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Records and Descriptions of North American Crane-Flies (Diptera) Part V. Tipuloidea of theGrand Teton National Park and Teton National Forest, Wyoming
Author(s): Charles P. Alexander
Source: American Midland Naturalist, Vol. 33, No. 2 (Mar., 1945), pp. 391-439
Published by: The University of Notre Dame
Stable URL: http://www.jstor.org/stable/2421338
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Recordsand Descriptionsof NorthAmerican
Crane-Flies(Diptera)'
Part V. Tipuloidea of the Grand Teton National Parkand
Teton National Forest, Wyoming
CharlesP. Alexander
In continuationof the seriesof papersconsideringthe rich crane-flyfaunaof the westernNorth Americanstates and provinces,the presentreportconcerns one of the most beautifuland strikingsectionsof the entire UnitedStates, the GrandTeton National Park and vicinity. As was done in previous instalments,illustrationsof new and little-knownspeciesare provided
since it is certainthat such aid materiallyin the correctdeterminationof the2variousforms. The recentpublicationof the "Crane-fliesof Connecticut"
has made availablekeys and figuresfor the speciesnow known from northeasternNorth America. With this as a basis it is intendedin the present
paperand succeedingreportsunderthis title to figuresuch speciesthat havenot been so illustrated.
General Account
The GrandTeton National Park, embracingthe more strikingpeaks of
the Teton Range,northwestern
Wyoming,has an area of approximately150
squaremiles or 96,000 acres. The park has a length of 27 miles, with awidth varyingfrom 3 to 9 miles. Its northernextremitylies approximately
11 miles south of the southernboundaryof the YellowstoneNational Park
(see Part IV of this seriesof papers;Alexander1943). Despite the close
proximityof the two parks,a markeddifferencein the crane-flieswas foundthat cannotbe attributedto seasonalcollecting,since the majorseriesof flies
the same time
availablefor studyfrom both areaswas takenat approximately
the late springand early summerfauna. The apparentof year,representing
reason for the differenceslies in certainvery distinct ecologicalconditions
existingin the two areas (compareStations 1 and 2 of presentpaper). As
indicatedby the presenttitle, I am includingnot only the speciesof craneflies known from the National Park but also from certainselectedstations
1 The preceding part under this general title was published in the American Midland Naturalist 30:718-764; 1943.
References in the text refer to the bibliography at the conclusion of the generalaccount.
In all cases in this report where no collector is given, the specimens were securedby the author. M. M. Alexander-Mrs. Charles P. Alexander.
2 Alexander, Charles P. The Diptera or true flies of Connecticut. Tanyderidae,
Ptychopteridae, Trichoceridae, Anisopodidae, Tipulidae. Conn. State Geol. and Nat.
Hist. Surv. Bull. 64:183-486, index 501-509; figs. 18-55 (with 389 individual illustrations);: 1942 (Dublished in 1943).
391

392

THE AMERICAN MIDLAND NATURALISTin the adjoining Teton National Forest, a vast area bordering the Park on
the east and extending northward to the limits of the Yellowstone National
Park and the Shoshone National Forest. Very recently a further noteworthy
addition to the national holdings in the Grand Teton section was made by
the setting aside of the Jackson Hole National Monument, established byorder of President Franklin D. Roosevelt in the spring of 1943, including
nearly 220,000 acres of land to the south and southeast of the Park in theJackson Hole country.
As was indicated in the preceding part, the first white man to see theTetons was John Colter, who, in the winter of 1807-1808, passed through theJackson Hole at the foot of the Teton Range. In the autumn of 1811, theAstorians, under the leadership of Mr. Wilson Price Hunt, crossed the range
through the Teton Pass. Washington Irving, in "Astoria", has described
this in detail. Two short paragraphs may be quoted from this classic.
September 15, 1811. "In the course of the day they came to a height thatcommanded an almost boundless prospect. Here one of the guides paused,
and, after considering the vast landscape attentively, pointed to three mountain peaks glistening with snow, which rose, he said, above a fork of theColumbia River. * * * These remarkablepeaks are known to some travelers
as the Tetons." And still later in September. "An important point in theirarduous journey had been attained, a few miles from their camp rose the threevast snowy peaks called the Tetons."
Atwood (1940) speaks of the Grand Teton National Park as being "oneof the scenic wonderlands of the world." To anyone visiting the area in midJune, when the mountains are still chiefly snowclad, the above characterization does not seem exaggerated. In its essentials, the National Park consistsof the Teton Range, with its series of giant peaks separated by deep canyons,at the foot of each of the latter with a beautiful lake. From north to south,
these lakes draining into Cottonwood Creek and thence into the Snake River,include Leigh (altitude 6,870 feet); String or Beaver Dick (6,867 feet);
Jenny (6,779 feet) and Bradley (7,061 feet). Still farther to the south at thefoot of Death Canyon and draining directly into the Snake is one of the mostbeautiful of all the lakes, Phelps (6,615 feet). The mountains that concern
us chiefly are, from north to south, Mount Moran (12,100 feet), separated
from the St. John Range by Indian Paintbrush Canyon and Leigh Canyon;
the various peaks of the St. John Range between Leigh Creek on the northand Cascade Creek on the south; the great mass of the Teton Range proper,with the various giants, Owen, Grand Teton (13,766 feet), Teewinot, Middle
Teton, South Teton and Nez-Perce, with others. Still farther to the south thepeaks become lower but are still impressive and very beautiful. The southernend of the range is marked by the Teton Pass (8,429 feet), permitting travel
into Idaho.
From the eastern or Jackson Hole side, the Teton Range presents an unusually precipitous front, rising abruptly from the floor of the valley (about
6,500 feet) to the maximum height of more than 13,760 feet. Glaciers of theIce Age have played the chief role in sculpturing the valleys and altering thefloor of the Hole. Each of the various main canyons contained its own local

ALEXANDER:NORTH AMERICAN CRANE-FLIES,

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glacier, all emptying onto the floor of Jackson Hole. The signs of the laststages of the glacier are to be seen everywherein the moraines, outwash plains,lakes and canyons.
The various canyons permit easiest access into the heart of the range.
Those most used by our party in 1941 and 1942 were Indian Paintbrush Canyon, Cascade Canyon and Death Canyon, the latter the most southerly. Cascade Canyon, between the St. John peaks and the three Teton giants was
especially favorable and accessible. Cascade Creek, often a roaring mountaintorrent from melting snow, flows down the canyon and may be followed forseveral miles along a well-maintained park trail. Near the head of the northfork of Cascade Creek, one reaches a beautiful mountain tarn, Lake Solitude(9,024 feet), close to the extreme western boundary of the park. Shortly before it flows into Jenny Lake, Cascade Creek forms one of the most beautiful
falls in the region, Hidden Falls, where numerous crane-flies were taken, as
discussed later. The Indian Paintbrush and Death Canyon trails were similarly
followed for several miles and excellent collecting found, particularly at
middle altitudes.

Life Zones.-Of the seven life zones or faunal areas recognized by Merriam and his successors, four are to be found within the geographical limits
considered in this report.
Transition (Foothills). The valley of the upper Snake River in theJackson Hole country, between the towns of Moran and Jackson. This includes the floor of the "Hole" to an altitude of about 6,700 feet. This isnoteworthy as being one of the chief winter reserves of the American elk inthis region. Station No. 1 (Arizona Creek) is definitely in this zone;
Station No. 2 (Moran bog), while lying within the limits of the Transitionshows definite features of the next zone.
Canadian (Montane). Much of the country surrounding the foot of the
Teton Range, including the various lakes as discussed before; thence up thelower mountain slopes and canyons to the upper limits of large tree growth,
or approximately 8,000-9,000 feet, dependent on exposure. Lake Solitude(9,024 feet) and Twogwotee Pass (9,650 feet) show some features of thehighest Canadian, yet appear to fall in the succeeding zone. Virtually allcollections of crane-fliesmade by the author and colleagues in 1941 and 1942
are from the Canadian zone.
Hudsonian (Subalpine). The relatively narrow belt of scrubby tree
growth above the Canadian zone and below the limits of extreme timberline.
As mentioned, Lake Solitude and Twogwotee Pass seem to pertain to thiszone.
Arctic-Alpine (Alpine). The high mountain country above extreme
timber line on the highest peaks and ridges. Because of the unusual numberof major mountain peaks in the Teton Range, a considerable proportion of
the park lies within this zone.
(References to life zones of the Tetons: Bailey, 1930: 9-15, colored map
showing the life zones; Cary, 1917, colored map; best sources for this region.)

394

THE AMERICAN MIDLAND NATURALISTThe more recent classification of North America into biotic provinces(Dice, 1943) places all of northwestern Wyoming in the Montanian Province, which here attains its southern limits, thence extending far to the northwest to include the western third of Montana, northern half of Idaho, muchof the southern half of British Columbia, together with the adjoining portionsof Alberta, and a comparatively narrow arm extending westward in northern
Washington. The distribution of Tipulidae confirms that this arrangementof states and provinces is a natural one.
The Fenneman (1931) classification of the United States into physicaldivisions places the Grand Teton region in the Middle Rocky Mountain
Province of the Rocky Mountain System. Mulford's division of the country
on the basis of plant-growth regions (Van Dersal, 1938: 16-27, map) places
all of northwesternWyoming in the Northern Rocky Mountain Region.
All streams within the Park limits eventually drain into the Snake River,a chief tributary of the Columbia. All collections were made to the westof the Continental Divide.
Collecting Stations.-Although collections of crane-flieswere made in many
scattered places within the area under consideration, a few localities proved
to be of such importance that they were made the basis of special investigation.
The chief collections available for this study were those made in 1941
by the author and Mrs. Alexander, with the appreciated co-operation of Mr.
and Mrs. Walter H. Harrison, of Amherst, Massachusetts, our companions
on many trips to various sections of the United States and Canada. These
collections were made between the dates of July 1st and 15th, representing
the late spring and early summer fauna. In 1942, Mrs. Alexander and I
again camped in the Park and collected between the dates of July 1st and10th. As indicated in an earlier paper, this second trip was a great disappointment in that I suffered a serious foot injury on July ist, this necessitating
hospital treatment in Jackson and restricting possible collecting trips to a very
few where transportationby car was possible and but little walking or hobbling
was required in order to secure at least a few specimens (Station 1, Arizona
Creek; Station 3, Hidden Falls; Station 6, Twogwotee Pass: this last proved
to be of exceptional interest and added several species of these flies to thosepreviously known from the area). Besides the above lots of material, totallingsome thousands of specimens, only one further major series from the Teton
Area has been available to me. This included several specimens taken by
Mr. and Mrs. John L. Sperry, of Riverside, California, taken chiefly along
Pilgrim Creek, near Moran, and in the Indian Paintbrush Canyon, in thePark proper. Many of these latter specimens were captured at a date later
than any indicated above and represent the late summer or early autumn
fauna. Among these late season species, especially, were found various additions to the present list.
The base stations established by us in 1941 and 1942 are as follows:
1. Arizona Creek; altitude 6,790 feet, July 1, 2, 8, and 15, 1941; July 4,
1942. Arizona Creek rises on Wildcat Peak in the Teton National Forest, andALEXANDER: NORTH AMERICAN CRANE-FLIES,V.

395

flows southwest into Jackson Lake. At the point where Route 89 (287)crosses the creek unusually rich collecting was found. The country is definitely
in the Transition lifezone, which probably accounts for the fact that manyof the species of Tipulidae found here were different from those of theCanadian or Hudsonian zones in either the Yellowstone or Grand Teton
National Parks. The sparse woody cover along the creek and nearby consists especially of Engelmann spruce, Picea engelmannii (Parry) Engelm.,and alpine fir, Abies lasiocarpa (Hook.) Nutt., with mountain alder, Alnus
tenuifolia Nutt., species of willow, Salix sp., and bog birch, Betula glandulosa
Michx.; further growths of Shepherdia, Cornus stolonifera Michx., and Lonicera involucrata (Richards) Banks. Most of the Tipulidae were swept fromthe rank herbage consisting primarily of Equisetumnarvense L., E. hiemale L.,
Smilacina stellata (L.) Desf., Actaea arguta Nutt., Heracleum lanatum
Michx., Pedicularis groenlandica Retz., Castilleja sp. (red flowered), andRudbeckzaoccidentalis Nutt. Some of the more characteristicTipulidae of
this station include Dicranoptycha quadrivittata Alexander, Dicranota (Dicranota) montana Alexander, Limnophila (Elaeophila) aleator sp. nov.,
Hexatoma (Eriocera) velveta (Doane), Gonomyia (Gonomyia) aciculifera
Alexander, Rhabdomastix (Sacandaga) neolurida Alexander, Erioptera (Psiloconopa) gaspicola (Alexander), E. (P.) shoshone sp. nov., E. (Ilisia) bikpinigera Alexander, E. (I.) manitobensis Alexander, E. (I.) zukeli Alexander, Ormosia megarhabda Alexander, 0. tetonica sp. nov., Molophilus(Molophilus) harrisonisp. nov.; M. (M.) perfiaveolus Alexander. The above
list provides an idea of the extreme richness of this particularstation. Labelled"Arizona Creek."
2. Moran. Teton National Forest; a large bog, 2.4 miles north of thevillage of Moran; altitude 6,800 feet, July 2 and 5, 1941. Around the bog
margin numerous lodgepole pine, Pinus murrayana Balf., with fewer alpine
fir and much aspen, Populus tremuloides Michx. On the hummocks of thebog proper, extensive thickets of willow, the dominant shrub. The herbage
included extensive beds of Equisetum arvense between the willow thickets;
Limnorchis borealis (Cham.) Rydb., L. viridiflora (Cham.) Rydb., Camas?ia
esculenta Lindl., Geranium RichardsoniiF. & M., Mimulus Langsdorfii Sims.,
and Pedicularis groenlandica, the latter in the very wet places. Some of themore interesting crane-fliesincluded Tipula sulphurea jacksonensissubsp. nov.;
Dicranota (Polyangaeus) megalops sp. nov.; Limnophila (Phylidorea) platyphallus Alexander. Labelled "Moran."
3. Hidden Falls, Cascade Creek, Grand Teton National Park; altitude
6,900-7,000 feet, July 3, 4, 1941; July 8, 1942. Collections under this heading include especially those made on shaded earthen banks along the trail
below the falls. These banks were shaded by alpine fir, Greene MountainAsh, Sorbus scopulina Greene; Rubus parviflorusNutt.; Acer glabrum Torrey;
species of Ribes, as R. lacustre (Pers.) Poir., and R. parvulum (Gray) Rydb.,
and other low shrubs. On the moist banks beneath this shade occurred arank growth of liverworts, including Marchantia, and various mosses. In
this habitat occurred Ornithodes harrimani Coq., Dicranota (Polyangaeus)
subapterogyneAlex., Archilimnophila subunica (Alexander) and others. Also

