2.6-3 mm.

Coloration above, dark green with a brown pattern; beneath, light green with indistinct
transverse brown lines; on dorsum a pale longitudinal mark on sides of posterior ring, cross-ing sutures between segments onto extreme base of anterior ring of following segment, thedark area of each segment thus appearing cruciform, this cross-shaped mark spotted andmarbled with darker in transverse rows; base of gills and center of spiracular disk hght green
in hving, healthy larvae.

Form moderately terete, each segment with transverse rows of small, prominent tubercles,
some of which are provided with setae.

On dorsum of posterior ring a subterminal row ofsLx tubercles, the middle one on each side with two setae, the remaining four tubercles unise-

tose; two rows of smaller naked tubercles at base and middle of posterior annulus; basal
annulus with four transverse rows of small naked tubercles; pleurites with three tubercles,a small seta on basal ring ventrad of basal tubercle, and two setae on posterior ring ventrad

and cephalad of posterior tubercle; sternites on posterior ring with six
rior row, the two middle ones very tiny.

setae, four on ante-

Spiracular disk almost as in T. coUaris, T. ignohUis,

and similar species, surrounded by six lobes fringed with moderately long hairs; inner faceof lobes somewhSt pale; ventral lobes with a narrow, capillary, dark brown line, extending

from tip toward base; lobes narrowly and more or less indistinctly margined with brown;
two brown spots at base of each ventral lobe, underneath each spiracle; dorsal and laterallobes jutting backward at tips into fleshy conical points.
Spiracles circular, moderately
large, separated by a distance about equal to one and one-half diameter of one.
Anal gills
with eight branches, rather short and stout, with two lateral divergent branches on eitherside and an inner pair with one ventral and one posterior branch (Plate XCII, 512).

Charles Paul Alexander

1016

Head capsule and mouth parts of almost normal tipuline appearance.

Mentum withouter plate forming the unusually long apical point, inner plate adding three teeth on either

margins bulging.

side,

Antenna with apical papillae somewhat flattened, surrounded bythree sense pegs.

(Described from abundant specimens,

Coy Glen, Ithaca, New York, April 23,

1914.)

Tipuline No. 2

The larva discussed below is known only from a single, apparently
grown, specimen. The writer has no clue as to which species it

fully

represents, altho from the larval structure it is obviously allied to Tijmla
seleneLarva.

Meig. of Europe.

— Length, 23 mm.
Diameter, 3 mm.

Coloration, a rather uniform pale yellowish or reddish brown.

Body covered with a rather sparse, long pubescence, setae unusually long and powerful.
Chaetotaxy as follows: tergites (Plate XCVl, 534) with two strong lateral setae near posterior margin and an additional one at extreme lateral margin of ring; pleural setae, one on
basal ring, two, one behind the other, on posterior ring; sternites with a transverse pair of
powerful setae near extreme lateral margin, and two anterior pairs of much smaller setae.
Spiracular disk

(PlateXCVI, 535) with four elongated, cylindrical, chitinized horns which

are narrowed to the acute, blackened, slightly curved tips; longer dorsal pair rather closely

approximated, lying

almostparallel,with tips alittlecurved dorsad; shorter and more

slender lateral horns directed ventrad at tips, at base on inner face with a powerful seta,

the large black spiracles lying above base oflateral horns; ventral lobes,

ifpresent, very

Anal gills not protruded in the only specimen available.
Head capsule and mouth parts rather normal but showing the following points of difference

blunt and indistinct.

from the usual Tipulatype: Mentum almost completely split, apical point long and narrow;the three lateral teeth on either side blunt, lateral pair tending to be reduced.
Hypopharynx with five blunt teeth. Antenna short, stout, length only about twice
diameter, at tip with a subglobular, feebly chitinized papilla.

(Described from a single larva found beneath a stone in a field near Taughannock Falls,

Tompkins County, New York,May 1, 1912.)

Genus Nephrotoma Meigen (Gr. kidney + I cut)
1800
1803
1834

Nouv. Class. Mouch., p. 14 {nomen nudum).
Nephrotoma Meig. Illiger's Mag., p. 2(52.

Pales Meig.

Pachyrrhina Macq.

Hist. Nat. Ins., Dipt., vol.

The large genus Nephrotoma,

1, p.

88.

some one hundred and fifty
in all respects.
The wiitercannot attempt to separate the immature stages of the genus from thosedescribed species,of Tipula,isincludingvery close to Tipulaon the scanty material that has been available for study.

The Crane-Flies of New York— Part II

described the hfe
In Europe Bcling and others haveei<.htofthe'commonerbeneath a covering

species.

The

1017

histories of about

following species live

m earth,cornicina (Lmn.), hneataof leaf mold: analis (Schum.),The
pratensts (Lmn.).
maculata (Meig.), lunulicornis (Schum.),
the
of
some
wood,
in decaying
following have been described as living(Lmn.),
crocata(Linn.),
cornicina
records apparently being in error:
Some of the species are mjurious
qvadrifaria (Meig.).
(Scop.)

lineata (Scop.),of coniferous plants.
to young seedlings, especially thosespecies, N.ferruginea (Fabr.), has
In North America the commonestHart
relations to agriculture.
been discussed several times in its economic
immature
theof
excellent description(1898 [1895]: 218-219) gives anandb:206) gives supplementary notes(1915-17
Mallochwhile
stagesearth.
or
sand
in
live
The immature stages
figures of the same species.
from a larva found in moss on Plummers
bred
was(Loew)
virescens
N. eucera (Loew)
R. C. Shannon.
Island Maryland, on April 5, 1913, by
taken under leaf
larvaepolymera (Loew) have been reared fromN

and N.
mold in woods by Mabel M. Alexander.

The Crane-Flies of New York— Part II

1019

BIBLIOGRAPHY AND REFERENCES CITED(Anonymous)

Injurious1899.

6:56-69.

(Anonymous)

Grubs onLondon Bd. Agr.

insects

and fungi.

oats.

London Bd. Agr.

Journ.

Journ.

11:215-216.

1904 a.

(Anonymous)

Leather-jacket grubs.
1904 b.
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Instr.

Adams, Charles C.
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Dept.

Agr.

An ecological study of prairie andIllinois State Lab.

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IrelandNat. Hist.

Bui. 11 :33-280.

and Tech.

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AmsLiE, C. N.

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1906.

Notes on the swarming of a species of crane
39:26-28.

flv.

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Aldrich, J. M. a catalogue of North American Diptera (or two-winged
flies).
Smithsonian misc. coll. 46: no. 1444:1-680. (Reference on p. 771905.

105.)

Alexander, Charles P. Fulton County (New York) Tipulidae
1910.
Ent. news 2 1 247-254.
'

—

—

I.

:

Notes on Limnobia -parietijia O. S.

Ent. news 22:85-86.

Fulton County (New York) Tipulidae (Dipt.).
23:66-73. 1912 a.

—

1911.

11.

Ent. news

A bromeliad-inhabiting crane-fly (Tipulidae, Dipt.).
23:415-417. 1912 b.

Ent. news

II.