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THE AMERICAN MIDLAND NATURALIST

near the falls occurred various low cliffs and rock outcroppings, on the dripping faces and slopes of which occurred a sparse but characteristiclithophytic
association. Some of the characteristic Tipulidae of this habitat included
Limonia (Dicranomyia) citrina (Doane), Elliptera astigmatica Alexander,
and Phyllolabis lagganensis Alexander. Labelled "Hidden Falls."
4. Cascade Canyon trail to Lake Solitude; boggy area at 8,200 feet,
July 9, 1941. An extensive oxylophytic area on the mountain side, withseveral streams and cold springs. The tree growth included scattered low
trees of alpine fir and whitebark pine, Pinus albicaulis Engelm. Shrubs
scattered about included Salix sp., dominant; Betula glandulosa; Spiraea
densiflora Nutt., and three very characteristicheaths, Kalmia polifolia Wang,
Phyllodoce empetriformis (Smith) Don., and Menziezia ferruginea Smith.
In the streamlets of the boggy areas, various herbs as Limnorchis borealis, L.
viridiflora,Camassia esculenta, Mitella pentandra Hook., and Saxifraga arguta
Don. Characteristic crane-flies included Prionocera primoveris Alexander,
Tipula (Oreomyza) absaroka Alexander, Erioptera (Ilisia) rainieria Alexander, and others. It should be observed that there is a marked resemblance
between this station and the Emerald Pool Station, Yellowstone (Alexander,
1943: 721-722); there the dominant heath was a Ledum while here the latter
was replaced by the three genera listed above. Labelled "Cascade Trail."
5. Jenny and Leigh Lakes; altitude 6,780-6,870 feet; July 3, 4, 6, 10,
12, 1941; Jenny Lake partly repeated in 1942. Collections made along small
mountain streams flowing into these lakes. Approximately a half dozen suchstreams flow down the eastern slopes of the St. John Range into the west sideof Jenny Lake; several other similar streams flow from the foothills of Rockchuck Peak at the northern end of the St. John Range northward into thewestern arm of Leigh Lake. Some of the streams are presumably temporary,
fed by the melting snows from the slopes above, but others are unquestionably
permanent. The largest of these is Cascade Creek, described under Station
3. In such places as the above, the evergreen forest cover is more boreal in
aspect, kinds and number than on the opposite shore of Jenny Lake, the lodgepole pine being replaced chiefly by alpine fir, Engelmann spruce and whitebark pine. Characteristic shrubs include Alnus tenuifolia, Ribes lacustre,
Sorbus scopulina, Rubus parviflorus, and Acer glabrum. Characteristicherbsof the area are Equisetum arvense, Streptopus amplexifolius (L) DC, Mitellapentandra, Saxifraga arguta and Heracleum lanatum. Some of the more noteworthy Tipulidae of this habitat were Tipula (Bellardina) jepsoni sp. nov.,
Pedicia (Tricyphona) degenerata (Alexander), Dicranota (Dicranota) tetonica sp. nov., Ula paupera Osten Sacken, Austrolimnophila badia (Doane),
Limnophila tetonica sp. nov., Gonomyia (Idiocera) shannoni Alexander,
Erioptera (Empeda) tristimonia Alexander, Erioptera (Psiloconopa) aperta(Coquillett), and various species of Ormosia, as 0. albertensisAlexander, 0.
dedita Alexander and 0. hallahani Alexander. Labelled "Jenny Lake, Leigh
Lake."
6. Twogwotee Pass, Teton National Forest; altitude 9,650 feet; July9, 1942. This single day's collecting provided the best high-altitude material

ALEXANDER:NORTH AMERICAN CRANE-FLIES,

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397

found in the area. Close to the summit of the pass on this date, numerous
masses of snow and ice persisted and conditions were very definitely those ofearly spring. A great similarity was noted between this station and the one
at a slightly lower altitude and earlier date reported at Sylvan Lake, Yellowstone, altitude 8,000 feet, June 21, 1941, as previously described (Alexander,
1943: 720). At the present station, Caltha rotundifolia (Huth.) Greene,
was in full bloom and many were in full flower while actually protruding
through masses of snow; also in full flower were Trollius albiflorus (Gray)
Rydb., and Ranunculus eximius Greene. The forest cover consisted principallyof alpine fir and Engelmann spruce, with some limber pine, Pinus flexilis
James. Great thickets of willow and adjoining wet areas produced many
crane-flies. Other excellent collecting was found by sweeping along the margins
of small mountain streams close to the pass. Some of the more noteworthy
Tipulidae of this place and date included Tipula (Yamatotipula) spernax
Osten Sacken, T. (Arctotipula) twogwoteeana sp. nov., T. (Oreomyza) absaroka Alexander, T. (O.) pseudotruncorumAlexander, Ornithodes harrimani
Coquillett, Molophilus rostriferus Alexander, and various species of Ormosia,
as 0. absarokaAlexander and 0. paradisea Alexander. It seems certain that
this particular station will well repay further collecting at a later seasonal
date. A very few additional specimens from here were taken by Mr. andMrs. Sperry, and by Professor G. F. Knowlton and Mr. H. F. Thornley,
the last on September 13, 1941. Labelled "Twogwotee Pass."
I wish to express my deepest thanks to Mrs. Alexander, and to Mr. andMrs. Harrison and Mr. and Mrs. Sperry, for continued aid in collecting theserich materials in the family. Further acknowledgements of kindly aid andadvice are extended to the Park Naturalist, Mr. Carl E. Jepson, and theRanger Naturalist, Mr. Willis T. Smith, of Ogden, Utah. No one could be
more kindly and efficient than these gentlemen and their friendly co-operation
is gratefullv acknowledged.
REFERENCES
1934, 1941-Grand Teton National Park, Wyoming. U. S. Dept. Interior,Nat. Park Service, pp. 1-35, figs.; maps; 1934. Revised and abridged, pp. 1-15,
figs.; map; 1941.
1941-Map of Jackson Hole, Wyoming (Post-Register, Idaho Falls, Idaho);
excellent map of entire region.
1941-A bibliography of National Parks and Monumentswest of the Mississippi
River, vol. 1: Grand Teton National Park pp. 1-16, mimeographed (National
Park Service); approximately 133 titles.
1941-Climate and Man. 1941 Yearbook of Agriculture. U. S.
(VARIOUS AUTHORS)
D. A., pp. 1-1248; Wyoming, pp. 1201-1210, 7 maps.
ALEXANDER, CHARLES P. 1943-Records and descriptions of North American craneflies (Diptera). Part IV. Tipuloidea of the Yellowstone National Park. Amer.
Midl. Nat. 30:718-764, 50 figs.
1940-The physiographic provinces of North America,
ATWOOD, WALLACE W.
pp. 536, 5 col. maps, 1 insert map, 281 figs. (frontispiece, The Grand Teton,in color, by Eugene Kingman).
CARY, MERRITT. 1917-Life zone investigationsin Wyoming. U. S. Biol. Surv., North
Amer. Fauna 42:1-95, col. map, 14 pls., 17 figs.; map shows life zones ofstate, including the Grand Tetons.
Yellowstone National Park, historical
CHITTENDEN, HIRAM MARTIN. No date-The
ANON.

398

THE AMERICAN MIDLAND NATURALIST

and descriptive, pp. 1-356, map (third revision); map includes the northernpartof the Grand Tetons, as far south as Leigh Lake (J. E. Haynes, St. Paul,
Minn.). A comparable work under the same title, 286 pages (fourth edition,
1933; Stanford University Press.)
DICE, LEE R. 1943-The biotic provinces of North America, pp. 78, map.
FENNEMAN, NEVIN M. 1931-Physiography of western United States, pp. 1-534, 173
figs., 1 pocket map.
IRVING, WASHINGTON. Various dates.-Astoria,
pp. 15-464.
STAGNER, HOWARD R. 1938-A
naturalist's guide to Grand Teton National Park.
Jackson Hole Museum Association, Mus. Bull. 1 :1-91; trail maps; trips withmileages; brief survey of flora and vertebrate fauna.
VAN DERSAL, WILLIAM R. 1938-Native
woody plants of the United States, theirerosion-control and wildlife values. U.S.D.A. Misc. Publ. 303:1-362, 44 pls.,
2 pocket maps.

Systematic Account
PTYCHOPTERIDAE

Ptychopteralenis coloradensisAlexander,1937.-Twogwotee Pass, Station
6; 9,650 ft., July 9, 1942.
PtychopterapendulaAlexander,1937.-Arizona Creek,Station 1; 6,790
ft., July 1-5, 1941;July4, 1942;Moranbog, Station2, 6,800 ft., July 5, 1941.
Bittacomorphaclavipes (Fabricius,1781).-Pilgrim Creek, near Moran,
6,800 ft., in wet swales,July 5, 1942.
TRICHOCERIDAE

Trichocerasaltator(Harris, 1782) (gracilisWalker, 1848).-Twogwotee
Pass, Station 6; 9,650 ft., July 9, 1942.
Trichocera tetonensis sp. nov.-General colorationblackishgray, thepraescutumwithoutstripes;knobs of halteresbrownishblack;legs brownish
black to black;wings with a weak brownishtinge, sparselypatternedwith
darker,includinga more distinctcloud over the anteriorcord; male hypopygiumwith the bridgeof the basistylescomplete;dististyleon mesal facewith a flattenedcompressedblade; dense curved setulae on mesal face ofdististylebeyondthe marginalblade.
d. Length,about6 mm.;wing,7 mm.;antenna,about3 mm.
Rostrumand palpi black. Antennae (male) approximatelyone-halfthe
length of body, brownishblack,the outer segmentspalingto whitish. Headdarkbrownishgray.
Pronotumbrownishgray. Mesonotumdarkerblackishgray, especiallythe praescutumand scutum;praescutumwithout differentiatedpattern;scutellumand postnotumclearergray. Pleurablack,sparselypruinose. Halteres
relativelylong, stem obscureyellow, knob brownishblack. Legs with thecoxae dark brown;trochantersobscureyellow; remainderof legs brownish
blackto black. Wings (Fig. 1) with a weak brownishtinge, the prearcularand basal costal fields more whitened;stigma faintly darkened,lying distadof vein R2; a distinct pale browncloud over anteriorcord; cell Cu1 andposteriorcordsomewhatless evidentlyclouded;veins darkbrown,palerin thebrightenedportions. Venation: Scl ending shortlybefore level of R2, Sc.2

ALEXANDER:NORTH AMERICAN CRANE-FLIES,

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399

at near one-fifth the length of Rs; R2+3+4 about one-fourth longer than R2 3;cell Ml a little longer than its petiole; m-cu shortly before fork of M3+4; vein
2nd A angularly bent, its distal portion slightly sinuous.
Abdomen, including hypopygium, black. Male hypopygium (Fig. 2) withthe bridge of the basistyles, b, (so-called coxal bridge) complete, at point ofunion of the lobes produced caudad into a low compressed point. Dististyle,d, elongate, on mesal margin at near one-third the length with a flattenedcompressedlobe or blade that is provided with setae on both faces, about fifteen
longer setae on upper face, somewhat fewer on lower surface; style beyond
the blade a little expanded, with scattered elongate setae at apex and on outersurface; mesal margin as far basad as the blade with short, abundant, curved
setulae. Gonapophyses, g, appearing as relatively broad, elongate, flattened
blades.

Fig. 1. Trichocera telonensis sp. nov.;
venation.
Fig. 2. Trichocera telonensis sp. nov.;
male hypopygium.
Fig. 3. Trichocera garrelti Alexander;
male hypopygium. (Symbols: b, basistyle;d, dististyle; g, gonapophysis.)

Holotype, c3, Hidden Falls, Station 3, 6,900 ft., July 8, 1942 (Alex-

ander).
The most similarspeciesis TrichoceragarrettiAlexander,1927, with awide distributionin southernCanadaand the northernUnited States (Idaho,
Utah). This latterdiffersespeciallyin the structureof the male hypopygium(Fig. 3). Here the bridgeof the basistyles,b, is much largerand more extensive,involvingthe entire mesal face of the basistyle,completelyfused at
the midlineand hereprovidedwith veryabundantlong pale setae. Dististyle,d, with the blade of mesal face much broader,semicircularin outline, theconvexlyroundedouter marginwithoutsetae;setulaeof mesal face of stylelongerand much more numerous,formingan elongateband from near theapex extendingbasadto beyondthe blade of the style, as illustrated.
TIPULIDAE
TIPULINAE

PrionoceraprimoverisAlexander,1943.-Moran bog, 6,800 ft., July 5,
1941; CascadeTrail, Station 4, 8,200 ft., July 9, 1941; TwogwoteePass,
Station6, 9,650 ft., July 9, 1942, the latterpart of the type series.

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THE AMERICAN MIDLAND NATURALIST

Nephrotoma ferruginea (Fabricius, 1805).-Jenny Lake, near camp,
6,780 ft., July 1, 1941.
Nephrotomalugens erythrophrys(Williston, 1893).-Twogwotee Pass,
9,650 ft., July 9, 1942 (M. M. A.).
Tipula (Bellardina) commiscibilisDoane, 1912 (contaminataDoane,1901).-Jackson, Wyoming,August 17, 1941 (J. L. & G. H. Sperry).
Tipula (Bellardina) jepsoni sp. nov.-Allied to faustina; antennal
flagellumbrownishblackto black;mesonotalpraescutumlight gray,with four
dark gray stripesthat are narrowlyborderedby brownishblack;interspaces
with conspicuoussetigerouspunctures;wings creamyyellow, marbledwith
pale grayishbrownand darkerbrown;Rs long, from one-halfto two-thirds
longer than m-cu; petiole of cell M, short, less than one-half m; malehypopygiumwith the ninth tergiteproducedinto a glabrousblackenedtoothon eitherside of midline;setae of tergal lobes relativelyshort, black, from
simple punctures;outer dististyleunusuallylarge and complex,with aboutfour lobes or brades,two of which are providedwith dense and abundant
blacksetae.
d. Length,about22 mm.;wing,21.5 mm.;antenna,about3.8 mm.
W. Length,about29 mm.; wing, 24 mm.; antenna,about4 mm.
Frontalprolongationof head relativelyshort,gray above,more infuscatedon sides; palpi black. Antennaesubequalin length in the two sexes; scapebrownishyellow, more darkeneddistally;pedicelyellow; flagellumbrownish
black to black;flagellarsegmentswith basal enlargementspoorly developed;
longestverticilsmuch exceedingthe segments. Head gray, with a conspicuous median,dark brownstripe; posteriorvertex behind eyes less distinctly
infuscated.
Pronotumbroadlydarkbrownmedially,with a capillarypale centralvitta,
the sidesof scleritepalingto yellow,sparselypruinose. Mesonotalpraescutumwith the restrictedgroundcolor light gray,with conspicuousbrownsetigerous
punctures;four dark gray stripes that are narrowlyborderedby brownish
black,the mediandark line broaderon cephalichalf, becomingvery narrow
to subobsoletebehind;humeraland lateralportionsof praescutummore orless distinctlyinfuscated;scutum with median portion brownishgray, eachlobe with two dark gray areas that are narrowlyborderedby dark brown;
scutellumgray, with a capillarydark brown median line, this obsolete on
caudal portion;mediotergitelight gray on cephalictwo-thirds,darkergray
behind,with a capillarybrownmedianvitta and poorlyindicatedbrownclouds
on eitherside of basal half of disk; pleurotergitegray, the katapleurotergite
abruptlyyellow pollinose. Pleura with the broad dorsopleuralregion lightyellow; remainderof pleura dark gray, variegatedwith lighter gray on theand the more posteriorventralsclerites. Halteres dark
dorsal sternopleurite
brown,the baseof stemand apexof knobslightlymorebrightened.Legswith
the coxae gray, the bases of fore pair more infuscated,of the middle pairbrownishgray;femorayellowto reddishyellow,
less extensivelyso; trochanters
blackened,the amountsubequalon
the tips ratherbroadlyand conspicuously
morethenarrowlyblackened;tarsi black,tips
all legs; tibiae reddishbrown,

ALEXANDER:NORTH AMERICAN CRANE-FLIES,

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401

the basitarsisomewhatpaler on proximalportions;tooth of claws (male)broad but acute. Wings (Fig. 4) with the ground color creamyyellow,
conspicuouslymarbledwith pale grayishbrownand darkerbrown;darkest
areasrelativelyextensive,much more so than in faustina,especiallythe postarculardarkening;areaat anteriorcord virtuallyconfluentwith stigma;yellow
bandbeyondcord wide;outertwo-fifthsof cell R5 pale; darkenedareain cell
Cu distinct, subtendedby yellow ground areas. Venation: Rs long, fromabout one-halfto two-thirdslongerthan m-cu;petiole of cell Ml short, lessthan one-halfm; m-cuslightlyless obliquethan in faustina.

Fig. 4. Tipula (Bellardina) jepsoni sp. nov.; venation.
Fig. 5. Tipula (Bellardina) jepsoni sp. nov.; male hypopygium.
Fig. 6. Tipula (Bellardina)faustina Alexander; male hypopygium. (Symbols: id, inner dististyle; od, outer dististyle; f, tergite.)

Abdomen with basal tergite brownishyellow, darkerand more pruinose
on posteriorportion;succeedingtergitesreddishor reddishyellow,the lateralbordersgray, narrowlyborderedinternallyby dark brown,on the outer segments this latter becomingmore extensiveand conspicuous;basal sternites
similarlyreddishbrown;outersegments,includinghypopygium,moreuniform
segmentsmoreopaqueblack,pruinose;brownishblack. In female,subterminal
ovipositorwith cerci long and straight,more than twice the length of thehypovalvae,their ventralmarginfringedwith long pale setae. Male hypopygium (Fig. 5) with the tergite,9t, somewhatas in faustina;tergitetranson eitherside with a slenderglabrousverse,with a gentlecaudalemargination;
blacktooth, whichin slide mountsare bent cephalad;on the ventralside oftergitewith a broaddepressedplate that is notchedat apex to producetwo
obtuselobes;setae of tergiterelativelyshort, black,from small simple punctures. Outer dististyle,od, unusuallycomplex,on a slide mount appearing
aboutas figured;aboutfour lobesand blades,of whichtwo are providedwith
unusuallyabundantand dense black setae. Inner dististyle, id, with thesensoryarea consistingof only eight or nine pegs. Notch of ninth sternite

402

THE AMERICAN MIDLAND NATURALISTvery shallow, its margin blackened and microscopically corrugated or roughened; no modified flange at base of dististyles.
In faustina (Fig. 6), the ninth tergite, 9t, narrows outwardly, its caudal
margin with an irregular V-shaped notch, the lobes thus formed terminating
in slender, sparsely hairy lobules; surface of tergal lobes provided with abundant long pale setae from conspicuous punctures to produce a roughened appearance; ventral surface of tergite with two very broad, obtuse, darkened
lobes that are separated from one another only by a narrow circular notch;
ventral surface of tergal lobes with scattered transverse groups of from fourto six small setulae, appearing as microscopic rows or bunches. Outer dististyle, od, relatively large but comparatively simple, with only two or threelobes, the vestiture relatively pale. Inner dististyle with the posterior portionof main body relatively stout. Sternal tube unusually short and broad.
Holotype 6, Jenny Lake, Station 5, 6,780 ft., July 4, 1941 (Alexander).
Allotopotype, S caught with the male but not mating, pinned on same pin.
Paratopotype, 1 3, July 6, 1941 (Alexander).
I am very pleased to name this handsome crane-fly in honor of the Park
Naturalist of the Grand Teton National Park, Mr. Carl E. Jepson. All of
the type specimens were swept from rich herbage close to the margin of a large
stream just north of Cascade Creek. The species is closest to Tipula (Bellardina) faustina Alexander, 1941, which differs especially in the venationand pattern of the wings, and in the structure of the male hypopygium, ascompared above. In the subgenus Bellardina Edwards, 1931, there are morethan a dozen species in the Rocky Mountain and Pacific Coast areas, all being
large and unusually handsome flies.
Tipula (Yamatotipula) albocaudata Doane, 1901.-Arizona Creek, 6,790
ft., July 1, 1941.
Tipula (Yamatotipula) colteri Alexander, 1943.-Twogwotee Pass, Station 6, 9,650 ft., July 9, 1942.
Tipula (Yamatotipula) continentalis Alexander, 1941.-Cascade Trail,
Station 4, 8,200 ft., July 9, 1941.
Tipula (Yamatotipula) spernax Osten Sacken, 1877.-Twogwotee Pass,
9,650 ft., July 9, 1942; several swept from willow thickets.
Tipula (Yamatotipula) sulphurea jacksonensis subsp. nov.
c5. Length, about 13 mm.; wing, 11.5 mm.; antenna, about 4.5 mm.
S. Length, about 14-15 mm.; wing, 12 mm.
Differs from typical sulphurea in coloration and in slight structuraldetails.
Frontal prolongation of head above gray, with a narrow, obscure yellow,
median vitta; sides of prolongation more reddish yellow; nasus conspicuous,
especially in female, tufted with long yellow setae. Antennae black, with only
the base of scape slightly reddened; first flagellar segment shorter than insulphurea. Head gray to blue gray, with a narrow brown median vitta.
Praescutal stripes more clearly defined and contrasting with the ground;
scutellum conspicuously infuscated, especially at base; postnotum clear lightALEXANDER:NORTH AMERICAN CRANE-FLIES,

V.

403

gray, more darkened posteriorly, the lateral border of mediotergite and katapleurotergite light yellow, anapleurotergitedarkened. Legs with the apices offemora broadly and conspicuously blackened, on the fore legs including the
distal half or approximately so, on the remaining legs somewhat narrower;
tibiae brown, with blackened tips; tarsi black. Wings with the stigma darker
brown and more conspicuous; a more or less distinctly darkened seam alongvein Cu, in both cells M and Cut. Abdomen with the sublateralblack tergal
stripes broaderand more conspicuous, leaving broad lateral borders;mid-dorsal
tergal stripe narrowerbut very striking.
Holotype, c, Moran bog, Station 2, 6,800 ft., July 5, 1941 (C. P. Alexander). Allotopotype, ? pinned with type. Paratopotypes, 1 c, 3 $ Y.
In its general appearance,the present fly is rather different from the typicalform but upon a careful analysis proves to be nothing more than a race ofthe more eastern and northern sulphurea Doane, 1901. The almost uniformly
blackened antennae and the increase in amount of gray color on the thorax
provide the most evident charactersfor the separation of this race.
Tipula (Tipula) spenceriana hardyi Alexander, 1943.-Moran, August 723, 1941 (J. L. & G. H. Sperry).
Tipula (Tipula) pendulifera Alexander, 1919.-Moran, in wet areas,
August 26-27, 1942 (J. L. & G. H. Sperry). Known hitherto only from
Colorado and Alberta. Very recently discovered in Quebec (La Ferme, in lateAugust, A. Robert); not hitherto recorded from northeasternNorth America.
Tipula (Arctotipula) twogwoteeana sp. nov.-General coloration ofthoracic notum gray, the praescutum with four stripes that are bordered withdark brown; antennae very short,; head and mesothorax with unusually long
and abundant black setae; halteres brownish black; legs with femora brown,
the tips blackened; wings pale brown or brownish gray, variegated with darker
brown and cream-coloredareas, the latter including incomplete bands at originof Rs and beyond the stigma; Rs gently curved, only a little longer than m-cu;
cell 1st M5 small; petiole of cell Ml long; abdomen dark gray, the caudal
and lateral borders of the segments yellow; ovipositor with short compressed
cerci, their tips obtusely rounded.
??. Length, about 16 mm.; wing, 16 mm.; antenna, about 2.2 mm.
Frontal prolongation of head relatively short, dark gray, conspicuously
clothed with long black setae; nasus conspicuous; palpi black, very sparsely
pruinose. Antennae (female) very short, black, the basal segments sparsely
pruinose; flagellar segments with basal enlargements poorly indicated, verticils
long and conspicuous. Head dark brownishgray, with conspicuous erect setae,
especially conspicuous surrounding the antennal fossae and on the conspicuous genae; vertical tubercle undeveloped; anterior vertex broad, nearly fourtimes the diameter of scape.
Pronotum dark gray, variegated with lighter gray, the sides with conspicuous erect setae. Mesonotal praescutum with the ground color gray, with four
stripes, the intermediate pair more brownish, narrowly bordered by darkbrown, the double median margin broad and conspicuous; lateral praescutal

404

THE AMERICAN MIDLAND NATURALIST

stripes with their restricted centers dark gray, the borders brown; interspaces
and lateral borders with conspicuous long black setae; scutum dark graymedially, the lobes somewhat lighter gray, with poorly defined darker central
areas; elongate setae of scutum relatively restricted, more numerous alongmesa1 edges of lobes behind the suture and on lateral portions of sclerite;
scutellum dark plumbeous gray, the parascutella more infuscated; mediotergite short, light gray, with a darker median stripe, the surface of sclerite
on anterior two-thirds with very conspicuous erect setae. Pleura and pleurotergite gray, the mesepisternum, including both anepisternum and sternopleurite, with conspicuous black setae, as in the subgenus; dorsopleural region
weakly infuscated. Halteres brownish black, the extreme base of stem brightened. Legs with coxae light gray, clothed with abundant elongate, chiefly palesetae; trochantersdark gray; femora brown, the tips blackened, somewhat more
extensively so on the fore femora; tibiae dark brown, the tips more blackened;tarsi black; legs relatively short and stout (female). Wings (Fig. 7) with
Fig. 7. Tipula (Arciolipula) taogDoteeana sp. nov.; venation.
Fig. 8. Tipula (Arciotipula) tnogwoteeana sp. nov.; ovipositor, lateral.
Fig. 9. Tipula (Vesliplex) baliopteroides sp. nov.; male hypopygium.
Fig. 10. Tipula (Vesliplex) baliopfera Loew; male hypopygium.

(Symbols: c, cercus; h, hypovalva; id, inner dististyle; 1, tergite.)the groundcolorpale brownor brownishgray,very restrictedlypatternedwith
darkerbrown and with somewhatmore extensivecream-coloredareas; thedarkestareasincludethe stigmaand confluentareasover anteriorcord and atoriginof Rs; the palergroundis somewhatmore intenseand conspicuousin
outer radialfield and as areasin cells M and Cu on eitherside of vein Cu;
markingsare most conspicuousas a narrowpost-stigma1
the cream-coloredband extendingfrom costa into cell R5 and as a more conspicuousoblique
bandbeforeoriginof Rs, extendingfromcosta to vein Cu nearouterends ofcells R and M; furthersmall pale areasin extremeouter end of cell R5, in
vicinityof vein M3+4,in basal half of cell Cu and in marginsof anal cellsofon eitherside of vein 2nd A; prearcularfield deeperyellow. Macrotrichia
veins extremelyreducedin number,as in the subgenus,beyondthe cord with
only aboutthreeor four very scatteredtrichiaover the lengthof distalsectionof R4+5; squamawith one or two setae. Venation: Rs gently arcuated,only
a little longer than m-cu; R1+2 entire; cell 1st M2 small, pentagonal; petiole

threeof cell Ml long, aboutthree-fifthsthe lengthof the cell or approximately
times m; m-cuon M4 some distancebeyondbase;cell 2nd A wide.
Abdomendarkgray,the lateraland caudalbordersof the segmentsnarrow-

ALEXANDER:NORTH AMERICAN CRANE-FLIES,

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405

ly pale yellow; tergites with vestiture much reduced, with scattered small black
setae only; sternites with vestiture similarly inconspicuous, appearing as longer
pale scattered setae; genital segment brownish yellow. Ovipositor (Fig 8)
with cerci, c, short, compressed, the tips obtuse, margins smooth; hypovalvae,h, very short and blunt; lateral border of ninth tergite produced slightly caudad
as a small lobe over the base of cercus.
Holotype, ?, Twogwotee Pass, Station 6, 9,650 ft., July 9, 1942 (C. P.
Alexander); ovipositing in wet earth at margin of small mountain torrent.
The present fly is readily distinguished from other regional members of
the subgenus by the handsomely patterned wings. The fly bears a general
resemblanceto Tipula (Oreomyza) pseudotruncorumAlexander, with which it
was associated in nature, but the subgenericcharactersreadily separate the twoflies. It should be emphasized that such charactersinclude the abundant bodysetae of Arctotipula as contrasted with the more nearly glabrous condition in
Oreomyza, this condition being well shown in the two species of Tipula here
discussed.
Tipula (Vestiplex) baliopteroides sp. nov.-Allied to balioptera;basal
flagellar segments more or less bicolored; wing pattern whitish subhyaline andrestricteddark brown on a pale brownishgray ground; abdomen chiefly yellow,
t4e tergites trivittate with dark brown; male hypopygium with a very conspicuous blackened tergal saucer, armed at anterior and posterior angles withstrong spines, with a further more or less developed discal spine near base of
median notch; inner dististyle stout-stemmed.
3'. Length, about 13-15 mm.; wing, 14-16 mm.; antenna, about 5 mm.
Frontal prolongation of head dark brown above, sparsely pruinose, paler
beneath; nasus elongate; palpi brown, the terminal segment passing into black.
Antennae with proximal three segments yellow, the succeeding segments
usually bicolored, the basal swelling black, the remainder yellow, at about the
fifth to seventh flagellar segments the color becoming uniformly dark; in one
paratype, the bicolorous nature is lost almost on the second or third segment;
flagellar segments rather strongly incised. Head gray, clearer in front, witha conspicuous brown longitudinal line extending from the low vertical tubercle
to the occiput, narrowed behind.
Pronotum dark gray. Mesonotal praescutumgray, with four slightly differentiated clearergray stripes that are conspicuously margined with brown; scutal
lobes each with two similar brown ringed areas; posterior sclerites of notum
gray, with a median brown vitta. Pleura chiefly gray, the dorsopleural membrane yellow; vague patterns of paler gray on the dorsal sternopleuritein somespecimens. Halteres with stem obscure yellow, knob infuscated. Legs withthe coxae pale, gray pruinose; trochanters yellow; femora obscure yellow, thetips brownish black, the amount subequal on all legs; tibiae and tarsi obscureyellow, the terminal tarsal segments darkened. Wings with the ground pale
brownish gray, variegated by slightly darker brown and more extensive whitish
subhyaline areas; the darker pattern includes the stigma, adjoining portionof anterior cord and a small spot over origin of Rs; the pale marks include a
broad band beyond stigma, extending from costa to cell 1st M2, including

406

THE AMERICAN MIDLAND NATURALIST

the bases of cells of the outer radial field; the white obliterative band at cord
is especially conspicuous from outer end of cell R, across 1st M2 into base of
M3; other pale areas at near two-thirds cell M and near basal third of cells
Cu and 1st A; no darkened area at arculus nor any whitening in cells immediately beyond; prearcular and costal fields clearer yellow; veins brown,
more brownish yellow in the brightened fields. Venation: Es long, nearly
three times m-cu; R1+2 long; inner end of cell 1st M2 pointed; petiole of cell
Ml nearly twice m.
Basal abdominal tergite brownish yellow, sparsely pruinose, restrictedly
patterned with darker brown; succeeding tergites bright yellow with three darkbrown stripes that are more or less interrupted at the posterior borders of thesegments; sublateral stripes widened posteriorly; lateral tergal margins pale;sternites yellow, the outer segments more infuscated and pruinose; tergites
eight and nine, and, in cases, tergite seven, more darkened; outer portions ofhypopygium (Fig. 9) with the tergal saucer, 9t, heavily blackened and very
conspicuous, its greatest length nearly equal to the maximum width, narrowed
anteriorly; posterior angles produced caudad into strong horns, the anterior
angles similarly produced, these horns directed chiefly cephalad; margin ofsaucer with further spinous points and denticles; caudal margin with a quadrate
notch, at or near base of which arises a single median horn or spine, as inbalioptera; in the type, as figured, the horn appears to lie very close to thebase of the notch, in other specimens somewhat more removed from themargin. Outer dististyle narrower than in balioptera. Inner dististyle, id.
with the stem much stouter and broader, the fringe of yellow setae sparse andchiefly confined to the distal portion. I have shown the inner dististyle (Fig.
10, id) of balioptera for comparison; here the stem is narrower, the entire
outer surface with abundant long delicate pale setae from very pale and inconspicuous punctures.
Holotype, 3, Pilgrim Creek, near Moran, 6,800 ft., July 5, 1942 (M. M.
Alexander). Paratypes, 1 3, Arizona Creek, Station 1, 6,790 ft., July 8,1941 (C. P. Alexander); 1 3, Gothic, Colorado, 9,500 ft., July 1, 1934
(C. P. Alexander).
The present fly is readily distinguished from the more northern, subarctic
Tipula (Vestiplex) balioptera Loew, 1863, by the hypopygial characters,
especially the details of the tergal saucer and the inner dististyle. The latter
species is known from Saskatchewanand Alberta, the types being from English
River (now Churchill River), Saskatchewan, at near 560 N.Lat., and fromthe then Russian America, all collected by Robert Kennicott.
Tipula (Schummelia) subtenuicornis Doane, 1901.-Jenny Lake, 6,780
ft., July 4-5, 1941; Leigh Lake, 6,870 ft., July 10-12, 1941; Hidden Falls,
7,000 ft., July 8, 1942; Death Canyon, 7,800 ft., July 14, 1941.
Tipula (Oreomyza) absaroka Alexander, 1943.-Cascade Trail, Station
4, 8,200 ft., July 9, 1941; Twogwotee Pass, Station 6, 9,650 ft., July 9, 1942;
part of the type material (Alexander, 1943: 729).
Tipula (Oreomyza) alia Doane, 1911.-A

single female, Death Canyon.