Biology of the North American crane flies (Tipulidae, Diptera).
Liogma nodicornis Osten Sacken. Journ. ent. and zool. 6:105-120.

1914.

The biology of the North American crane flies (Tipulidae,
Journ. ent. and zool. 7:1-9.
III. The genus Ula Haliday.

Diptera).

1915 a.

A second bromeliad-inhal)iting crane-fly
1915 b.
Ent. news 26:29-30.

(Tipulidae,

Diptera).

The biology of the North American crane flies (Tipulidae
IV. The tribe Hexatomini. Journ. ent. and zool. 7:141-160.

Diptera).

1915 c.

Charles Paul Alexander

1020

New or little-known crane-flies from the United States andAcad. Nat.

Part 2.
Tipulidae, Diptera.
1916 a.
Proc. 67(1915) 458-514.

Canada:Sci.

Philadelphia.

:

New or little-known

from the United States andDiptera. Part 3. Acad. Nat.

crane-flies

Canada:Tipulidae, Ptvchopteridae,
Proc' 68 486-549.
Sci. Philadelphia.
:

1916 b.

Notes on the genus Dicranoptycha Osten Sacken (Tipulidae,
1919 a.
Ent. news 30 19-22.

Diptera).

:biology of the North American crane-flies (Tipulidae,
V. The genus Dicranoptycha Osten Sacken. Journ. ent.
andzool. 11:67-74. 1919 b.

The

Diptera).

Crane-flies.

In

1913-18, 3C:3c-30c.

Report of the Canadian Arctic(Reference on p. 17c-23c.)
1919 c.

The crane-flies of New York.

onomy

the adultof

25:765-993.

flies.

Cornell

Part I. Distribution and taxUniv. Agr. Exp. Sta. Memoir

1919 d.

Alexander, C. P., and Lloyd, J. T.
(Tipulidae,

crane-flies

expedition,

Diptera).

Journ. ent. and zool. 6:12-37.

The biology of the North American
The genus Eriocera Macquart.

I.

1914.

I.
The larva of
19:541-560. 1908.

Anthon, Soester
Journ. morph.

Ctenophoraangustipennis

Loew.

Averin, V. G. [Russian title.] Operations to be undertaken in August.
Charkov Zemstvo, Ent. and Phytopath. Bur. Bulletins on the pests
Bui. 5:6-9.
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Baer, W.

Die Bedeutung dcr insektenfressenden Vogel fiir die ForstAus der Natur 9:659-671. 1913.

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Directions for collecting
Bui. 67:1-135.
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andpreserving

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Banks, Nathan, Greene, C. T., McAtee, W. L., and Shannon, R. C.
District of Columbia Diptera:
Syrphidae. Biol. Soc. Washington.
Proc. 29:173-203.

Barber, H. S.

Notes on a wood-boring S5rrphid.

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Barthou,
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—

.

1916.

Ent. Soc. Washington.

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Les ennemis de Framboisier.

Monit. hort. 37-^-248-249.

— Part II1021

Zur Kenntniss der Kopfbildung der Dipteren.

Wien. ent.

The Crane-Flies of New York
Becher, E.

Ztg. 1:49-54.

1882.

Becker, Richard.

Zur Kenntnis der Mundteile und des Kopfes derZool. Jahrb., Anat. 29:281-314.

Dipteren-Larven.

Becker, Th.

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Note preliminaire sur un diptere nouveau de Belgique.

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Beling, Theodor. Acht neue Arten deutscher zweifliigelicher Insecten.
K.-K. Zool.-Bot. Gesell. Wien. Verb. 23 Abhandl., p. 547-560. 1873 a.
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Boitrag zur Naturgeschichte verschiedener Arten aus der Familie
K.-K. Zool.-Bot. Gesell. Wien.
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Kleiner Beitrag zur Naturgeschichte der Land- und Gartenwirthschaft schadlichen Insecten. Wien. ent. Ztg. 6:61-63.
1887.

Bengtsson, Simon.

Studier ofver insektlarver.

I.

om larven af Phalacrocera replicata (Lin.), p. 1-117.
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Tillkannedomen

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Ueber sogen. Herzkorper bei Insectenlarven, zugleich ein Beitragzur Kenntniss der Blutgewebe, p. 1-23.
Kongl. Svenska Vetensk.
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Bezzi, M. Biospeologica. XX.
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Dipteres.

Ditteri cavernicoli dei Balcani.
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1-1004.

etgen.,Atti 53:

207-230.

Riduzionesci.

enat. 7:85-182.

scomparsa

delle

ali

Un nuovo genre di ditteri subatteri.
57:19-28.

1918 a.

ditteri.

Riv.

Soc. Ital. Sci. Nat.

Atti

negli

insetti

1916.

Charles Paul Alexander

1022

Stiidi sulla ditterofauna nivalc delle alpi italiane.

Soc. Ital. Sci.

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oneconomic

Pflan-(Reference

zoology,p.

1-192.

Orphnephila testacea Macq., ein Beitrag zur Kenntnis
Ann. biol. lacustre 4:53-87. 1909-10.

dcr fauna hygropetrica.

Thompson, William R.

Sur un diptere parasite de la larve d'unSoc. Biol. [Paris].
Compt. rend. 78:87-89. 1915.

mycethophilide.

Tillyard, R.
p.

The biology of dragonflies, p.

J.

314-315.)

1-396.

(Reference on

1917.

Tonnoir, a. Notes sur les Ptychopteridae (Dipt.).
Ann. 59: 115-122. 1919.

Soc. Ent. Belgique.

Etude sur Ptychoptera alhimana (Diptere Nemocere).
1914-16.
Arch. zool. exp. et gen. 55: Notes et revue, p. 81-94.

TopsENT, Emile.

The ancestry of insects with particular referenceschilopods and trilobites.
Amer. journ. sci., 4th ser.: 42: 373-387.
1916.

ToTHiLL, John D.
toTrybom, Filip.

Spar

af

Tidskr. 18(1897): 63-64.

tipulidlarver
1898.

a

sandigstrandmark.

Ent.

Sur les chenilles de Tipula nuisables aux bles. Tygodnik
1834.
rolniczo-technologiczny Kurowskiego [Warsaw], no. 41.

Tyzenhaus, C.

Vannay, C.

Contribution a I'etude des larves et des metamorphoses
1902.
des dipteres. Lyon Univ. Ann., 2d ser.: 1**: 178.

Vassiliev, E. M. [Russian title.] Report on the work of the entomological ])ranch of the myco-entomological experiment station, Kiev.
1915.
Rept. 1915:1-74.

Verrall, O. H. Another hundred new British species(Reference on p. 23.)
1912.
Ent. mo. mag. 48:20-27.
Viallanes, H.

ofDiptera*Sur I'appareil respiratoire et circulatoire de quelques
Acad. Sci. [Paris]. Compt. rend. 90:1180-1182.

larves de dipteres.
1880.

Anomalie analogue a la precedente constatee chez un
Syrphide Platychirus peltatus Meig. In Etudes dipterologiques. Wien.
1910.
ent. Ztg. 29:81-82.

Villeneuve, J.
:

Pachyrhina iridicolor Schummel jako skudce fepy cukrove.
Casopis Ceske Spolecnosti Ent. 2:70-75. 1905.

Vimmer, A^

The Crane-Flies of New York— Part II

1041

Pachyrin a Tipulin.
Srovndvacl studie o ustnim ustroji larev
190G.
3:37-49.
Ent.
Casopis Ceske Spolecnosti
Praghypopharyn^u nekterych larev z podrddu Orthorrhaphe.
20:1-11.
1911
no.
32,
Jos.,
Rozpr. Ceske Ak. Frant.

Wahl B

SchnakenlarvenalsStat. [Vienna].

Baden,

1914.

Bericht der Hauptstcllo fiir Pflanzenschutz

Wahl, C, and Muller, K.
in

1914.

p. 70.

1917.

1-130.

Amer. ent.

Crane fly larvae.

Walsh, Benj. D., and Riley, Chas. V.
:

Ark. Zool. IV

Det olandska alvarets djurvarld.

Wahlgren, Einar.

1

Pflanzensch.

K.

Pflanzenschadlinge.

Mitt. 1914:4.

18G8-69.

100.

Ward Henry Baldwin, and Whipple, George Chandler.
water biology, p. 1-1111.

Washburn, F. L.

Chionea(Reference on p. 944.)

1918.

Minnesota.

Can.

valga

in

Fresh39:103.

ent.

1907.

Webster F M.
Indiana.

Ins.

Larvaeof

3:12-14.

life

acrane-fly

destroying young

wheat

in

1891.

Reports
Report of entomological work of the season of 1891. In
and experiments in the practical work of the division,
Ent. liul.
made under the direction of the entomologist. U. S. Div.

of ol^servations1892 a.

26:72-73.

Crane flies (Tipuliae) as food of the robin.
Sta.

Bui. 43:131.

In Ohio Agr. Exp.

1892 b.

Crane flies: Leather jackets. In Underground insect destroyers
Ohio Agr. Exp. Sta. Bui 40:238-246. 1893 a.
of the wheat plant.
- Methods of oviposition in the TipuUdae. Ohio Agr. Exp. Sta.
Bui. (Tech. ser.) F: 151-154.

1893 b.

Weed Clarence M. The feeding habits of the chipping sparrow.
New Hampshire Coll. Agr. Exp. Sta. Bui. 55:99-110. 1898.
Insektlivet

Wesenberg-Lund,

C.
ence on p. 329-351.)

Westhoff, F.

iferskevande,

p.

1-524.

1915.

zur Kenntniss der Westfalischen
1879.
Wcstfiil. Prov. Ver. 8:1-16.

Beitrage

Abt. TipuUnae.

(Refer-

Arten

der

Weston, Robert Spurr, and Turner, C. E. Studies on theunpolluted stream.
of a sewage-filter effluent by a small and otherwise

digestion

Massachusetts Inst. Tech. 10:1-96.

12(Reference on p. 53.)

1917.

Charles Paul Alexander

1042

Westwood, J. 0.

An

introduction to the modern classification(Reference on p. 512-514.)
1840.

of

insects 2:1-587, +1-158.

Weyenbergii, H.

Beitrage
cephalen Dipteren-Larven,

ziir

Anatomie und Histologic der Hemi-

p. 1-G2.

1872.

The food of the common mole.

White, P. B.

no. 5:21:401-407.

Bd. Agr. London.

Journ.

1914.

Williamson, E. B.

Directions for collecting and preserving specimensdragonflics for museum purposes.
Univ. Michigan, Mus. Zool.
Misc. pub. no. 1:5-15.
1916.
of

Williston, Samuel W. Manual of the families and genera of North
American Diptera, 2d ed., p. 1-167. (Reference on p. 29-31.) 1896.

Manual of North American Diptera, 3d ed., p. 1^05.
ence on81-86.)

p.

(Refer-

1908.

WiNNERTz, J.

Beitrag zu einer Monographic der Gallmiicken.
entomologica 8:154-322.
(Reference on p. 281.)
1853.

WuLP, F. M. van der.
soorten van Diptera.

Linnaea

Jets betreffende de entwikkeling van een tweetal
Soc. Ent. Pays-Bas.
Mem. 1 15-19. 1857.
:

Dipterologische Beitrage. Part 2.
En bcskrifning af\sirvenai Cylindrotonia distinctissima.
Oken's Isis 1842:807-810. 1842.

Zeller, p. C.

Zetterstedt,

Johann Wilhelm.

descripta 10:3711-4090.

Diptera Scandinaviae disposita et(Reference on p. 4014, 4016, 4023.)
1851.

EXPLANATION OF ABBREVIATIONS USED ON PLATES
A=antenna
C=cardoPl^pleuron

PM = prementum

Lb=l;ibium, or labial lobes

S = stipes

Momentum

SM = submentum

Aid = mandible

Sp = spiracle

Mx = maxilla

St = sternite

P = palpus, or palpal sheath

T = tergite

Memoir 34, An Economic Study of Farm Layout, the fourth preceding number in this series of publications, was mailed on January 31, 1021.

Memoir 35, Some Effects of Potassium Salts on Soils, the third preceding number in this series of pubHoations, was mailed on January 29, 1921.

Memoir 36, Resistance of the Roots of Some Fruit Species to Low Temperature, the second preceding numberin this series of publications, was mailed on January 19, 1921.

A

Alemoir 37,
Modified Babcock Method for Delermininij Fat in Butter, the next preceding number in thisscries of jiublications, was mailed on December 10, 1920.

Plate XII

Memoir 38

Pec//C(7r/a/4nf'o char/P/cr(7r70/?/yc/7a:^\/^euc/o/)vm
T/pu/'/Toe/?-/77//<'i^«-7//7u/o/e/ ^rrcesfor

pupal hull.
1, Eriocern npinnsn emerging from2, Antncha
2-3, Spiracular disk and tracheation of larvae:
saxicola (apneustic); 3, DicranoUi hiniaculala (metapneustic),
after Miall4,

Phylogenetic tree

.

1013

Memoir 38

PROTOPLASA FITCHII, SUPPOSITION
5, Dorsal

aspect; G, spiracular disk;

7,

head, dorsal aspect;
anal gill

1044

8,

head, ventral aspect; 9,

Memoir 38

Plate XFV12

PTTCHOPTERA RUFOCINCTA
Larva:

10, dorsal aspect; 11,

Pupa:

13, lateral aspect; 14, ventral aspect; 15,

breathing tube extended; 12, head, ventral aspect
female cauda, ventral aspect

1045

s

£ ft

o
3

o
p<
>>

73

O

X!
CO*"

CO

^
w

^I
oo"

CO

^ -6

CO

CD

9

3

8

S,

n

a^

m

^ "S ss £="S "^ fc s

b2

-

-

I- u
S S ojoT O
Ji.