ALEXANDER:NORTH AMERICAN CRANE-FLIES,

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407

7,000 ft., July 14, 1941. In this, Rs is longer than in any other specimen that
I have seen but the identification seems correct.
Tipula (Oreomyza) fundata sp. nov.-Belongs to the borealis or unca
group; most similar to coloradensis and criddlei; antennal flagellum uniformly
blackened; mesonotum gray, the praescutum and scutum conspicuously patterned; tips of femora narrowlyblackened;wing pattern only weakly developed;
abdomen with a conspicuous median, dark brown stripe that becomes obsolete
beyond segment six; male hypopygium with the lateral appendage bearing apair of blackened spinous points; gonapophyses strong, terminating in a spatulate head.
d.
mm.

Length, about 10-13 mm.; wing, 11-15 mm; antenna, about 4-5.2

Y?. Length, about 15 mm.; wing, 14 mm.

Frontal prolongation of head pale, sparsely pruinose; nasus elongate, black;
palpi black. Antennae of moderate length; scape and pedicel yellow, flagellum
black. Head gray, paler gray on front and anterior vertex.
Pronotal scutum brownish gray, with a narrow median infuscation; scutellum yellow. Mesonotal praescutum with the ground color gray, the stripes
barely indicated, best marked by brown borders that form an irregularpattern,
the intermediates widened on anterior portion, very narrow behind, theposterior ends recurved; lateral brown stripes forming strongly curved hookshaped lines; scutum gray, the lobes with scarcely indicated darker gray areasthat are narrowly bordered by pale brown; scutellum and mediotergite gray,with a vague capillary median darkening. Pleura extensively gray, the posteriorsclerites, including the katapleurotergite,more yellow; dorsopleural membrane
buffy yellow. Halteres with stem obscure yellow, knob brownish black. Legs
with the coxae brownish gray; trochanters yellow; femora yellow, the tips
rather narrowly but very conspicuously blackened, the amount subequal on all
legs; tibiae and basitarsi obscure yellow, the tips very narrowly infuscated;outer tarsal segments blackened. Wings with the ground color pale brown,
restrictedly patterned with darker brown and whitish subhyaline; prearcularfield and cell Sc much clearer yellow; stigma dark brown; small darker brownareas at origin of Rs and over anterior cord; somewhat paler clouds in outerradial field, outer end of cell ls't M2 and in cells M and Cu, the two latter
alternating with whitish subhyaline spots; whitish post-stigmal band passing
through cell 1st M2 into bases of adjoining cells; outer end of cell R5 slightly
brightened; veins brownish black, yellow in the flavous areas. Venation:
Rs variable in length, from approximately twice to about three times m-cu.
Abdominal tergites yellow, the first more infuscated; a conspicuous, darkbrown median stripe, ending at about segment six, narrowly interrupted bypale caudal. borders of the segments; sternites and subterminal tergites moreyellowish; ninth segment and eighth sternite chiefly black; genital shield black;
cerci long and slender. Male hypopygium (Fig. 11) relatively large. Ninthtergite, 9t, with the caudal margin truncate, with a small median notch; eachlobe with a conspicuous outer horn on ventral surface, this directed chiefly
caudad. Outer dististyle, od, truncated at apex. Inner dististyle, id, with

408

THE AMERICAN MIDLAND NATURALIST

the setae of dorsal crest relatively inconspicuous, excepting a group or pencil
immediately back of the beak. Lateral appendage, la, with two blackened
teeth or denticles, as in coloradensis and criddlei; pendulous lobe moderately
elongate, broad at base. Gonapophyses, g, strongly developed, broad at base,much narrowed before a conspicuous dilated apical spatula. Eighth sternite
with caudal margin weakly biemarginate to form three blunt lobes that arefringed with abundant long pale setae, those of the central lobe longer andmore conspicuous.

Fig. 11. Tipula (Oreomyza)
fundata sp. n(jv.; male hypopygium.

(Symbols: g, gonapophysis;
id, inner dististyle; la, lateralappendage; od, outer dististyle;s, sternite; t, tergite.)

Holotype, 3, Buffalo Creek, below Twogwotee Pass, 8,000 ft., July 9,1942 (C. P. Alexander). Allotype, ?, Richel Lodge, Montana, August 20,
1942, (J. L. & G. H. Sperry). Paratypes, 1 3, with the allotype; 1 3,
Plateau of Beartooth Mts., Wyoming, 10,000 ft., July 29, 1942 (j. L. & G.
H. Sperry); 3 ', Arizona Creek, Station 1, 6,790 ft., July 4, 1941 (C. P.
Alexander); 3 3, Moran bog, Station 2, 6,800 ft., July 2-5, 1941 (M. M.
Alexander); 3 3, Death Canyon, 7,800 ft., July 14, 1941 (C. P. Alexander);1 ', Twogwotee Pass, 9,650 ft., July 24, 1942 (J. L. & G. H. Sperry).
The two species with which the present fly requires comparison are Tipula(Oreomyza) coloradensis Doane, 1911, and T. (O.) criddlei Dietz, 1914.
Both of these have the lateral appendage of the male hypopygium having two
upper teeth or points, much as in the present fly. The former fly is describedas having the entire abdomen yellowish brown, darker posteriorly, without theconspicuous central darkened tergal stripe; ovipositor with the cerci long andslender, as in the present fly. The latter species, described from Manitoba,
has the abdominal pattern much as in the new form; ovipositor with unusuallyshort valves, the cerci and hypovalvae being of equal length. Both of these
species are described and figured as having the gonapophyses slender, with theapical portion curved into a sickle, not at all as in the insect here defined.
Tipula (Oreomyza) gaspensis Alexander, 1929.-This small fly had been
known only from eastern Canada. It was found commonly at various stations
throughout the Tetons. Arizona Creek, 6,790 ft., July 8, 1941; Moran bog,
6,800 ft., July 2, 1941; Hidden Falls, 7.000 ft., July 4, 1941; jenny Lake,
Station 5, 6,790 ft., July 6, 1941; Leigh Lake, Station 5, 6,870 ft., July 12,
1941; Indian Paintbrush Canyon, 8,000 ft., July 13, 1941; Death Canyon,
7,800 ft., July 14, 1941. As was the case with Quebec specimens, in thismaterial Rs similarly varies in length within rather surprisingly wide limits.

ALEXANDER:NORTH AMERICAN CRANE-FLIES, V.

Tipula (Oreomyza) ingrata Dietz, 1914.-Arizona
6,790 ft., July 2, 1941 (M. M. Alexander).

409

Creek, Station 1,

Tipula (Oreomyza) serta Loew, 1863 (albonotata Doane, 1901).-Pilgrim
Creek, near Moran, July 25, 1942 (J. L. & G. H. Sperry).
Tipula (Oreomyza) pseudotruncorumAlexander 1920.-General coloration
gray, the praescutumyellowish gray with four brownish black stripes, the intermediate pair with an interpolated median dark vitta; antenna short, flagellum
uniformly black; head, scutellum and mediotergite with a central brown vitta;
knobs of halteres dark brown; femora brownish yellow, the tips narrowly
blackened; claws (male) toothed; wings yellowish white, heavily clouded withpale brown and darker brown; Rs long, exceeding twice m-cu; R1+2 entire; cell
1st M2 large; petiole of cell M, short, subequal to or shorter than m; abdominal tergites gray, heavily patterned with brown, the posterior borders of
the segments yellow; male hypopygium with the beak and outer basal lobeof the inner dististyle heavily blackened; eighth sternite with lateral lobes thatbear conspicuous hair tufts.
c.

Length, about 14.5-15 mm.; wing, 16-17 mm.; antenna, about 3 mm.

Frontal prolongation of head relatively long, dark gray; nasus very short;palpi black. Antennae (male) short; scape elongate, transverselycorrugated;pedicel brown; flagellum uniformly black; flagellar segments short, with basal
enlargements feebly developed; verticils longer than the segments; terminal
segment reduced to a small thimble-shaped structure. Head gray; vertex
infuscated medially; vertical tubercle very low and simple.
Pronotum brownish gray, more infuscated above. Mesonotal praescutum
yellowish gray with four brownish black stripes, with a further narrow central
vitta, the intermediate three stripes all narrowed behind and connected at thesuture; lateral stripes narrow, with slightly paler, more brownish gray centers;
humeral region in vicinity of pseudosutural foveae more or less darkened;scutum yellowish gray, each lobe with two brownish gray areas, the more
cephalic one much smaller; scutellum and postnotum clearer gray, with anarrow, interrupted median brown vitta. Pleura and pleurotergite clear gray;dorsopleural membrane yellow, conspicuously variegated with brown areas
along both the dorsal and ventral edges. Halteres brownish yellow, knobs
dark brown. Legs with coxae light gray; trochantersobscure yellow or brownish yellow, the tips narrowly blackened, the amount subequal on all legs;
tibiae dark brown, the tips more blackened; tarsi black; claws (male) toothed.
Wings with the ground pale yellowish white, heavily clouded with pale brown
and with fewer dark brown areas; the darkest clouds include the stigma anda confluent cloud over the anterior cord, origin of Rs, and a spot in basesof cells R and M at arculus; paler brown clouds in distal half of outer radial
field, as seams along outer medial veins, most of cell R, and two conspicuous
clouds in cells M and Cu, separated from one another by a ground area about
opposite origin of Rs; cell 1st A conspicuously clouded, cell 2nd A virtuallyclear except for a narrow axillary margin; the creamy areas include a broad,nearly complete band beyond cord, extending from C to the posterior margin

410

THE AMERICAN MIDLAND NATURALIST

in outer medial field; cells C and Sc; prearcularcells and conspicuous areas incells M to 1st A; veins dark brown, Sc paler at base. Macrotrichia on outerveins, from R4+5 to 2nd A inclusive. Venation: Rs long, exceeding twice
m-cu; R1+2 entire; cell 1st M2 large, exceeding in length vein M3 beyond it;
petiole of cell Ml short, subequal to or less than m.
Abdominal tergites gray, clearest laterally, heavily patterned with brown,
on the first segment this including a narrow mid-dorsal area; tergite two witha continuous dorsal and interrupted sublateral stripes; succeeding segmentswith dorsal and sublateral stripes more extensive and becoming confluent; allsegments with a more yellowish posterior border, this more distinct and broader
on segments four to eight, inclusive; sternites more uniform brownish gray,with conspicuous yellow posterior borders;ninth segment more brownish gray,the posterior margins of both the tergite and sternite more reddish yellow;
dististyles chiefly pale. Male hypopygium (Figs. 12, 13) relatively small andof simple structure; ninth tergite entirely separated from ninth sternite; basistyle complete; accessory sclerite small but developed. Ninth tergite, 9t,
generally rectangularin outline, transverse; caudal margin produced into two
flattened yellow lobes that are separated from one another at the midline only
by a very narrow notch; posterior borders of lobes obliquely truncated, theirsutrfacewith the setae sparse and very small. Ninth sternite, 9s, extensive,

Figs. 12, 13. Tipula (oreom_za)
pseudoiruncorumAlexander; male hypopygium.

(Symbols: b, basistyle; g, gonapophysis; id, inner dististyle; od, outerdististyle; s, sternite; 1, tergite.)with a medianline of pale membrane,on caudalportionwidenedto form a
V-shapedincision;caudomesalmarginsof notch with a small lobe on eitherside, this bearingabundantlong yellow setae that virtuallyfill the whole
incision. Basistyle,b, not producedcaudally,its outer portionprovidedwith
severallong setae. Outer dististyle,od, relativelybroadand flattened. Innerdististyle,id, with both the beak and the outer basal lobe heavilyblackened
and unusuallyshort and compact;posteriorcrest of style thin, pale yellow,

ALEXANDER:NORTH AMERICAN CRANE-FLIES,

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longitudinally striate, its posterior border microscopicallyerose. Gonapophyses,
g, appearing as a pair of very thin, flattened plates. Eighth sternite, 8s,
moderately sheathing, the median area filled with pale membrane; each posterior angle bearing an oval lobe that is completely detached from body ofsternite by pale membrane; each lobe bearing numerous strong setae, someof which are grouped into a more conspicuous apical tuft or pencil directedcaudad and slightly mesad, more or less decussate across midline with its
mate of opposite side.
This rare fly was taken at Twogwotee Pass, Station 6, 9,650 ft., July 9,1942 (M. M. Alexander). It is most closely related to Tipula (Oreomyza)
shoshone Alexander, 1945, and T. (O.) ternaria Loew, 1864, both of which
it somewhat resembles in general appearance. It is readily told by the natureof the wing pattern and by the structure of the male hypopygium, particularlyof the ninth tergite, inner dististyle and eighth sternite. Mrs. Alexander captured these specimens while they were flying high among the spruces on aforested slope not far from a cold mountain torrent. A few others were observed but always so high that they could not be netted. The species had notbeen figured or adequately describedhitherto.
Tipula (Oreomyza)

trivittata laetifica subsp. nov.

W?.Length, about 26 mm.; wing, 20.5 mm.
Close to typical trivittata Say, 1823, differing especially in the distributionof the wing pattern which gives the fly a distinctive appearance.
Arcular darkening restricted; dark crossband at near one-third the lengthof wing widely separated from the brown spot at origin of Rs; wing apex only
restrictedly darkened, the areas in outer ends of cells R2 and R3 solid, in theremaining cells representedonly by narrow but conspicuous dark seams to theveins, leaving the central portions of cells R5 to M4, inclusive, white. In
typical trivittata, these cells are dark with the exception of M3 which is traversed by the complete white band before the stigma and cord; the other cells,in cases, have restricted pale areas but are not whitened as in the present fly;the dark spot at origin of Rs is confluent with the dark band at near one-third
the wing length.
Holotype,?, Jenny Lake, Grand Tetons, 6,780 ft., August 9, 1942

(J. L. & G. H. Sperry).
The discovery of the male sex of the present fly will show whether there
are any hypopygial charactersto support the distinctions indicated above. I
have seen typical trivittata from the Black Hills, South Dakota (Harney Peakand vicinity, 6,500 ft., June 15, 1941, Alexander).
Tipula (Lunatipula) bisetosa percita subsp. nov.
Differs from typical bisetosd Doane, 1901, chiefly in minor hypopygial
details, especially of the eighth sternite. In the present fly, this shows thecushion between the major lateral setae as transverse,its caudal border truncate
or even weakly bilobed, the entire cushion with more abundant setae.