S fc ^"^

S ? «3

-

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J

1048

o
'O

Plate XVIII

Memoir 38

BITTACOMORPHA CLAVIPES
dorsal aspect
Larva: 41, labium; 42, mandible and antenna,
tubercles; 47, arrangement ol leg
Pupa: 43, ventral aspect; 44-46, types of abdominal
sheaths; 48, male cauda, dorsal aspect1049

IMemoir 3S

Plate XIX

l>**4SSSKJ?€l*B^3«f'^i^^ry^i/st^"^

BsIF

49

50
TRICHOCERA SP. (rEGELATIONIS, SUPPOSITION)
Larva:

40, dorsal aspect; 50, spiracular disk, lateral aspect; 51, spiracular disk, dorsal
aspect; 52, head, ventral aspect (after De Meijere) 53, head, dorsal aspect (after
De Meijere)
Pupa: 54, lateral aspect; 55, female, ventral aspect; 50, female cauda, lateral aspect;

1050

Memoir 38

YvV

Plate XXpti

tf

Memoir 38

Plate XXII71

ANTOCHA SAXICOLA, PUPA
69, Lateral aspect;70,

male cauda, dorsal aspect; 71,
aspect (diagrammatic)

1053

fifth

abdominal segment, dorsal

D. "

b.

„^

C +J
b.
a;

> CI
i) >
Soo"

01
U u

a>
to

,

~3
^ o
I s

CM

1054

h^IAh

Memoir 38

Plate XXIV

LIMNOniA CINCTIPES
Larva:

79,

head capsule, dorsal aspect; SO, hypopharynx; SI, mentum; S2, antenna; S3,

spiracular disk
Pupa: 84, female, lateral aspect; 85, female cauda, dorsal aspect1055

Plate XXV

Memoir 38
Y.j,/k/iiii'»ww

LIMNOBIA CINCTIPES
Larva:

8G, labrum-epipharynx; 87,

mandible and maxilla; 88, mandible; 93, spiraculardorsal aspect
Pupa: 89, female, ventral aspect; 90, pronotal breathing horn, lateral aspect; 91,
Cauda, lateral aspect; 92, male cauda, dorsal aspect

disk,

105G

male

Plate XXVI

Memoir 38

101

102
LIMNOBIA TRIOCELLATA AND

S"£ ;X:!t™:^5!tb?ul:;;pthary.x;

FALLAX90, n,e„.a,„; 97, antenna; 98,n^dible;horn; 102, female
101, pronotal breathing
aspect103, female Cauda, lateral^C^'to /«««.. pupa:
Cauda, doreal aspect;

L.

100,

mouth parts;

1067

1058

be <^

P-

t)

CD

rM

I

^

^

«3

Memoir 38

Plate XXIXDICRANOMYIA BADIA, PUPA
118, Lateral aspect;

119, mouth parts; 120, female cauda, dorsal aspect

1060

o,
-

o

83

Ol

s
s

rvj

1061

Plate XXXI

Memoir 38

RHIPIDIA RRYANTI, RHAMPHIDIA MAINENSIS,

AND RHAMPHIDIA FLAVIPES

Rhipidia bryanti, pupa:
128, mouth parts; 129, pronotal breathinghorn; 130, male cauda, lateral aspect; 131, male cauda, dorsal aspert
Rhamphidia mninensis, larva: 132, lateral aspect; 133, mentum;
135, mandible; 136, spiracular disk

Rhamphidia flampes, larva:l.'U,

antenna

10()2

^

'-'-5

o -a

00

1063

Plate XXXIII

Memoir 38

143

144

RHAMPHIDIA FLAVIPES, PUPA
abdominal segment, dorsal aspect (diagrammatic); 144,
abdominal segment, ventral aspect (diagrammatic)

142, Lateral aspect; 143, fifth
fifth

1004

Plate XXXIV

Memoir 38

145

ULA ELEGANS, LARVA

spiriicular disk

1035

1066

Plate XXXVIMemoir 38

EPIPHRAGMA SOLATRIX, LARVA
158,

159, mentuin; 100, prcmentuin;
102, maxilla; 103, spiracular disk

Head capsule, ventral aspect;

1067

IGl,

hypopharynx;

Memoir 38

Plate XXXVII

166

51-

p\

\
5p

^^

L
^V7/7^^^N^^^^Ni^,

167
EPIPHBAGMA SOLATRIX 'AND E. FASCIPENNISEj)i]ihrnqma solalrix, pupa:

IGi, lateral aspect; IGG, head of male, ventral aspect; 167,
segment (diagrammatic)
Epiphragma fascipennis, pupa:
165, cephalic crest, lateral aspect

fifth abdominal

1068

00

<

Memoir 3S

Plate XXXIXPSEUDOLIMNOPHILA LUTEIPENNIS AND
PseudoHmnophila luteipennis, larva:

172,

mentum:176, spiracuiar disk

PseudoHmnophila inornala, larva:

174,

antenna1070

P.

INORNATA

173,

hypopharynx;

175, mandible;

S S

1071

Plate XLI

Memoir 38

185

184

186

188

187

189
DACTYLOLABIS DE^fTICULATA,D.

WODZICKII, ANDD.

CUBITALIS

Dadylolabis denticulala (after Mik):
181, pupa; 1S2, larva, head capsule, ventral aspect;
183, larva, dorsal aspect
Dactylolabis wodzickii (after Nowicki):
184, pupa; LS,'), larva, head capsule; ISO, larva,
spiracular disk
Daclylolahiii cuhi(nli,<i, pupa:
187, pronotal breathing horn; 188, second abdominal segment, showing spiracle; 189, male cauda, dorsal aspect1072

Plate XLII

Memoir 38

LIMNOPHILA (dICRANOPHRAGMa) FUSCOVARIA, LARVA
190, Cephalic and caudal ends, dorsal aspect; 191, head capsule, ventral aspect; 192, labrum
epipharynx; 193, antenna; 194, mandible; 195, spiracular disk

1073

13

1074

I

1075

Plate XLVMemoir 38

LIMNOPHILA (lASIOMASTIx) MACROCERA, PUPA
209, Male, lateral aspect; 210,

mouth parts; 211, male cauda, dorsal aspect; 212, female
Cauda, lateral aspect

107fi

Plate XL VI

Memoik 38

214
213

'

LIMNOPHILA PICTIPENNIS,

L.

219

PUNCTATA, AND ULOMOBPHA PILOSELLA

Limnophila pictipennis: 213, head capsule (after Brauer)
Limnophila punctata.: 214, spiracular disk (after Gerbig)
Ulomorpha pilosella, larva: 215, head capsule, dorsal aspect; 21G, labruni; 217, antenna;
218, mandible; 219, spiracular disk, dorsal aspect
1077

is
M>
.3

.

£ S

a-3
<s S
« o

COTS

N

o3

2ta

1078

:^

i

w

-"

CO

r/T

H

to

(M

eS

=S

^

03

o3

c

.

O

a;

o

Ph

o
to

CM
en
CM

c3

o
'S.

o
CO
a

'

-^^^
TO

t-i

-^

(M
O)

/11

^ -*-'

t-t

a

5
H

S to;
6 g

ft

g (NgJ"2 •&§•

*

c3

8 e e

!^

S~

•ci •<>' *^ 'c^
S-

5-

C ^^^

vS '^ '?^ "^0,0,0, a.