412

THE AMERICAN MIDLAND NATURALIST

In typical bisetosa,the cushion is roughly semicircularin outline, its
posteriorborderevenly roundedto stronglyconvex, the surfacewith fewer
setae.
Holotype, 6, JennyLake,GrandTetons, 6,780 ft., August 10, 1942 (J.
L. & G. H. Sperry). Allotopotype,Y, August 6, 1942. Paratopotype,6,August 9, 1942 (Sperry).
Tipula (Lunatipula) diversa Dietz, 1921.-Jenny Lake, around camp,
July 1-9, 1941;a few, flyingbeneaththe evergreens,associatedwith the much
morenumerouspellucidaand sinistra.
Tipula (Lunatipula) johannus sp.nov.-Allied to barbata;generalcolorationgray,the praescutumwith four narrowbrownstripes;capillarydarkgray
tips of femoranarrowlyblackened;lines on head,scutellumand mediotergite;
wingswith a strongbrownground,the obliterativeareasat cord and beyond
stigma unusuallylarge and conspicuous;abdominaltergitesyellow, trivittate
with brownishblack;sterniteswith an unusuallybroad black centralstripe;
malehypopygiumwiththe lobesof the tergiteobliquelytruncate;ninthsternite
with the lobes large, with conspicuouspencils of long reddishsetae; innerdististylewith beak long and slender,outer basal lobe long and narrow,unmedianarmed;eighthsternitewith the outermajorbristlestronglytuberculate;sternalplate transverse,its centralportionproducedinto a roundedlobe.
6. Length,about12.5 mm.;wing, 12 mm.;antenna,about3.5 mm.
Frontalprolongationof head darkbrown,sparselypruinoseat base above;nasusshortbut distinct;palpiblack. Antennaewith scapeand pedicelyellow,
flagellumblack;flagellarsegmentsmoderatelyincised,a little longerthan theverticils. Head gray,with a conspicuousblackvitta on vertex.
Pronotumgray, with a narrow,dark brown,median dash. Mesonotal
praescutumgraywith four narrowbut conspicuousentirebrownstripes;intermediatepair narrowto obsoleteat their cephalicends, widest at near midlength, separatedby a ground vitta of about the same width; a dusky triangularareain humeralregion;scutumgray,each lobe with two disconnecteddark brownishgray areas;scutellumand mediotergitegray with a capillary
black median vitta. Pleura and pleurotergitegray; dorsopleuralmembrane
buffy. Halteres with stem dusky, narrowlyyellow at base, knob blackened,
its apexa triflepaler. Legs with coxaepale, graypruinose;trochantersyellow;
femorayellow, the tips narrowlyblackened,a very little more extensiveon
fore legs;tibiaeyellow,the tips morenarrowlyinfuscated;basitarsidarkbrown;
remainderof tarsiblack;claws (male) toothed. Wings with a strongbrownish ground,deeperthan in allied species,especiallyin the cells beyondcord;
stigma oval, still darkerbrown;basad of cord the dark groundis more restrictedto the centersof the cells, the vicinityof the veins broadlypale, thisincludingalmostall of cell Cu and the broadbasesof the Anal cells; cells C
and Sc uniformlydarkenedexcept at their whitenedouter ends, cell Sc alittle moreyellowish;whitishobliterativeareasalong cord and beyondstigma
unusuallyextensiveand conspicuousagainstthe ground,includingalmost allof cell RI; dark centralarea of cell 1st A with a whitishelongatemarginal
streak;veins dark brown,brownishyellow in the costal and prearcularfields.

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Venation: Rs moderatelylong, more than one-halflonger than m-cu;R1+2
entire;petioleof cell Ml shorterthan m; cell 1st M2 relativelylong, its outerend pointed.

Fig. 14. Tipula (Lunalipula)johannussp. nov.; male hypopygium.

(Symbols: g, gonapophysis;
id, inner dististyle; od, outerdististyle; s, sternite; t, tergite.)

Abdominaltergiteswith first segmentgray pruinose;succeedingsegments
yellow, conspicuouslytrivittatewith brownishblack, the mid-dorsalstripe
narrowlyinterruptedby pale caudalbordersto the segments;sublateralstripes
similarlybroken,conspicuouslywavy on the more proximaltergites;lateralborderspale; sternitesyellow,with an unusuallybroadand conspicuousblack
mid-ventralstripethat is only narrowlybrokenat the incisures;hypopygium
brownishyellow. Male hypopygium(Fig. 14) complex,as in the barbata
subgroup. Ninth tergite,9t, with the median incisionunusuallydeep andnarrow;laterallobes obliquelytruncate,their outer angles subacute. Ninththe lower
elongate-oval,
sternite,9s, withthe lobe veryconspicuous,transversely
marginwith unusuallylong and conspicuousreddishsetae, the inner group
longer,morecompactand forminga strongbrushor pencil. Outer dististyle,od, long and narrow,as in the subgroup. Inner dististyle,id, with the beaklong and slender;dorsal crest high, with all portionsevenly rounded;outerbasal lobe long and slender,twisted, unarmedwith teeth or spines. Gonapophyses,g, withthe outersclerotizedknobsrounded;innercompressedblades
producedinto a slenderapicalpoint. Eighth sternite,8s, with the armature
lobes conspicuous;at outer lateralportiona smallof the outer semi-detachedcylindricalpedunclebearinga single unusuallystrong and powerfulspinous
seta, its surfacewith microscopictubercles;remainingsetaelong and conspicuous, their marginsfimbriate,those lying closest to the mesal portionof lobe
arisingfrom expandedbases;medianplate of scleritesclerotized,transverse,
the centralportionof the posteriorborderproducedinto a roundedlobe.
Holotype, 6, Jenny Lake, Grand Tetons, 6,790 ft., August 10, 1942
(J.L. S G. H. Sperry).
I am very pleasedto name this fly in honor of Mr. John L. Sperry,towhom I am greatly indebtedfor numerousinterestingWestern Tipulidae.
The fly is alliedto Tipula (Lunatipula)barbataDoane, 1901, and to T. (L.)

414

THE AMERICAN MIDLAND NATURALIST

subbarbata Alexander, 1927, more particularly to the latter. The structureof the male hypopygium, especially as regards the inner dististyle, is more as
in s'ubbarbata,but it differs from this latter fly in almost every detail of the
hypopygium; as regards the inner style, the present species differs in the moreelongate beak, the evenly rounded dorsal crest, and the unarmed outer basal
lobe. The shape and armature of the paired lobes of the ninth sternite of
the present fly is entirely distinct from those of either of the allied species
mentioned.
Tipula (Lunatipula) macrolabis macrolaboidesAlexander, 1918.-Pilgrim
Creek, Moran, 6,800 ft.,>July 25, 1942 (J. L. & G. H. Sperry). Jenny Lake,
Station 5, 6,780 ft., July 4, 1941; August 8-9, 1942 (Sperry); Hidden Falls,
7,000 ft., July 4, 1941 (W. H. Harrison).
I had considered this fly as being identical with macrolabis Loew, 1864,
but now believe it representsa distinct race, best differentiatedby the unusually
narrow arm of the basistyle, at its apex this being only a little wider than thebasal portion. In typical macrolabis, this blade is distinctly widened into a
paddle. From Snodgrass' description and figures of the hypopygium ofspectabilis Doane, 1901, based on type material from Idaho (Trans. Amer.
Ent. Soc., 30:220-221, figs. 156, 158-161; 1904), it is evident that the latter
is a synonym of typical macrolabis.
Tipula (Lunatipula) pellucida Doane, 1912 (clara Doane, 1901).-Shoresof Jenny Lake, July 1-5, 1941; String Lake, 6,870 ft., July 7, 1941; Hidden
Falls, 6,900 ft., July 4, 1941. This is the commonest crane-fly among thelodgepole pines of the eastern shore of Jenny Lake, often associated with T.
(L.) sinistra Dietz.
Tipula (Lunatipula) saxemontana Alexander, 1945.-Arizona Creek, Station 1, 6,790 ft., July 2-8, 1941; Pilgrim Creek, Moran, 6,800 ft., July 5, 1942.
Tipula (Lunatipula) sinistra Dietz, 1921.-Jenny Lake, among the lodgepole pines, 6,780 ft., July 1-12, 1941; July 5, 1942, associated with pellucida;
Hidden Falls, 6,900 ft., July 4, 1941; String Lake, 6,750 ft., July 7, 1941;
Leigh Lake, 6,870 ft., July 12, 1941.
Tipula (Lunatipula) splendens Doane, 1901.-Pilgrim Creek, Moran,
6,800 ft., July 5, 1942; Jenny Lake, 6,780 ft., August 8, 1942 (j. L. & G. H.
Sperry); Leigh Lake, 6,875 ft., July 12, 1941; Indian Paintbrush Canyon trail,
7,500 ft., July 13, 1941, the last relatively abundant in unusually dry places
along the trail; String Lake, 6,870 ft., July 7, 1941, associated with Limnophila (Phylidorea) adusta Osten Sacken.
Tipula (Lunatipula) unicincta Doane, 1901.-Pilgrim Creek, near Moran,
6,900 ft., August 9, 1942 (J. L. & G. H. Sperry).
Tipula (Lunatipula) willis-smithi sp. nov.-Size relatively large (wing,
male, over 15 mm.); general coloration yellowish gray, the praescutum withthree brownish yellow stripes that are narrowly bordered by reddish brown;
antennal flagellum black; wings brownish gray, conspicuously patterned with

416

THE AMERICAN MIDLAND NATURALIST

Rs relatively long, approximately three-fourths longer than m-cu; cell 1st M2
relatively small, pentagonal; petiole of cell M1 subequal to or a little shorter
than m; m-cu at fork of M3+4.
Basal abdominal tergites yellow, the outer segments more reddish brown,
passing into dark brown on outer segments; basal tergite slightly pruinose;
tergites with three little-developed brown stripes, broader and more evident onouter segments; basal sternites clear yellow, the outer segments passing into
dark brown. Male hypopygium (Figs. 16-19) with both the tergite and basistyle entirely cut off from sternite by sutures. Ninth tergite, 9t, a transverse
plate that is completely divided medially by a furrow; caudal margin of either
half heavily blackened and produced into three strong black spinous points,
two of which lie more dorsad, the third immediately beneath the more mesialof these, close to midline. Ninth sternite, 9s, immediately ventrad of basistyle, produced caudad into a long powerful arm that gradually narrows to
an obtusely rounded point, the enlarged base with numerous long coarse setae,
the distal portion clothed with abundant microscopic setulae; ventrad andmesad of this arm projects caudad a second lobe that is shorter but still very
conspicuous, short-stemmed, expanded at apex into a head that is provided
with abundant reddish setae, the more dorsal ones longer and more delicate.
Outer dististyle, od, a small pale, relatively narrow spatula. Inner dististyle,id, about as figured, the actual beak very slender, blackened, slightly twisted,
separated from the shorter and stouter obtuse subapical beak by a roundednotch; outer basal lobe closely applied to main body of style, with a flattened,sclerotized plate lying in the notch; sensory area large, comprised of morethan 25 pegs, placed close to base of style. Gonapophyses appearing as longpowerful blackened spines. Eighth sternite, 8s, sheathing, its dorsal-cephalicregion strongly produced into a lobe; caudal margin with a deep U-shaped
emarginationthat is filled with membranein which is suspended a conspicuous
subquadrate lobe that is densely covered with microscopic setulae; its lower
or ventral surface with a central brush of long reddish setae, the outer or morecaudal ones becoming shorter and less conspicuous; lateral lobes of sternite
bearing a disconnected plate that is separated from the main body by a complete suture, at apex produced into a powerful black horn, probably composedof fused or fasciculate bristles; at base of this latter with several strong setae,
and on mesal face close to base of horn with two smaller hair tufts.
Holotype, c!, near Pilgrim Creek, Teton National Forest, in woods,
6,800 ft., July 5, 1942 (C. P. Alexander). Paratype, 8 , Jenny Lake, Grand
Tetons, 6,780 ft., August 10, 1942 (J. L. & G. H. Sperry).
This fine species is dedicated to the Ranger Naturalist, Professor Willis
T. Smith, of Ogden, Utah. The general appearanceof the male hypopygium
suggests species such as Tipula (Lunatipula) saxemontana Alexander, but theactual structure is entirely distinct. In the latter fly it is the basistyle thatis produced into an arm whereas in the present fly this structure is a direct
prolongation of the ninth sternite, ventrad of the basistyle. The conspicuously
patterned wings likewise are distinctive of the present fly.

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CYLINDROTOMINAE

CylindrotomapallescensAlexander, 1930.-Common in situations described under Station 5; Jenny Lake, 6,780 ft., July 3, 1941; Leigh Lake, 6,870

ft., July 10-12, 1941.
LIMONIINAE
LIMONIINI

Limonia (Limonia) cinctipes (Say, 1823) .-Moran,

August 7, 1941

(J. L. & G. H. Sperry).
Limonia (Limonia) indigena jacksoni (Alexander, 1917).-Jenny Lake,
6,780 ft., July 4, 1941; Leigh Lake, 6,870 ft., July 10-12, 1941; Indian Paintbrush Canyon, 7,500 ft., July 13, 1941.
Limonia (Limonia) sciophila (Osten Sacken, 1877).-Arizona Creek,
6,790 ft., July 3, 1941; Jenny Lake, 6,780 ft., July 3, 1941; Leigh Lake,
Station 5, 6,870 ft., July 10-12, 1941.
Limonia (Limonia) solitaria (Osten Sacken, 1859).-Moran bog, 6,800ft., July 5, 1941; Jenny Lake, 6,780 ft., July 1-3, 1941; Leigh Lake, 6,870 ft.,
July 10-12, 1941.
Limonia (Discobola) annulata (Linnaeus, 1758) (argus Say, 1824).Moran, 6,800 ft., August 23, 1941 (J. L. & G. H. Sperry).
Limonia (Dicranomyia) athabascae (Alexander, 1927).-Moranbog,
Station 2, 6,800 ft., July 2-5, 1941; bog near Jackson Lake, 6,770 ft., July
15, 1941.
Limonia (Dicranomyia) brevivena (Osten Sacken, 1869).-Moran
6,800 ft., July 3, 1941; Jackson Lake, 6,780 ft., July 2, 1941.

bog,

Limonia (Dicranomyia) citrina (Doane, 1900).-Along
Cascade Trail,
Grand Tetons, between 6,900 and 8,800 ft., July 4-9, 1941; again on July 8,
1942; especially numerous on wet rocks (Station 3) below Hidden Falls; atthese lower altitudes on July 4th, only females were found but males werestill flying at the higher levels.
Limonia (Dicranomyia) halterata (Osten Sacken, 1869).-Arizona
6,790 ft., July 1, 1941.

Creek,

Limonia (Dicranomyia) humidicola dampfi (Alexander, 1925).-Jenny
Lake, July 3, 1941; Leigh Lake, July 10-12, 1941; both discussed under Station 5. This is the same form as reported from the Yellowstone as being
humidicola (Osten Sacken, 1859); it now appears that the Rocky Mountainmaterials actually pertain to the subspecies listed above.
Limonia (Dicranomyia) longipennis (Schummel, 1829) (immemor Osten
Sacken, 1861).-Moran, bog, 6,800 ft., July 2, 1941; along Buffalo Creek,near Moran, July 9, 1942.
Limonia (Dicranomyia)

morioides (Osten Sacken, 1860).-Arizona

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THE AMERICAN MIDLAND NATURALIST

Creek, 6,790 ft., July 1-8, 1941; Moran bog, 6,800 ft., July 5, 1941; Buffalo
Creek, near Moran, July 9, 1942; Death Canyon, 7,800 ft., July 14, 1941.
bog,
Limonia (Dicranomyia) penicillata (Alexander, 1927).-Moran
6,800 ft., July 5, 1941; Buffalo Creek, near Moran, July 9, 1942; Twogwotee
Pass, 9,650 ft., September 13, 1941 (Knowlton & Thornley).
Limonia (Dicranomyia) venusta (Bergroth, 1888) (negligens Alexander,
1927) .-Moran, August 24, 1941 (J. L. & G. H. Sperry); Jenny Lake, 6,790
ft., August 5, 1942 (Sperry); Leigh Lake, 6,870 ft., July 10-12, 1941; Hidden
Falls, Station 3, 6,900 ft., July 8, 1942; a few of the individuals were still
teneral.
Antocha (Antocha) monticola Alexander, 1917.-Along
Snake R., Moose, Wyoming, July 6, 1942.

the banks of the

Elliptera astigmatica Alexander, 1912.-On wet rocky outcrops below
Hidden Falls, Station 3, July 4, 1941; July 8, 1942; resting and crawling about
among patches of Saxifraga and Lewisia.
Dicranoptycha quadrivittata Alexander, 1919.-Arizona Creek, 6,790 ft.,
July 8, 1941 (W. H. Harrison); along Pilgrim Creek, 6,800 ft., July 5, 1942;
July 25, 1942 (J. L. & G. H. Sperry); along the Snake R., Moose, Wyoming,
6,800 ft., July 6, 1942; among rich riverside vegetation.
PEDICIINI

Pedicia (Tricyphona) degenerata (Alexander, 1917) .-Jenny Lake, Station 5, 6,780 ft., July 4, 1941; Leigh Lake, 6,870 ft., July 12, 1941, very
common; Death Canyon, 7,000 ft., July 14, 1941; one male from the lastnamed station has the wings even more reduced in size than usual but stillwith the venation clearly defined though distorted ( c3. Length, about 6 mm.;wing, 4 x .38 mm.).
Pedicia (Tricyphona) sparsipuncta (Alexander, 1920).-Jenny
6,780 ft., July 3, 1941.