1079

Memoir 38

Plate XLIXPILARIA TKNUIPES, PUPA

do;S-

STe^'mtsrS: dT,S Zti

1080'^'*'^' "^^'^ '''"" »'^-™' segment,

1081

Plate LI

Memoir 38

243

H-:

248

m

245

244

246
247

249
HEXATOMA MEGACERA
Larva:

Pupa:

243, labrum; 244, antenna; 245, mandible; 246, spiracular disk
247, lateral aspect; 248, male, ventral aspect; 249, female, ventral aspect

10S2

1

^
(N

«

-

-:

S

J

i6

g « o

1083

Memoir 38

Plate LIII

259

260
ERIOCERA CINEREA, PUPA
25G, Male, lateral aspect; 257, cephalic crest of male, ventral aspect;
258, mouth parts;
259, fifth abdominal segment, lateral aspect; 260, male cauda, dorsal aspect; 261, female
Cauda, dorsal aspect1084

Plate LIV

Memoir 38

267(f
ERIOCERA SPIN03A,
Eriocera spinos'i,larva:

E.

262,

^^

269

270

CINEREA,

E.

271

LONGICORNIS, ANDspiracular disk;

2C)7,

E.

FULTONENSIS

h?ad capsule, dorsal aspect; 271,

mandible
Eriocera cinerei, larva: 263 and 2'U, spiracular disk; 270, mandible
Eriocera longicnrnis, larva:
265, spiracular disk
Eriocera fuUonensis, larva: 236, spiracular disk; 2'>S, labrum; 269, mandible

1085

Memoir 38

Pi-ate

LV

278

277

279
273

282

274

280

276
ERIOCERA LONGICORNIS,E.

FULTONENSI3, AND

E.

SPINOSA

Eriocera longicornis, pupa: 272, male, lateral aspect; 273, male, ventral aspect; 274,
female cauda, lateral aspect; 275, head of male, ventral aspect; 277, thorax of male, dorsal
aspect; 278, female, lateral aspect
Eriocera fuUonerisis, pupa: 276, female cauda, lateral aspect; 279, female, lateral aspect
Eriocera spinnsa, pupa: 280, female, lateral aspect; 281, male] cauda, dorsal aspect; 282,
male cauda, ventral aspect

1086

Plate LVI

Mkmoir 3S

288

285

ERIOCERA SPINOSA,E.

LONGICORNIS, ANDE.

FULTONENSIS(diagrammatic);

2S3, fifth abdominal segment, lateral aspect
Eriocera spinosa, pupa:x28-1, female cauda, dorsal aspect
lateral aspect (diagrammatic),
Eriocera longicornis, pupa: 2S5, fifth abdominal segment,.

male cauda, lateral aspect„:„„ififth abdominal
irroccm fuUonends, pupa 2S7, cephalic crest of male, ventral aspect 288,

2Si,

;

:

segment, lateral aspect (diagrammatic)1087

Plate LVII

294

295

293
292
PENTHOPTERA ALBITARSIS
Larva:

289, labrum; 290, antenna; 291, mandible; 292, spiracular disk, dorsal aspect;
293, spiracular disk, lateral aspect

Pupa:

294, pronotal breathing horn; 295, male cauda, lateral aspect

1088

Memoir 38

Plate LVIII

296
PENTHOPTERA ALBITARSIS, PUPA
296, Female, lateral aspect; 297, female, ventral aspect; 298, female cauda, dorsal aspect;
299, male cauda, lateral aspect1089

Memoir 38

Plate LIX

300

304
305
ADELPHOMYIA MINUTA (SUPPOSITION), LARVA
300, Labrum; 301, antenna; 302, mentum; 303, mandible; 304, maxilla; 305, spiracular
u; k

1090

Plate LX

Memoir 38

307

309

ADELPHOMYIA MINUTA (SUPPOSITION), PUPA
dorsal aspect; 309, male
306, Female, lateral aspect; 307, mouth parts; 308, female cauda,
Cauda, dorsal aspect; 310, male cauda, lateral aspect1091

Memoir 38

Plate LXI

311

315
PEDICIA ALBIVITTA, LARVA

311, Dorsal aspect; 312, head capsule, ventral aspect; 313, antenna; 314, maxillary palpus;
315, anal gills, ventral aspect1092

Plate LXII

Memoir 38

324

322
RHAPHIDOLABINA FLAVEOLA
Larva:

31G, mentum; 317, hypopharynx; 318, antenna; 319, mandible; 320, maxilla;

321, spiracular disk
Pupa: 322, mouthparts; 323, pronotal breathing horn, dorsal aspect; 324, pronotal

breathing horn, lateral aspect; 325, male cauda, dorsal aspect1093

Memoir 38

Plate LXIIITRICYPHONA INCONSTANS

mentum;Q^n*'"''^'
'l^^.
.mU,
spiracular
disk

327,

hypopharynx; 328, antenna; 329, mandible and maxilla-

Pupa: 331, mouth parts; 332, pronotal breathing horn,
dorsal aspect; 333, pronotal
breathing horn, lateral aspect; 334, female cauda, lateral
aspect
1094

CM
en

8§

o
en

I
S

»CO
CO

— a
_
<!

03

<!

g
s

•«

o
j3

o o3

33

CO •- c

^t^ o

^ aa
g^ e3 CO

2"3co
w CO

> tc *

<«

S

S " s

-o
•J

en
1095

° ..

03

a

Memoir 38

Plate LXV

344

349

346
MOLOPHILUS HIRTIPENNIS, LARVA
344, Lateral aspect; 345, head capsule, ventral aspect; 346, mental plate; 347, hypopharynx;
348, antenna; 349, mandible; 350, maxilla; 351, spiracular disk

1096

Plate LXVI

Memoir 38

355

352

354

uJ

356

MOLOPHILUS HIRTIPENNIS, PUPA
352, Female, lateral aspect; 353, male cauda, lateral aspect; 354, male cauda, dorsal aspect;
355, female cauda, lateral aspect; 356, female cauda, dorsal aspect1097

Memoir 38

Plate LXVII

357

358

363
ERIOPTERA MEGOPHTHALMA
Larva:

357, spiracular disk; 358, anal gills, ventral aspect
Pupa: 359, female, lateral aspect; 3(50, female, ventral aspect; 361, fifth abdominal segment, lateral aspect (diagrammatic); 362, male cauda, lateral aspect; 363, male cauda, dorsal

aspect1098

Plate LXIX

Memoir 38

372

369

ERIOPTERA CHLOROPHYLLA,E.

SEPTEMTRIONIS, AND

E.

ARMATA

Erioptera chlorophylla, pupa:
3G9, female, lateral aspect; 370, female cauda, lateral aspect
Erioptera septemtrionis, pupa: 371, male cauda, lateral aspect; 372, male cauda, dorsal
aspectErioptera armala, pupa: 373, female, lateral aspect

1100

Plate LXX

Memoir 38^=^ 376

378
ORMOSIA NUBILA,O.

INNOCENS, AND O. MEIQENII

.^

disk
Ormosia nubiia, larva: 374, mandible; 375, spiracular
aspect; 377, male Cauda, dorsalOrv^oda innocens, pupa: 376, female, lateral
aspect; 378, male cauda, lateral aspect
Ormosia meigenii, larva: 379, spiracular disk

1101

Memoir 38
Plate

LXXI

381

380

OBMOSIA NUBILA, PUPA
380, Female, lateral aspect; 381, cephalic crestof female, ventral aspect; 382

Cauda, dorsal aspect

1102

'

femalePlate LXXII

Memoir 38

385

384

>)wj£w/i»ryin»in\W^5^

383

386

387

ORMOSIA NIGRIPILA, PUPA
383

segment,
Male, lateral aspect; 384, pronotal breathing horn; 385, fifth abdominal
male cauda, dorsal aspect; 387, female cauda, laterallateral aspect (diagrammatic); 386,

aspect1103

Memoir 38

Plate LXXIII

395

HELOBIA HTBRIDA AND TRIMICHA PILIPES
Helobia hybrida, larva: 388, spiracular disk
Helobia hybrida, pupa: 389, female, lateral aspect; 390, mouth parts; 391, arrangementof leg sheaths; 392, male cauda, dorsal aspect; 393, female cauda, lateral aspect; 394, male
Cauda, lateral aspect
Trimicra pilipes, larva: 395, spiracular disk (after Gerbig)

1104

1105

U

Plate LXXV

Mkmoir 38

402

403

404

405

406
GONOMTIA ALEXANDERI AND ERIOPTKRINK NO.

1402, mandible; -^lOI?, spirafular disk
Erioptcrine No. I, larva: 40+, lateral aspect; 405, spiraeuiar disk and anal
406, lobe of spiraeuiar disk, enlarged

Gonomi/in alexanderi, larva:

1106

gills;

g

1107

d

-2

a

«

^

fe

^

c« .S

03

~

Sg

03

^

..

a;

Plate LXXVIII

Memoir 38

420

421

423

422

TRENTEPOHLIA PENNIPES AND T. BROMELIADICOLA
Trentcpohlia jiennipes, pupa: 419, lateral aspect (after De Meijere)
Trcntepohlia bromeliadicola, larva (after Picado) 420, anal gills
Trentcpohlia bromeliadicola, pupa (after Picado): 421, female, ventral aspect; 422, male
CI id,"., lateral aspect; 423, female cauda, lateral aspect
:

1109

CD

Jail

Plate LXXXI

Mkmoir 38

442
ELEPHANTOMYIA WESTWOODI, PUPA
441, Female, lateral aspect; 442, female, ventral aspect1112

Plate LXXXII

Memoir 38

445

444

446

447

443
CHIONEA ARANEOIDES, LARVA (aFTER BRAUER)
mental plate; 446, spiracular disk,
443 Dorsal aspect; 444, mandible; 445, possibly'

lateral aspect; 447, spiracular disk, dorsal aspect

1113

^

Plate LXXXIII

Memoir 38

448

452

449

450

PHALACnOCERA REPLICATA AND TRIOCMA TRISULCATA
Phalacrocera replicata, larva: 448, lateral aspect
Phnlacrocera replicata, pupa: 449, male, lateral aspect;mouth parts
Triogma trisulcata: 452, larva (after Steinmann)

1114

-0)0,

male, dorsal aspect; 451,

Memoir 38

Plate LXXXIV

454

45!

456

453

CYLINDROTOMA SPLENDENS
Larva:

Pupa:

dorsal asp-ct 454, mandible
455, female, lateral aspect; 456, head of female, ventral aspect
4.>5,

;

1115

t
Ph

CD

O
Z

in

•J

^^J

o

in
1^

2

S

c|

CO

^

^

^

—

1117

Plate LXXXVII

Memoir 38

475

474

477

476

BRACHYPREMNA DISPELLENS, PUPA
474, Head, ventral aspect; 475, pronotal breathing horn; 476, arrangement of leg sheaths;
477, male cauda, ventral aspect

1118

Plate LXXXVIII

IVIemoir 3S

482
TANYPTERA FRONTALIS
Larva: ITS, inenturn; -479, antenna, 4S0, mandible; 481, fifth abdominal segment, dorsal
aspect (diagrammatic); 482, fifth abdominal segment, ventral aspect (diagrammatic); 483,
spiraciilar disk
Pupa: 4SJ-, female, lateral aspect; 485, mouth parts; 48(5, pronotal breathing horn; 487,
female cauda, dorsal aspect

1119

Plate LXXXIX

Memoir 38

LONGURIO TESTACEUS AND AESHNASOMA RIVERTONENSISLongurio testaceus, larva: 488, mentum; 489, hypopharynx; 490, spiracular disk
Lonyurio'testaceus, pupa: 491, male, lateral aspect
Aeshnasoma river tonensis, larva: 492, spiracular disk, showing branched anal gills (afterJohnson)1120

Platk XC

Memoir 38

496

493

497

495

494
HOLORUSIA RUBIGINOSA
Larva: 493, mentum; 494, hypopharynx; 495, antenna; 496, spiracular disk

Pupa: 497, male, lateral aspect1121

Memoir 38

Plate XCI

504

503

505

498

PRIONOCERA FUSCIPENNIS
Larva: 498,

men turn; 499, hypopharynx; 500, antenna; 501, mandible; 502, spiraculardisk

Pupa: 503, lateral aspect; 501,

tip ofpronotal breathing horn; 505, male cauda, dorsal

aspect1122

Plate XCII

Memoir 38

TYPES OF ANAL GILLS IN GENUS TIPULA, VENTRAL ASPECT
506, Tipula oropezoides; 507, T. nobilis; 508, T. caloptera; 500, T. dejecta; 510, T. usitata;511, T. ignobilis; 512, T. iroquois (supposition)

1123

2
In

in

Ah

'O

22
In

Memoir 38

Plate XCIV

523
522
TIPULA COLLARI3
Larva: 519, mentum; 520, hypopharynx; 521, fifth abdominal segment, dorsal aspect;
522, spiracular disk

Pupa:

52.3,

male, lateral aspect1125

Plate XCVMemoik 38

528

/ ^

f

529

f

\

524

527

526

525
TIPULA DEJECTA,T.

USITATA, AND

T.

TRIVITTATA

Tipula dejecta, larva: 524, menturn; 525, hypopharynx; 520, spiracular disk
Tipuln unUita, larva: 527, fifth abdominal segment, dorsal aspect; 528, spiracular disk
Tipula trivilluki, larva: 529, spiracular disk

1126

1127

-7

^

1128

INDEX(When more than one page reference is given, the more important references are italicized)
PAGE

Abdomrn of pupa
Abiioruinal spirai-Ies

Adflphomyaria
Adclpliomyia
aiuericana

cay u«aminutaAdipo-^e tissue

Ae ihnasoma
978,rivertonensis
Amphibia feeding on crane-flies
Anal gills of larva
Antennae of larva
Anthomyiidae feeding on crane-flies

T28,

Antochamonticola
798,

saxicola

Antocharia
Ants feeding on crane-flies
Arachnida feeding on crane-flies
Arachnida parasitic on crane-flies

Arrangement of families and lesser groups
Asilidae feeding on crane-flien
Aves feeding on crane-flies

.