Lake,

Ornithodes harrimani Coquillett, 1900.-Hidden Falls, Station 3, 6,900
ft., July 8, 1942, resting on the face of perpendicular wet rocks (M. M.
Alexander); Twogwotee Pass, Station 6, 9,650 ft., July 9, 1942; swept fromdense willow patches.
Dicranota (Dicranota) montana Alexander, 1920.-Arizona Creek, 6,790
ft., July 4, 1942 (M. M. Alexander); Hlyingclose to stream margin.
Dicranota (Dicranota) tetonicola sp. nov.-General coloration dark.
gray, the praescutumwith three blackish stripes; antennae short, 13-segmented;
knobs of halteres dark brown; legs black, the femoral bases restrictedly
brightened; wings with a strong blackish tinge, stigma oval, still darker; cellR3 sessile or virtually so, R2+3+4 usually lacking or reduced to a punctiform
element; cell M, present; male hypopygium with the lateral tergal arms very

ALEXANDER:NORTH AMERICAN CRANE-FLIEs,

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419

small, slender-stemmed, terminating in a small head; apex of basistyle produced into a stout dorsal lobe; interbase a strong powerful rod, at apex
expanded into an oval blade that terminates in a small apiculate point.
d3. Length, about 6.5 mm.; wing, 8 mm.; antenna, about 1-1.1 mm.
Rostrumblack, sparselypruinose;palpi black. Antennae short black throughout, 13-segmented; first flagellar segment cylindrical, nearly twice as long as
second; succeeding segments oval, with the lower face a little produced;
terminal segment reduced, about one-third as long as the penultimate; longest
verticils a little shorter than the segments; besides the verticils, an abundantvestiture of short erect dark setulae. Head dark brownishgray; anteriorvertex
broad; vertical tubercle low.
Pronotum dark gray. Mesonotum dark gray, the praescutum with three
blackish stripes, the median one vaguely divided by a paler vitta; scutal lobes
variegated by blackened areas. Pleura and pleurotergite gray, the ventral
sternopleurite darker; dorsopleural membrane infuscated. Halteres relatively
long, stem pale, knob dark brown. Legs with the coxae gray; trochanterstestaceous; remainder of legs black, the femoral bases restrictedly brightened.
Wings (Fig. 20) with a strong blackish tinge; costal border and a seam alongvein 'Cu still darker;stigma oval, even darkerbrown; veins dark brown. Venation: Sc1 ending opposite the supernumerary crossvein in cell RI, Sc2 a

Fig. 20. Dicranota (Dicranota) tetonicola sp. nov.; venation.
Fig. 21. Dicranota (Dicranota) tetonicola sp. nov.; male hypopygium.
Fig. 22. Dicranota (Polyangaeus) megalops sp. nov.; venation.
Fig. 23. Dicranota (Polpangaeus) megalops sp. nov.; male hypopygium.
(Symbols: b, basistyle; d, dististyle; i,interbase; id, inner dististyle; f, tergite.)

distancebeforethe originof Rs greaterthan the lengthof the latter;vein R2
erect; cell R3 sessile or with R2+3+4 indicated by a punctiform element; cell

Ml present;m-cuaboutits own lengthbeyondthe fork of M.
Abdomen black, sparselypruinose,the pleural membraneconspicuously
pale;hypopygiumlarge,black. Male hypopygium(Fig. 21) with the tergite,
9t, broad,concaveacrossthe margin,the lateralanglesproduced,terminatingarm with an expandedor capitatehead.
in a very small, slender-stemmed
Basistyle,b, with the apicallobe largeand stout, with long coarsepale setae;
interbase,i, a strongpowerfulrod, at apex expandedinto an oval bladethat
terminatesin a small apiculatepoint. Outer dististyle,od, broad,the distal

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THE AMERICAN MIDLAND NATURALIST

half with abundantblackenedspinoussetae. Inner dististyle,id, a flattened
elongateblade, its tip narrowlyobtuse.
Holotype, c3, JennyLake,\Station5, 6,790 ft., July 6, 1941 (C. P. Alexander).Paratopotype,1 c3. The types werecapturedby sweepingvegetation
alongbanksof CascadeCreekclose to the lake margin.
Dicranota(Dicranota)tetonicolais entirelydistinctfrom all otherNearctic speciesdescribedto this date. It is well distinguishedby the blackened
wings,with distinctivevenation,and by the structureof the malehypopygium,
especiallythe producedbasistyleand the shapesof the interbaseand lateralarms of tergite.
Dicranota(Plectromyia)reducta(Alexander,1921).-Leigh Lake,Station
5, 6,870 ft., July 10-12, 1941, abundant;Indian PaintbrushCanyon, 7,500
ft., July 13, 1941.
Dicranota (Polyangaeus) megalops sp. nov.-Fully-winged in both
sexes;generalcolorationdark brownor brownishblackthe praescutalstripes
poorly differentiated;antennaewith flagellarsegmentstending to fuse into
groupsof two or three,more especiallythe proximalones; eyes (male) large
and protuberant;
halterespale;tibiaewhitewith both ends narrowlydarkened;male hypopygiumwith the mediantergallobe very narrow,tippedwith aboutfour largesetae.
d. Length, about 4-5 mm.; wing, 4.2-5.2 mm.; antenna,about 0.7 0.9
mm.
9?. Length,about4-4.5 mm.;wing,about4.5-5 mm.
Rostrumand palpi black. Antennae with scape and pedicel brownish
black, flagellumyellowishbrownto dark brown;antennalsegmentsvarying
in numberfrom 12 to 14 segments;basalflagellarsegmentstendingto become
unitedinto one or morepairedsegments,or into a more extensivefusion, insome casesinvolvingthe proximalthreeflagellarsegments;in some cases,theoutermostpairof segmentslikewisemoreor less fused;unfusedsegmentsshortoval to oval. Head dark brownishgray;eyes of male very large and protuberant,broadlycontiguousbeneathbut widelyseparatedby the vertex;anteriorvertexaboutthreetimesthe diameterof scape.
Pronotumrestrictedin size, testaceousbrown. Mesonotumalmostuniform
darkbrownor brownishblack,the threepraescutalstripespoorlydifferentiated.
membrane
Pleurabrownishblack;dorsopleuralpale. Halterespale. Legswith
the coxae testaceous,the fore pair a little paler than the others;trochanters
yellow; femora brownishyellow, the tips narrowlyblackened;tibiae white,with both base and apex narrowlyblackened;proximaltarsal segmentspale
brown,the outersegmentsdarker. Wings (Fig. 22) broad,subhyaline,withcrossveinin cell Sc;
restricteddark clouds, as follows: Over supernumerary
origin of Rs; cord; R2 over all outer supernumerarycrossveins;veins pale
brown,darkerin the cloudedportions. Wings equallydevelopedin bothsexes.
Venation: Supernumerarycrossveinsin cells R3, R4 and M, additionalto
the usualone in cell Rj; venationunusuallyvariable,especiallyas regardscell

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Ml which,whilenormallypresent,may be lacking (as figured) or in processof loss by atrophyof vein M2.
Abdomen, including hypopygium,brownishblack. Male hypopygium(Fig. 23) with the medianregionof tergite,9t, producedinto a slenderlobethat bearsabout four elongatesetae at and near summit;lateraltergal arms
appearingas conspicuousflattenedblades,narrowestat base,expandedat apex
into a semicircularhead; still lateradof the insertionof the blade with two
smallerlobes on the incurvedmarginof the tergite. Basistyle,b, with twoslightlyunequalapicallobes, the outer one stouterthan the inner,both with
abundantshort blackenedspines;interbase,i, a strong powerfulblade, verybroadat base, at aboutthree-fourthsthe length bent almostat a right angle
and thencenarrowedto a long straightpoint;surfaceof interbasebeforethe
bend with abundantmicroscopicsetulae. Dististyle, d, in the notch formedby the lobes of the basistyle,appearingas a slenderyellowblade,broadestat
base,narrowedverygraduallyto the obtusetip whichbearstwo strongspinous
setae;elsewhereon surfacethe setaesparseand reducedin size.
Holotype, c, Moranbog, Station 2, 6,800 ft., July 2, 1941 (C. P. Alexander). Allotopotype, ?, pinned with type. Paratopotypes,1 c, pinned
withtype;numerousadditionaldc dc and a few V ?, July 2-5, 1941 (C. P. e
M. M. Alexander).
Dicranota (Polyangaeus)megalopsis readily told from the only otherdiscoveredRocky Mountain species, D. (P.) subapterogyne(Alexander,1943) by the large eyes of the males and the fully-wingedfemales. Fromthe still more western,D. (P.) maculata(Doane, 1900), it is told by theunusuallylargesize of the head, especiallyacrossthe eyes, and in the detailsof coloration,as the almostuniformdark tone of the body. From a recent
survey of the greatly developedDicranota fauna of Asia, particularlythe
Himalayanregion,it seemsbestto placevariousgroups,includingPolyangaeus
Doane, 1900, as subgeneraratherthan to attemptto maintainthem as fullgenera.
The presentfly was found in a very restrictedarea in the Moran bog.
Here they were found resting on herbs beneaththe willow thicketsand aseries of more than fifty specimenswas taken. Toward eveningthe males
appearedin small swarmsof from six to eight individuals,dancingclose to
the groundalongsidethe willowthickets.
Dicranota(Polyangaeus)subapterogyne(Alexander,1943).-Jenny Lake,
6,780 ft., July 4, 1941; Hidden Falls, 6900 ft., 6,900-7,000ft., July 3-8,
1941;LeighLake,6,875 ft., July 10, 1941;Moran,bog, 6,800 ft., July2, 1941;
all part of type series. For an accountof the habitsand occurrenceof thisfly, with its nearlywinglessfemales,the precedingpart of this seriesshould
be consulted(Alexander,1943: 743).
Dicranota(Rhaphidolabis)cayuga(Alexander,1916).-Jenny Lake,6,780
ft., July 6, 1941; CascadeTrail, aboveStation4, 8,800 ft., July 9, 1941.
Dicranota (Rhaphidolabis)subsessilis(Alexander, 1921).-Jenny Lake,

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brightened costal border; veins brown, yellow in the costal interspaces. Venation: Rs long, angulated and sometimes spurred at origin; R2+3+4 relatively
long, subequal to or slightly longer than cell 1st M2, virtually in longitudinalalignment with Rs; R2 subequal to or longer than R2+3; cell 1st M2 relatively
small, m-cu at or before midlength.
Abdominal segments obscure brownish yellow or light brown, the lateraland posterior borders of the segments darker brown; sternites with the groundsomewhat clearer yellow; a narrow darker subterminal ring; hypopygium
medium brown. Male hypopygium (Fig. 27) with the basistyle, b, relatively
stout, without elongate setae as in angustior. Outer dististyle, od, broader
than in superlineata, the outer flange broadly attached, relatively small andFig. 24. Litnnophila (Elaeophila) aleator sp. nov.; venation.
Fig. 25. Limnophila bigladia
sp. nov.; venation.
Fig. 26. Limnophila fetoniicola sp. nov.; venation.
Fig. 27. Limnophila (Elaeophila) aleator sp. nov.; malehypopygium.

Fig. 28. Limnophila (Elaeophila) shannoni Alexander;
male hypopygium.
)

Fig. 29. Limnophila tetonicola
sp. nov.; male hypopygium.

(Symbols: b, basistyle; g, gonapophysis; id, inner dististyle; od, outer dististyle;p, phallosome; 1, tergite.)inconspicuous.Inner dististyle,id, with only the distal third sclerotizedandprovidedwith setae, the basal portion, exceptinga narrowband along theloweror cephalicborder,consistingof whitemembrane. Phallosome,p, withthe gonapophysesappearingas relativelylarge and conspicuousoval blades,much largerthan in superlineata;
phallosomicplate narrowedoutwardly,not
forming,squareshoulders,as in superlineata.
Holotype, 6, Arizona Creek,Station 1, 6,790 ft., July 2, 1941 (C. P.
Alexander).Allotopotype, ?, July 8, 1941. Paratopotypes,6 cd ?, 6,7906,800 ft., July 1-8, 1941 (C. P. & M. M. Alexander);paratypes,a few c6 S?
5 miles south of Glendale,Utah, along streamseepage,May 5, 1943 (G. F.
Knowlton).
The only generallysimilar regionalspecies is Limnophila(Elaeophila)
superlineata
Doane, 1900,whichis well-distinguishedby the broadwings,witha differentarrangementof pattern,and in the detailsof structureof the malehypopygium,as discussedabove.

424

THE AMERICAN MIDLAND NATURALIST

Limnophila (Elaeophila) angustior Alexander, 1919.-Arizona
6,790 ft., July 4-8, 1941.

Creek,

Limnophila (Elaeophila) shannoni Alexander, 1921.-The types were from
Moscow Mt., Idaho, collected July 25, 1920, by R. C. Shannon. I have notseen any additional specimens until the present material.
Leigh Lake, Station 5, 6,870 ft., July 10-12, 1941. I am transferringboth
this species and L. (E.) bifida Alexander, 1921, from the subgenus Idioptera
Macquart, 1834, where they were originally assigned, to Elaeophila Rondani,
1856. Such a treatment conforms to that of the late Dr. Fred W. Edwards(Trans. Soc. British Ent., 5, pt. 1:76-84; 1938) where, by a readjustment ofcharacters,the various British species previously referred to Idioptera on thebasis of length of the antennae in the male sex were placed in Elaeophila onother charactersof venation and hypopygial structure. Both of the Nearctic
species above mentioned have the male antennae elongate and in this respectdiffer widely from the remaining rather numerous species of Elaeophila, being
evidently more nearly allied to the northern European L. (E.) trimaculata(Zetterstedt, 1838).
Male hypopygium (Fig. 28) with the median region of tergite, 9t, produced caudad into a rounded or oval lobe that is densely clothed with setae.
Outer distyle, od, simple, appearingas a blackened blade, expanded outwardly,
its apex truncated; outer apical portion with microscopic blackened spinulae,
the extreme angle more or less produced. Inner dististyle, id, subequal inlength to the outer style, gradually narrowedto the obtuse tip. Gonapophyses,
g, appearing as oval plates, the apex smooth and evenly rounded, the inner orcaudal margin more roughened. Phallosome and aedeagus short.
Limnophila (Phylidorea) adusta Osten Sacken, 1859.-String Lake, bog,
6,870 ft., July 7, 1941. The first record of this eastern North American
species from the west; hitherto not known from west of Wisconsin and Illinois.
Limnophila (Phylidorea) claggi Alexander, 1930.-Moose Ponds, near
Jenny Lake, 6,700 ft., July 6, 1941; Jenny Lake, 6,790 ft., July 3, 1941; LeighLake, Station 5, 6,870 ft., July 10-12, 1941; Cascade Trail, in Engelmann
Spruce forest, 8,000 ft., July 9, 1941.
Limnophila (Phylidorea) platyphallus Alexander, 1926.-Moran bog, 6,800ft., July 2-5, 1941.
Limnophila (Phylidorea) tepida Alexander, 1926.-Moran bog, 6,800 ft.,
July 2-5, 1941; Cascade Trail, in boggy areas in the Engelmann Spruce forest,
8,000-8,200 ft., July 3-9, 1941. Closely allied to L. (P.) flavipila Doane,
1900.
Limnophila bigladia sp. nov.-Allied to mcdunnoughi; general coloration dark gray, the praescutum with four brown stripes; fore femora extensively blackened; wings cream-colored, with a restricted but conspicuous
brown pattern; outer radial field darkened; area near base of cell R very reduced; cell 1st M2 rectangular; abdominal sternites yellow; subterminal seg-

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425

mentsblackened;male hypopygiumwith the outerdististylebroadlyflattened;
gonapophysesappearingas relativelysmall, slender, divergent blades thatexceedthe aedeagusin length.
c<. Length,about 11-12 mm.; wing, 10-11.5 mm.; antenna,about 1.61.7 mm.
$ . Length,about13 mm.;wing, 12 mm.
Rostrumblack,pruinose;palpi brown,passinginto black. Antennaewith
scapeand pedicelblack;flagellumdarkbrown,the outersegmentseven darker;
relativelylargeand crowded,with verticils
basalflagellarsegmentssubglobular,
on theirdorsalsurfaceonly; on aboutthe sixth and succeedingsegmentswith
verticilson both faces, very long and conspicuous,the segmentselongate.
Head gray, the setigerouspuncturesbrown;head only moderatelynarrowed
behind;anteriorvertexaboutthreetimes as wide as the diameterof scape.
Pronotumrelativelysmall, gray. Mesonotumdark gray, the praescutumwith four brownstripes,the humeralregionlightergray, enclosingthe large,
conspicuous,black pseudosuturalfoveae; scutal lobes with weakly darkened
centers;posteriorscleritesof notumlight gray. Pleuragray, the dorsopleuralregiononly slightlybrightened. Halteresuniformlypale yellow. Legs withfore and middle coxae dark gray, brightenedat apices,posteriorcoxae more
extensivelypale; trochantersreddish;fore femorain types extensivelyblackwith aboutthe proximalthird yellow;middle andened, as in nigrofemorata,
hind femora usually with the tips more narrowlyblackened,especiallythe
middle pair; in other specimens,all femoraequallyand broadlyblackened;
tibiaebrown,the tips narrowlyblackened;tarsiblack. Wings (Fig. 25) withthe ground color creamy,the prearcularand costal fields cleareryellow; arestrictedbut conspicuousbrownpattern,includingthe stigmaand a complete
band at cord; outer end of cell 1st M2; wing-tip, especially in outer radial

field;a largecircularspot at originof Rs; a smallcloudnearouterend of cell
1st A adjoiningthe vein; a small to scarcelyindicateddarkeningin cell R
adjoiningthe vein, aboutmid-distancebetweenarculusand originof Rs, thismuch smallerthan in nigrofemorata.Venation: Rs squareand spurredat
origin; R2+3+4 relativelyshort, up to about one-halflonger than the basal
section of R5; cell 1st M2 rectangular,longer than in related species.

Abdomen yellow; basal tergite and proximalring of second darkened;succeedingtergitesbrownwith the basal lateralportionsyellow;on the fifthand succeedingtergitesmoreuniformlybrown;sternitesclearyellow,the intermediatesegmentswith a weakdarkareaon posteriorborder;eighthand ninth
segmentsblack;styli paler. Male hypopygiumwith the dorsal lobe of basistyle muchproduced. Outer dististyleof aboutthe same shapeas in mcdunnoughi, broadestacrossthe proximalhalf, the greatestwidth at about twothirds the length; apex unequallybidentate. Gonapophysesappearingas
relativelysmall, slender,divergentbladesthat exceedthe aedeagusin length.
Holotype, c6, Death Canyon, Grand Tetons, 7,800 ft., July 14, 1941
c3
(C. P. Alexander).Allotopotype,9, with the type. Paratopotypes,25 6;paratypes,1 9, ArizonaCreek,Station 1, 6,790 ft., July 1, 1941 (C. P. Alexander); 1 dc, Mount Rainier,Washington,2,900 ft., July 28, 1940 (H. &426

THE AMERICAN MIDLAND NATURALIST

M. Townes); 1 6', 1 ?, Banff, Alberta, July 18-25, 1922 (C. B. D. Garrett);
other material in this series in the Canadian National Collection (as mcdunnoughi).
Although closely allied to Limnophila mcdunnoughi Alexander, 1926, andL. nigrofemorataAlexander, 1927, the present fly is distinguished by various
characters, particularly in the structure of the male hypopygium. Frommcdunnoughi it differs in the venation and pattern of the wings and legs, and,especially, in the conformation of the gonapophyses, which in mcdunnoughi
are unusually large and flattened, much more complex in structure. Fromnigrofemorata, it differs in the structure of the hypopygium, particularly theouter dististyle and the gonopophyses (see figure, Proc. U. S. Nat. Mus., 72,
art. 2, pl. 1, fig. 7; 1927; outer dististyle). In their general appearance all
three species much resemble large, showy species of the subgenus Phylidorea
but form a distinct and isolated group of forms that it seems better not toattempt to assign to any particular subgeneric group at the present time.
The main series of types was swept from low grasses and rushes near thebanks of Death Creek, near the Forks. The males were flying very close to
the ground and were much in evidence.
Limnophila occidens Alexander, 1924.-Leigh Lake, Station 5, 6,870 ft.,
July 10-12, 1941; Indian Paintbrush Canyon, 8,000 ft., July 13, 1941; Cascade
Trail, 8,000 ft., in Engelmann spruce association, July 3, 1941.
Limnophila tetonicola sp. nov.-Size large (wing, male, 10 mm. ormore); general coloration of notum brownish gray, the posterior sclerites andthe pleura clearer gray; antennae relatively short, the proximal three flagellar
segments enlarged, the outer ones very slender, with conspicuous verticils;
wings brownish yellow, the oval stigma dark brown; Rs very long, subequal tothe distal section of R5; cell Mt longer than its petiole; m-cu at or beyond
tnidlength of cell 1st M2; abdomen brownish black, the basal sternites with
their central portion obscure yellow; male hypopygium with the outer dististylea nearly straight rod, with a suberect spine before the decurved apex.
d. Length, about 9.5-11.5 mm.; wing, 10.5-13.5 mm.; antenna, about
1.9-2.1 mm.
Rostrum black, brownish gray pruinose; palpi black. Antennae relatively
short; scape brownishblack above, lighter brown beneath; pedicel and flagellum
black; pedicel large, pyriform; basal three flagellar segments large, especiallythe first, outwardly becoming progressively smaller, subcylindrical or with thelower face slightly produced; succeeding segments passing into long-cylindrical,
with long conspicuous verticils, the longest exceeding the segments in length;
terminal segment a little longer than the penultimate. Head brownish gray;anterior vertex relatively wide, about three times the diameter of scape; head
only moderately narrowed behind.
Pronotum moderately massive; scutum brownishgray; scutellum brightened,
obscure yellow medially, infuscated on sides. Mesonotum almost uniform
dark brown, the praescutum without clearly differentiated stripes, the interspaces about concolorous with the stripes, the central region before suture moreALEXANDER:NORTH AMERICAN CRANE-FLIES,

V.

429

ish black,the remainingcoxaetestaceousyellow;trochantersyellow;femora,
tibiaeand basitarsibrownishyellow,somewhatdarkerat theirtips; remainderof tarsi black. Wings (Fig. 30) with a grayishtinge, the prearcularandcostal fields more yellowed;stigma pale brown,ill-delimitedand poorly indicatedagainstthe ground;veins brown. Venation: Sc relativelyshort, Sc1
endingoppositethe originof Rs, Sc2 a distancebeforethis originthat is subequal to m; R2+3+4only slightly arcuated;cell 1st M2 rectangular,only slightly

arcuated;cell 1st M2 rectangular,only slightly widenedoutwardly,subequalin length to vein M4 beyondit; m-cuabout one-fifthto one-sixthits length
beyondthe fork of M.
Abdominaltergitesdarkbrown,the incisuresof the outer segmentspaler;sternitesand hypopygiumyellow. Male hypopygium(Fig. 36) with the outerdististyle,od, shorterthan in extensivena,its darkenedflange longer. Innerdististyle,id, with a single simple blackenedspine. Gonapophysesunequal.
Apex of aedeagusshapedas in the group,the subterminallobes obtuse,thetips nearlyhyaline.
Holotype, c3f,Arizona Creek,Station 1, 6,790 ft., July 4, 1942 (C. P.
Alexander). Paratopotype,6.
Gonomyia(Gonomyia)vafrais rather
Althoughin its generalappearance,
differentfrom G. (G.) extensivenaAlexander,there can be no questionof
betweenthe two flies. The hypopygialdifferenceshave
the close relationships
been discussedabove. In extensivena,the wings are broader,with the individualcells correspondingly
wider,and with vein Sc extendingto a varying
distancebeyondthe originof Rs.
Rhabdomastix(Sacandaga) leonardiAlexander, 1930.-Arizona Creek,
6,790 ft., July 1-8, 1941, July 5, 1942; abundanton low willows along thestream.
Rhabdomastix(Sacandaga)neoluridaAlexander,1943.-The type material
was fromArizonaCreek,6,700-6,790ft., July 8, 1941. Additionalspecimensfrom the same stationJuly 5, 1942.
Rhabdomastix(Sacandaga)subcaudataAlexander,1927.-Cascade Trail,
GrandTetons, belowLake Solitude,8,800 ft., July 9, 1941.
Erioptera(Empeda) tristimoniaAlexander,1943.-Jenny Lake, Station
5, 6,780 ft., July 6, 1941.
Erioptera(Erioptera) septemtrionisOsten Sacken, 1859.-Leigh Lake,
6,870 ft., July 10-12, 1941; Death Canyon,7,000 ft., July 14, 1941 (M. M.
Alexander).
Erioptera (Mesocyphona) distincta Alexander, 1912.-Arizona Creek,
Station 1, 6,790 ft., July 1, 1941.
Erioptera(Symplecta) cana (Walker, 1848).-Arizona Creek,6,790 ft.,
July 2, 1941;Moranbog, 6,800 ft., July 5, 1941;Leigh Lake, 6,870 ft., July
10-12, 1941;PilgrimCreek,nearMoran,6,800 ft., July 5, 1942.

ALEXANDER:NORTH AMERICAN CRANE-FLIES,

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431

Holotype, ', Arizona Creek, Station 1, 6,790 ft., July 2, 1941 (C. P.
Alexander). Allotopotype, i, pinned with the type. Paratopotypes 6 c ?,pinned with the types; numerous additional paratypes, 6,790-6,800 ft., July
1-8, 1941 (C. P. & M. M. Alexander; W. H. Harrison); July 5, 1942 (C. P.
Alexander).
Fig. 30. Conomyia (Gonomyia) vafra sp. nov.;
venation.
Fig. 31. Erioptera (Psiloconopa) shoshone sp. nov.;
venation.
Fig. 32. Eriopfera (Ilisia)
bispinigeraAlexander; venation.
Fig. 33. Eriopfera (Ilisia)
manitobensis Alexander;
venation.
Fig. 34. Erioptera (Ilisia)
rainieria Alexander; venation.
Fig. 35. Erioplera (Ilisia)
zukeli Alexander; venation.
Fig. 36. Gonomyia (Gonomyia) vafra sp. nov.; malehypopygium.
Fig. 37. Erioptera (Psiloconopa) shoshone sp. nov.;
P male hypopygium.
Fig. 38. Erioptera (Ilisia)
bispinigeraAlexander; malehypopygium.
Fig. 39. Erioptera (Ilisia)
rainieria Alexander; m a I ehypopygium.
Fig. 40. Erioplera (Ilisia)
dorothea Alexander; maleg hypopygium.
Fig. 41. Erioptera (Ilisia)
zukeli Alexander; male hypopygium.

(Symbols: b, basistyle; g, gonapophysis; id, inner dististyle; od, outer dististyle;p, phallosome.)

The presentvery distinctcrane-flybelongsto the typicalgroupof the subgenus,now knownto be representedby severalspeciesdistributedthroughout
the Holarctic Region. In our fauna, the only related species is Erioptera(Psiloconopa) cramptonella(Alexander, 1931), of eastern Canada,which
differsin the largersize, venation,and in the very differentmale hypopygium.
In its open cell M2, the presentfly agreesmore closelywith E. (P.) pusilla(Schiner,1865) of Europe,and E. (P.) verna (Alexander,1920), of Japan.
In recentyearsit has becomeincreasinglyevidentthat it will be difficult
or impossibleto maintainas distinctcertainsubgeneraof Erioptera,such as
IlisiaRondani,HoplolabisOsten Sacken,and PsiloconopaZetterstedt. I have

432

THE AMERICAN MIDLAND NATURALIST

discussed this problem in other papers and it seems virtually certain that eventually all three groups will be reduced to a single name, the oldest being
Psiloconopa. In this particularsubgenus several very distinct groups of species
are found, some of which are well representedin the Rocky Mountain region.
Erioptera (Ilisia) bispinigera Alexander, 1930.-Arizona Creek, Station
1, 6,790 ft., July 2-15, 1941. Hitherto known only from the unique typespecimen from Montana. The wing venation is shown (Fig. 32). Male
hypopygium (Fig. 38) with the basistyle, b, at apex produced into a small
conical point that bears several very long setae; mesal face of style near basewith a pale fleshy lobe provided with several short setae; remainder of stylewith scattered elongate setae. Outer dististyle, od, a blackened blade, thestem relatively short, the distal two-thirds expanded into an oblique spatula,
its outer margin with parallel rows of subappressed comb-like teeth. Innerdististyle, id, shorter and smaller, outer or lateral margin produced into two
powerful black spines, the more basal one slightly larger, both spines slightly
recurved; surface of style with numerous conspicuous setae. Phallosome, p,consisting of a powerful blackened plate on either side, broad at base, at near
midlength with the outer portion produced into a long slender spinous rod,the tip acute, the outer margin before apex with a few appressed teeth; on
mesal edge at point of narrowing of style with a smaller knoblike projectionthat is tipped with numerous short blackened spines to produce a macelike
appearance.
Erioptera (Ilisia) margarita Alexander, 1919.-A few specimens taken
along Pilgrim Creek, near Moran, 6,800 ft., July 5, 1942 (M. M. Alexander).
One male shows an interesting abnormality of venation. The right wing(mounted on slide) has cell M2 open by the atrophy of the basal section ofvein M3, producing a Mesocyphona type of venation; the left wing has thecell open by the atrophy of m, producing the normal Erioptera type.
Erioptera (Ilisia) manitobensis Alexander, 1929.-Arizona Creek, Station
1, July 1-8, 1941; numerous. The wing venation is shown (Fig. 33).
Erioptera (Ilisia) rainieria Alexander, 1943.-One male, Cascade Trail,
Station 4, 8,200 ft., July 9, 1941. This was in the alpine fir-whitebarkpineheath association discussed before.
I had hitherto referred this species and the more southern dorothea to thesubgenus Hoplolabis Osten Sacken but it now appears preferrable to place
both in the older subgenus Ilisia Rondani. If Hoplolabis is to be recognized
as a subgenus distinct from Ilisia or Psiloconopa, it must be strictly on thebasis of the spurred or divided cell 1st M2, which would restrict the groupto the subgenotype, armata Osten Sacken, 1859, and two further species,
bipartita Osten Sacken, 1877, and asiatica Alexander, 1919. At best it would
appear that Hoplolabis may be maintained as a weak subgenus for convenience only.
The differences between dorothea and rainieria are best shown in the
structure of the male hypopygium, as follows:
E. (I.) dorothea Alexander, 1914. Male hypopygium (Fig. 40); apicallobe of basistyle, b, very slender, extending caudad beyond the level of theALEXANDER:NORTH AMERICAN CRANE-FLIES,

V.

433

outermost arm of the outer style. Outer dististyle, od, entirely blackened, thethree arms unequal, the central one short, about one-half the length of theouter; outer arm with microscopicspinulae; lower arm longer and more slender,
the tip obtuse. Inner dististyle, id, a long slender rod, entirely pale, strongly
bent at midlength to appear more or less boomerang-shaped,the tip very
obtuse. Gonapophyses, g, blackened, on either side with a subquadrateplate,from the cephalic lateral angle of which arises a long slender spine that is bent
on its base, directed caudad and slightly mesad, very gradually narrowed tothe acute blackened tip; basal plate at outer lateral angle produced into two
blackened teeth. (Colorado-New Mexico).
E. (I.) rainieria Alexander, 1943. Male hypopygium (Fig. 39); apicallobe of basistyle, b, slender, extending caudad about to the level of the outermost arm of outer dististyle. Outer dististyle, od, entirely blackened, with avery short base that is produced into three short arms of approximately thesame shape; outer arm obtuse at tip, its outer surface with appressed acutespinulae, its lower surface merely crenulate; central arm of about the samelength, more narrowed outwardly, the lower surface crenulate; lower arm
shorter and more strongly curved, thickest at near midlength, thence narrowed
to a stout acute point. Inner dististyle, id, a pale stout clavate lobe, strongly
widened on distal half. Gonapophyses, g, blackened, with two principal arms,
a more basal slender spine directed laterad, and a flattened outer blade thatis twisted at near midlength. Ninth tergite large, its caudal margin with adeep U-shaped notch, the lateral lobes more broadly obtuse. (WyomingWashington). The wing venation is shown (Fig. 34).