.

.

Bibliography
Birds feeding on crane-flies
clavipes

Bittacomorphellajonesi

Bittacomorphinae
Bittaeomorphinae, keys to genera of
Body form of larva

773,

Brachypremna
dispellens

984,

eocenica

Breathing horns of pupa

denticulata.

780
779
779
745
984
985
984
754

837,

montana ....'.
wodzickii

837,

Dicranomyia
badia
dunietoruiii
foliocuniculator

macateei

721
783
784
779

Bittacomorphacubitalis

rara

734
693
1019

820,
798, 799,
799,

pilipennis

simulans

799,
820,
798,
799,

stultatrinotataumbrata
Dicranophragma
Dicranoptycha
winnemana

910
950

Chionea
alpinaaraneoides
valga
Circulatory system

911,

Claduradolicatula
flavoferruginea
Coleoptera feeding on crane-flies
Collecting

947,

Comparative morphology
Ctenophoraangustipennis
apicataf estiva

flaveolata
nigriroxa
pectinicornis
vittata

978,
978,
978,
978,
978,
978,

Ctenophoraria
Cylindrotoma
distinctissima

splendens
tarsalis

960,
708, 960,

951
961

950
7G0
947
948
948
729
740
766
986
9se
986
986
986
986
986
986
986
966
967
967
967

852
852
854
853
855
864
819
824
810
820
820
820
820
819
820
861

799,

Dicranoptvcharia
Dicranota
bimaculata
Dicranotae

899,
895,

Dictenidia

bimaculata

978,Digestive systemargus
caesarea

798,

Ditomyiidae
Dolichopeza
albipes

828
829
828
906
906
906
987
987
758
800
727

DiotrephaDiptera feeding on crane-flies
Dipterous parasites of crane-flies
Discobola

978,

731
815
815
815
788
981
981

982
978

americanasylvicola

Cheilotrichia imbuta

950
960

D

800
800
799
729
727
732
701
728
721

792,

Dactylolabaria
Dactylolabis

799

B
Bacteria in crane-fly larvae
Beling, Theodor, life of

PAGE
Cylindrotominae
Cylindrotominae, keys to genera of

756
757
895
895
898
898
896
763
993
993
726
748
751
732

Dolichopezaria

981

Duration of pupal existence.

720

Economic importance.
Egg.
Elephantomyaria
Elephantomyia

909,

.'

.

.

westwoodi
EUipteraomissa798, 806,

Ellipteraria

Emergence of adult

Empedomorpha
empedoides
Empididae feeding on crane-flies
Enemies of crane-flies
Epipharynx of larva
Epiphragma
fascipennis
picta

837, 844,
837,

solatrix

Epiphragmaria
Erioceracinerea1129

f ultonensis

837,
838,

736
714
952
952
953
806
807
806
710
933
933
728
721
750
843
846

8U
844
843
881
886
890

Charles Paul Alexander

1130

loiiKicornis

70/,,

spiroaa
Eriocera, keys to species of

S37,
837,

SSS
883
882
91-1

Erioptera
arniata

915,
915,
915,
910,

calopterachlorophylla
flavescensgraphica
knabi

915,
910,
910,

lutoa
niaculata

nicgophthalma
parva
septomtrionis
sordidavcspertina
Erioptera. key to pupae of
Eriopteraria
Kiioptorine No. 1

915,
910,
915,

921

Eriopterini. keys to genera ofEuliiiuiopliila

External niorphology of larva
External morphology of pupa
External parasites

9U

Imaginal disks

Flagellate parasites of crane-flies

Internal morphology
Internal parasites of crane-flies

919_

Killing and preserving specimens

Geological record

Geranomyia
canadensis
diver.sa

rostrata
virescens

Gnophomyia
rufa
tripudians

911,
911,
911,

tristissima

Gonomyella

Gonomyia

938,
938,
938,

alexanderi
kansensis
subciijerea

sulphurella

938,
911,

tcnellaGonomyia, key to pupae of

744
752
764
816
817
817
817
817
934
93i
931,

935
938
940
939
941
938
940
938
938

H
Habroinastix cinerasoens

Head of larva
Head of pupa
911,

Hexatomabicolormegacera
nigra

saxonum
Hexatomaria

838,
838,

980
748
753
928
929
929
877
878
878
878
878
876

763
758
733

K
740

L
Labium of larva
Labruni of larva
Larva
Larval habitat
Lasiomastix
Leiponeura
Libnotes
perkinsi
798,
Life activities of crane-flies
Life histories of crane-flies, representative.
.

.

Limnobaria
Limnobaria, keys to genera ofLimnobia
annulusbifasciata
cinctipes

decemmaculata

G
General features of larva
General features of pupa

978,

877
835
836
763
735
993
994
921
729

I

915
922
920
915

763
713
717
726
734
732

Fungous parasites of crane-flies

Hexatoniiiii
Hexatoniini, keys to subtribcs ofHistoblasts
Historical summary of immature stages

Holorusia
rubiginosa
Hoplolabis
Hymenoptera feeding on crane-flies

F
Fat bodies, or adipose tissue
Feeding habits of adults
Feeding habits of larvae
Fish feeding on crane-flies

Hexatomaria, keys to genera of

922
91S
915
915
922
915

915
911
956
908
909
872
744
752
730

lOriopterini

Hclobia
hybridamacroptera

PAGE

page

Eriocera (continued)

fallaxflavipes

immatura

798,
798,
798,
810,
798,
798,

indigena
inustamacrostigma
nigropunctata
nubeculosa
obscuricornis
parietina

quadrimaculata
sexpunctata
solitaria
triocellata

tripunctata

798,
798,
798,
798,
798,
798,
798, 810,
798,

xanthoptera
798,
Limnobiinae
Limno.biinae, keys to tribes and subtribes of.
Limnobiini
Limnobiini, keys to subtribes ofLimnophila
858,
adustabryobia
837,
disparferruginea
fuscovaria
hyalipennis
lineola837,
837, 860,
837,
837,

macroceranemoralisoehracea
pallida.
pictipennis

punctata
sinistra

837,
837,
837,
837,
837,
837,

750
750
715
716
863
939
814
814
7lO
704
808
808
809
809
809
810
809
813
810
810
810
798

810
810
810
810
810
809
810
810
814
810
809
793
793
795
797
868
867
859
SGO
866
861
860
860
864
860
S60
S60
869
868
859

Thk Crane-Flies of New York— Pautultimaunica
Lininophilae
Limnophilaria
Limnophilaria, keys to gencraand subgenera of
Liogina960.
Rlabrata
960,
nodirornis
961,
Liogma, keys to species of
Lipsothrix icterica
Longiirio
testaceus

PAGE

711

Paramongoma

800
858
8o6
857
069

Parasitic natural enemies of crane-flies

00.9

971

970
910
900
99

Pedicaria
Pedicaria. key to genera of
Pedicia
albivitta

895,

contermina
rivosaPediciae

895,

Pediciini
Pediciini, key to subtrlbes ofPenthoptera

721
751
710
752
921
741
741
911

'

Maxillae of larva

M"sopyphoiia

Methods of rearing
Methods of study
Molophilus
bifilatus

Pilaria,

914
944
763
788
788

PolymeraursinusN
Neanotype
NoocladuraNcphrotoma

743
947
1018
980, 101
980, 1017
980, 1017

cornicina
crocataeuceraferruginea
histrio
iridicolor

737, 981,,

lineatalunulicprnis

980,
980,
980,

maculata
polymera
pratensis
quadrifaria
virescens

1017
1017
980
980
1017
1017
1017
1017

980, 101
981, 1017

1017
743
762

Nepionotype
Nervous system

-.