E. (I.) zukeliAlexander, 1940.-Arizona Creek, 6,790 ft., June 1-8, 1941;
July 5, 1942. Hitherto known only from the unique female type from Idaho.
The present material agrees closely with the type except that in most specimens
the dark pattern of the wing along vein Cu is much more restricted and does
not involve cell M. The wing-venation is shown (Fig. 35). Male hypopygium (Fig. 41) about as shown. Outer dististyle, od, unequally trilobed, withtwo primary flattened blades, with a third smaller lobe on the upper marginof the lower of these blades; outer blade microscopically roughened on outerface, the inner blade more nearly smooth. Inner dististyle, id, entirely paleand fleshy. Gonapophyses, g, appearing as powerful curved lateral horns thatnarrowto acute spinous points, the inner portion close to the midline produced
caudad into smaller acute horns; outer margins of lateral horns with microscopic appressed denticles, the inner margin in cases with a more or lessdistinct denticle.
Cryptolabis (Cryptolabis) molophiloides Alexander, 1943.-Jenny Lake,
6,800 ft., July 13, 1941 (C. P. Alexander); part of the type series. As previously indicated, this series was taken in the dry evergreen forests near thecamp at the lake, the tree species being chiefly lodgepole pine, Douglas fir andalpine fir; no streams or other wet areas were present other than the lake
which was some distance away. The significance of these remarks lies in thefact that although we have no definite knowledge of the immature stages, wehave several reasons to believe that the larvae of members of the genus are434

THE AMERICAN MIDLAND NATURALIST

strictly aquatic. Where the immature stages of the present fly occur remains
very much in question.
Ormosia (Ormosia) absarokaAlexander, 1943.-Twogwotee Pass, Station
6, 9,650 ft., July 9, 1942; abundant along small mountain torrents that werefed wholly or in part by melting snow; associated with 0. (O.) paradisea
Alexander.
Ormosia (Ormosia) albertensis Alexander, 1933.-Arizona Creek, 6,790
ft., July 4, 1942; Jenny Lake, Station 5, 6,780 ft., July 3, 1941; Leigh Lake,
Station 5, 6,870 ft., July 10-12, 1941; Indian Paintbrush Canyon, 7,500-8,000ft., July 13, 1941.
Ormosia (Ormosia) dedita Alexander, 1943.-Hidden Falls, 6,900 ft.,
July 8, 1942; Jenny Lake, Station 5, July 6, 1941; Leigh Lake, Station 5,
July 10-12, 1941.
Ormosia (Ormosia) fusiformis (Doane, 1900).-Jenny
July 6, 1941.

Lake, 6,790 ft.,

Ormosia (Ormosia) hallahani Alexander, 1943.-Arizona Creek, 6,790 ft.,
July 4, 1942; Leigh Lake, Station 5, 6,870 ft., July 12, 1941; Indian Paintbrush Canyon, 8,000 ft., July 13, 1941. This very distinct fly is evidently
widely distributed in the northern Rockies; I have seen it from the Big Horn
Mountains, Wyoming, and the Black Hills, South Dakota.
Ormosia (Ormosia) manicata (Doane, 1900) (deviata Dietz, 1916; fuscopyga Alexander, 1924).-Moran bog, 6,800 ft., July 2-5, 1941; Jenny Lake,near camp, 6,790 ft., July 1, 1941.
Ormosia (Ormosia) megarhabdaAlexander, 1943.-Arizonation 1, 6,790 ft., July 8, 1941.

Creek, Sta-

Ormosia (Ormosia) onerosa Alexander, 1943.-Arizona Creek, Station 1,
6,790 ft., July 4, 1942.
Ormosia (Ormosia) paradiseaAlexander, 1920 (garretti Alexander, 1926).
-Twogwotee Pass, Station 6, 9,650 ft., July 9, 1942; taken by sweeping fir
and spruce along a small, clear mountain stream, very common.
Ormosia (Ormosia) sentis Alexander, 1943.-Arizona Creek, Station 1,
6,790 ft., July 4, 1942.
Ormosia (Ormosia) tetonica sp. nov.-Allied to innocens; generalcoloration blackened, heavily pruinose; antennae short, black throughout, basal
flagellar segments subglobular; halteres black, the knobs obscure yellow; legs
black, the femoral bases more reddish brown; wings yellow, restrictedly
patterned with brown; macrotrichiaof cells unusually sparse, restricted to outercells of wing; cell 1st M2 closed; vein 2nd A gently sinuous; male hypopygiumwith the ninth tergite complex, produced into two slender lobes that are separated by a rounded notch; dististyle a heavily blackened mace-like structure,provided with several blackened denticles.

436

THE AMERICAN
MIDLANDNATURALIST

spicuousU-shaped notch, the tips of the lobes almost touching and thus
closing the notch or virtuallyso. Basistyle,b, with mesal face near apex
bearingan oval lobe providedwith severallong setae. Dististyle,d, a massive
blackenedstructure,the apex and mesalface heavilyblackenedand produced
on mesal face at near
into severalteeth to producea mace-likeappearance;
one-thirdthe lengthbearinga small duskylobe that is providedwith several
setae, two of whichare of unusuallength, much longerthan the lobe itself.
Gonapophysesappearingas flattenedplateswith obtusemargins.
Holotype, c, ArizonaCreek,Station 1, 6,790 ft., July 4, 1942 (M. M.
33 c c3, with
Alexander). Allotopotype,Y, with the type. Paratopotypes,
the type (C. P. & M. M. Alexander);5 dp $, July 1-8, 1941 (C. P. Alexander);the main serieswas taken while flying close to the low overhanging
banksof the stream,just abovethe water;one weeklaterthey had completely
disappeared.
The presentfly is very distinctfrom the two easternNearcticspeciesthat
are most nearlyallied, Ormosia(Ormosia) apicalisAlexander,1911, and 0.
(0) innocens (Osten Sacken, 1869). The great reductionin numberof
in the wingcells is particularly
noteworthy. In both alliedspecies
macrotrichia
these trichiaare much more abundantand are distributedover virtuallythe
entire wing surface,as is commonin the genus. From their reductionin
numberin the presentfly it wouldseem reasonableto expectthat an Ormosia
in the wing cells may be discoveredor that suchwholly lackingmacrotrichia
specieshave alreadybeen describedbut are now placedin the closely related
genus Erioptera,based on the lack of such trichiain the wing cells. It isthus obviousthat the genericdistinctionsbetweenOrmosiaand Eriopteraare
not as well-markedas had formerlybeen believed.
Whetherthe presentfly and its two Nearcticrelativesfall in the subgenus
ScleroproctaEdwards,1938, remainssomewhatin questionbut probablywill
be found to be the case. The chief charactersof this subgenus,other thanthe ones confinedto the malesex, are the glabrouspostnotumand the cephalicpositionof the tuberculatepits of the praescutumwhichlie beforethe transverse level of the pseudosuturalfoveae. However,it should be noted thatvariousothergroupsof Ormos'ia
similarlyhavethe abovementionedcharacters,
these includingthe nigripilagroupand the speciesthat centerabout0. (O.)
prava Alexander,1940, includingthe westernNearctic 0. (O.) hallahani
Alexander.Whetherit is advisableto erectsubgenerafor thesevariousisolatedgroupsof Ormosiaseems somewhatquestionableto me at the presenttime.
It appearscertainthat therewill be some notablediscoveriesin this genus in
the Indo-Chinese
regionthatmay still furthercomplicatethis matter.
Molophilus (Molophilus) colonus Bergroth,1888.-Moran bog, Station
2, 6,800 ft., July 2-5, 1941.
Molophilus (Molophilus) falcatusBergroth,1888.-Jenny Lake, Station
5, 6,780 ft., July 3, 1941;LeighLake,Station 5, 6,870 ft., July 10-12, 1941;
IndianPaintbrushCanyon,7,500 ft., July 13, 1941.

ALEXANDER:NORTHAMERICANV.
CRANE-FLIES,

437

Molophilus (Molophilus) harrisoni sp. nov.-Belongs to the gracilisgroup and subgroup;allied to nitidus; general coloration dark brown or brownish black, sparsely pruinose; head dark brown; pronotal scutellum and thepretergites variegated with yellow; halteres pale yellow; legs brownish black;wings with a strong blackish tinge; R2,3 and basal section of R4+5 long,
petiole of cell M3 short; male hypopygium with the ninth tergite terminating
in a flattened-depressedplate, its caudal margin with microscopic teeth; mesal
lobe of basistyle obtuse; inner dististyle a blackened rod, narrowed on outerportion and provided with abundant blackened spines; aedeagus long andslender.
d. Length, about 3.8-4.2 mm.; wing, 4-4.5 mm.; antenna, about 1-1.1mm.
'?. Length, about 4.5 mm.; wing, 4.8 mm.
Rostrum and palpi black. Antennae relatively short, black throughout;flagellar segments oval; longest verticils slightly exceeding the segments inlength, chiefly unilaterally distributed. Head dark brown.
Pronotum dark brown, the lateral borders of scutellum and the pretergites
obscure yellow. Mesonotum dark brown to brownish black, the surface sparsely
pollinose; scutellum dark. Pleura dark brown, sparsely pruinose; dorsopleural
membrane obscure yellow to brownish yellow. Halteres pale yellow. Legs
brownish black. Wings (Fig. 43) with a strong blackish tinge, the prearcularfield a trifle more brightened; veins and macrotrichiabrownish black. Venation: Sc1 ending about opposite R2; R2+3 long, a trifle exceeding basal sectionof R4+5; cell M3 deep, its petiole variable in length, from subequal to m-cu
to one-half longer than this element; vein 2nd A relatively short, gently
sinuous, ending before level of m-cu.
Abdomen, including hypopygium, black. Male hypopygium (Fig. 45)
with the ninth tergite, 9t, conspicuous, narrowed outwardly, terminating in aflattened-depressedplate, the caudal margin of which is concave and irregularly
toothed. Basistyle, b, with the dorsal lobe, db, at apex narrowed intoa cylindrical yellow rod that terminates in a powerful black spine, at base oflatter with several conspicuous setae; ventral lobe, vb, long, clavate; mesal lobe
short and obtuse. Outer dististyle, od, an elongate blackened rod, uinequallybifid at tip, both lobes obtuse at their tips. Inner dististyle, id, a shorter blackrod, stout on basal half, the outer portion conspicuously narrowed, the surfaceof outer portion with abundant blackened spinous points; surface of enlarged
basal portion with scattered setae. Aedeagus unusually long and slender..
Holotype, c, Arizona Creek, Station 1, 6,790 ft., July 8, 1941 (W. H.
Harrison). Allotopotype, 9, pinned with type. Paratopotypes, 6 c6 S, July
1-8, 1941 (C. P. Alexander & W. H. Harrison); 1 ?, July 4, 1942 (M. M.
Alexander).
This very interesting and distinct fly is named in honor of Mr. Walter H.
Harrison, who has participated in virtually all of our collecting trips during
the past ten years. The nearest ally is the more western Molophilus (Molophilus) nitidus Coquillett, 1905, which differs in the still larger size (Holotype, male, length, 4.5 mm.; wing, 6.2 mm.), more intensely black coloration

438

THE AMERICAN MIDLAND NATURALISTof the bodyand appendages,and, especially,the structureof the male hypopygium. The type of nitiduswastakenat Fieldbrook,California,May 18, 1903,
by H. S. Barberand is preservedin the United States National Museum(Type No. 8347). I am greatlyindebtedto Dr. Henry Townesfor making
a slide mountof the hypopygiumof this type and sendingme a drawingthat
showsthe essentialstructures. Fromthis (Fig. 46) it may be noted that theninth tergite,9t, terminatesin a simple furcula,withoutdenticles;both thedorsal,db, and mesal, mb, lobes of the basistyle,b, are producedinto long
slenderspines of approximately
the same shape. The upper branchof theouterdististyle,od, likewiseis a long slendersinuousrod that narrowsgradually to the acutetip. The drawingfurtherindicatesthat the aedeagusis quite
differentin shapefrom that of harrisoni,near its apex bearinga lateralspur
or branchimmediatelybeforethe decurvedtip. From this figureof the typeI havedeterminedthat nitidusis represented
in my own collectionfromnorthern Californiaand Oregonand that thereis a still differentundescribed
speciesin extremesouthernCalifornia.
Molophilus(Molophilus) perflaveolusAlexander,1918.-Arizona Creek,
Station 1, July 5, 1941; SnowshoeCabin,Wister Draw, at foot of Death
Canyon,6,800 ft., July 14, 1941. The speciesis exceedinglyclose to M. (M.)
auricomusAlexander,1926, of the southeasternUnited States.
Molophilus(Molophilus) rostriferusAlexander,1943.-Twogwotee Pass,
9,650 ft., July 9, 1942;in swampymeadowsnearJacksonLake,6,800 ft., July5, 1942;partof type material.
Molophilus (Molophilus) spiculatus sigmoideus subsp.nov.Very similarto typicalspicUlatusAlexander,1918, differingonly in slight
hypopygialdetails. The outerdististylehas the conspicuoussigmoidshapeinboth racesbut in the presentfly the innerstyle is shorterand broader,particularly acrossits base. In typicalspiculatts the blackenedouter portionis approximatelyone-halfthe total lengthof the outerstyle and the spiculaeof thesurfaceare more sparse,particularlyon basal portion and sides, being bestdevelopedon outer surfaceand on outer half of loweredge. In the present
race,the blackenedouterportionof the innerstyle is only aboutone-thirdthe
lengthof the outerstyle whilethe spiculaearemoreabundantand moreevenly
distributed,occurringvirtuallyto the base.
Holotype, 3, CascadeTrail, GrandTetons, 8,500 ft., July 9, 1941 (C.
P. Alexander). Paratopotypes,3 $?. Paratypes, 3 ?, Moose Ponds, near
JennyLake,6,700 ft., July 6, 1941;LeighLake,Station 5, 6,870 ft., July 1012, 1941; Indian PaintbrushCanyon,7,000-8000ft., July 13, 1941; TwogwoteePass, Station6, 9,650 ft., July 9, 1942 (C. P. Alexander).
The classificationof the westernNearcticmembersof the gracdilisgroup,
pubipennissubgroup,is fully as complicatedand involvedas is the conditionin the speciesof easternNorth America. Bergr6th,in 1888, describedfrom
southernAlaska three species,colonus,falcatusand paulus,that, despitemy
inabilityto studythe typesnow preservedin Finland,I feel havebeencorrect-

ALEXANDER:NORTH AMERICAN CRANE-FLIES,

V.

439

ly identified. The species most in question is colonus and it seems certainthat what Coquillett (Harriman Expedition Report, 1900) determined as thisfrom Alaska and later from British Columbia, is not the true colonus but
actually pertains to the new subspecies of spiculatus described above. Thislatter species has the wings unpatternedwhile Bergroth'sdescription of colonus
calls very definitely for a fly with marked wings. I have identified as colonus
a species previously recorded (Alexander, 1943: 762) from the Yellowstone
and representedin the Teton series. It appears further that comatus (Doane,1900) is a synonym of colonus. Molophilus squamosus Alexander, 1919,
from the extreme southern parts of California appears to represent a subspecies of colonus. The present fly, spiculatus, has the outer dististyle of themale hypopygium more strongly sigmoid than any of these allied species.
Molophilus paulus Bergroth, of which I have seen a broken male from thetype locality (Sitka, Alaska) has the antennal structure and vestiture distinct
from all of the above mentioned members of the pubipennis subgroup. Unfortunately, the abdomen of this specimen is broken and the hypopygium
cannot be described.