837,
837,

quadrata

Oil
91

reconditatenuipes
keys to species ofPisces feeding on crane-flies

837,

836,

geniculata
georgiae
magnifica

Polymeraria
836,
Predatory natural enemies of crane-flies
Prionocera
fuscipennis
parri

978,
978,

Progononiyia
Protoplasa
fitchii

Pseudolimnophila
luteipennis

837,

albimana
contaminata

773,

lacustris
lenis

773,

paludosa
rufocincta

Ptychopteridae
Ptychopteridae, keys to subfamlUeo of
Ptychopterinae

Pupa
Odonata feeding on crane-flies
Orimarga
Orimargula
Ormosia
haemorrhoidalis
innocens

nodulosa

910,
923,
910,
923,
923,
910,

nubilavaria

910,

lineatameigenii
nigripila

Ormosia, key to pupae of

Or jpeza
obscuraOviposition

727
799
799
922
023
9$5
923
928
927
923
923
923
923
982
983
713

866
766
763
872
872
872
875
874
873
872
726
893
893
893
893
893
721

995
996
996
938
769
769
818
851

inornata

Pseudolimnophilaria
Ptychoptera

Pupal duration

O

802
961
962
961

Pilaria
discicoUis
fuscipennis

013
912

Muscular system
Mycetobia
Mycetobiinae

960,

Phylogeny

910,
910,

Mongomaanalisreplicatatipulina

Phylidorea
Phylogenetic considerations

910,

hirtipennis

obscurusochraceus

838,

Phalacroccra

Mammalia fording on crane-flies
Mand.ble of larva
Mating

944
730
899
899
899
900
900
900
899
894
894
891

albitarsis

M

1131

P

PAGE

Liiiinophila (continued)

II

8o0
848
773
774
773
773
775
773
775
772
773
773
719
720

R
References cited
Reproductive system
Respiratory organs
Resting habits

Rhabdomastix
flava

schistacea

1019
762
760
712
942
942
911,
94£

Rhamphidaria
Rhamphidia
flavipes
longirostrisniainensis

Rhaphidolabina
flaveola

799,
799,

830
830
833
830
831
901
901

Charles Paul Alexander

1132

PAGE
Rhabidolabis
cayugatenuipes

895.

Rhipidiabryantidomestica799,

fidelisniarulata

799,
799,

uiiisoriata

Uhvphidac
Rhyphiiiao

Rhyplmspunctatus

907
907
907
825
826
826
826
826
826
787
787
787
788
942
728
746
734
957
958
958
958
958
958
957
711

Scatophagidae feeding on crane-fliesS|)ira<'ular disk of larva

Sporozoan parasites of crane-flies
Styringomyia
ceylonica

didymajacobsoni

venustavittata

Styringomyiini

Swarming

979,
979,

macrolabis

marmorata

maxima979,.959

737, 979,

979
999

micansnigra
nobilisnubeculosa
ochracea
oleracea
oropezoidcj

paganaPaoandaga

999
999
978
1014
999
979,

lunataluteipennis
lutescenspabulina

S

P^^aE

Tipula (continued)

paludosa
parva
peliostigma
pruinosa
rufinasayi
scriptaseleneserta
signata

1004
979,
979,
737, 979,

1001
979.
979.
737, 979,
7.37.

979,
979,
710. 979,
979.
979. 999,
979.

simplex
subnodioornis

979.

taughannock
tephrocephalaT
Tachinid parasites of crane-flies
Tanyderidae

731
769

Tanyptera

988
988
988
988
988
988
945
946
798

atrataatrata ruficornis

978,
978,
978,

flavicornis
frontalis

fumipennis
Teucholabis

978'.

complexa
Thaumastoptera
Thorax of pupa

911,
716,

754
998

Tipulaabdominalis
angustipennis

980, 1011
1000

980

annulicornis
arctica

980, 1001

1004
738, 980, 1001

bclla
bicorniscaesia999
978,980, 1005

caloptera

cayugacoUariscunctans

738, 980,

dejectadilatata
eluta

978,
980,

flavicansflavolineata
fuliginosa
fulvipennis

gigantea
hortensis

hortulana

978,
978,
'.

978,
979,

1000
1002
1001
1006
999
1001
980
999
1000

999
979
999
999

tricolor
trivittata

truncorum
ultima980.

usitatavariicornis
variipennis
vernalis
vittatawinnertzii

980,
980,
980,
980,

muliebrisregelationis

inf uscata

980

immaculata

1014

inconstanspaludicola

longicornis

979,
979,

999
999
980

788,

auripennis

schinerivernalis

999
999
999
999
980
990
792
974
977
981
978
768
955
955
956
943
944
943
943
94^
944
789
790
790
790
789
1001

Tricyphona.

1010
979

lateralis

911,
911,

hiemalis

ignobilis
inibecillairoquois
irrorata

911.

cruciferella

Trichocerinae
Trichotipula

1000
999
1008

791
792,

Tipulodina pedata
Tipuloidea, keys to famlUej ofToxorhina
madagascariensis

humeralis
leucoxena
pennipes
Trichocera
f uscata

95.9

999
999
1000
999
1016
980
999
738
999
1013

.980. 1009
999
979.
980. 1001

unca

Trentepohlia
bromeliadicola

999
999
999
979

1000
1000
.

umbrosa

Tipularia
Tipulidae
Tipulidae, keys to subfamilie.T ofTipulinae
Tipulinae, keys to genera of
Tipulini

999
999
999

895.

895.

903
904
904
904
903
904
904

The Crane-Flies of New York
Trimicra911.

pihpes..
reciprocauia {continued)

932

elegans(^^3

macroptera

933
J73

Triogma«'="'P^

S::ia;V.V.;.V

PAGE

X7I
.•..••••.••.•.••.•.

^1.

Types of the iiuiuature stagesuUlaboLtophha.V.V.V.'.V.V.V.V. ...... -SSV,

II
743— Part

II

1133
^^^^
837,
837.

Ulariauioniorphanilosella

Ulo-norphae

84O
839
838
869
869
869

W
839
S40

Wasps feeding on ciaue-flies729r ^^

9 f i

*^B

1

>

=4

\

•.I

i