Revision of the Stonefly

Family Nemouridae (Plecoptera):

A Study of the World Fauna

at the Generic Level

RICHARD W. BAUMANN

m

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 211
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SMITHSONIA N CONTRIBUTION S TO ZOOLOG Y • NUMBE R 211

Revision of the Stonefly

Family Nemouridae (Plecoptera):

A Study of the World Fauna

at the Generic Level

Richard W. Baumann

SMITHSONIAN INSTITUTION PRESS

City of Washington

1975
ABSTRACT

Baumann, Richard W. Revision of the Stonefly Family Nemouridae (Plecoptera): A Study of the World Fauna at the Generic Level. Smithsonian Contributions to Zoology, number 211, 74 pages, 186 figures, 1 table, 1975.—The Northern

Hemisphere family Nemouridae is revised and divided into two subfamilies:

Amphinemurinae, new subfamily, and Nemourinae. Of the 17 recognized genera

in the Nemouridae, 14 are described and figured and 3 are described as new:

Illiesonemoura, Indonemoura, and Mesonemoura; 373 species are assigned to

genera or listed as incertae sedis. Keys to subfamilies and genera are given for

males, females, and nymphs. Structures of male and female terminalia are

described and the terminology defined. A phylogeny is constructed showing the

relationships of the genera. Distributions are given for all species, the zoogeography of the family is discussed, and a composite map of the distribution of all

genera in Nemouridae is provided.

OFFICIAL PUBLICATION DATE is handstampcd in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year. SI PRESS NUMBER 6007. SERIES COVER DESIGN:

The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Baumann, Richard W.

A revision of the stonefly family Nemouridae (Plecoptera)

(Smithsonian contributions to zoology, no. 211)

Bibliography: p.

1. Nemouridae. 2. Insects—Classification. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology; no. 211.

QL1.S54 no. 211 [QL 505.3.N4] 591'.08s [595.7'35] 75-619077
Contents

Page

Introduction 1

Materials and Methods 2

Terminology 2

Acknowledgments 3

Morphology 3

Terminalia of Males 5

Terminalia of Females 6

Key to the Subfamilies and Genera of Nemouridae 7

Amphinemurinae, new subfamily 10

Amphinemura Ris 11

Indonemoura, new genus 12

Malenka Ricker 13

Mesonemoura, new genus 14

Protonemura Kempny 16

Nemourinae Newman, 1853 17

Illiesonemoura, new genus 18

Lednia Ricker 19

Nemoura Latreille 20

Nemurella Kempny 22

Ostrocerca Ricker 23

Paranemoura Needham and Claassen 24

Podmosta Ricker 25

Prostoia Ricker 26

Shipsa Ricker 27

Soyedina Ricker 28

Visoka Ricker 29

Zapada Ricker 30

Species Incertae Sedis 31

Phylogeny 32

Apomorphic Character States 32

Zoogeography 36

Epilogue ... . 38

Literature Cited 38

Figures 4-186 44

i n

Revision of the Stonefly

Family Nemouridae (Plecoptera)

A Study of the World Fauna

at the Generic Level

Richard W. Baumann

Introduction

The family Nemouridae is the largest in the

order Plecoptera, comprising 373 species in 17

genera. The family was erected by Newman in

1853, and he included most of the present Euholognatha in a large diverse grouping. Klapalek

(1905) divided the family into four families: Cap

niidae, Leuctridae, Nemouridae, and Taeniopterygidae. These four families and the Southern

Hemisphere family Notonemouridae were recently

defined as the superfamily Nemouroidea by Zwick

(1973a).

This study deals with the Nemouridae as proposed by Kimmins (1961) and adopted by the

International Commission on Zoological Nomenclature (1963). Kempny (1898) divided the European species into subgenera; Kimmins (1940),

lilies (1955), Winkler (1957), Hynes (1958), and

Aubert (1959b) recognized four full genera: Amphinemura, Nemoura, Nemurella, and Protonemura. The North American subgenus Paranemoura was described by Needham and Claassen

(1925), and Ricker (1952) revised the North

American Nemouridae fauna and divided the

species into 12 subgenera, including Paranemoura

and also Nemoura and Amphinemura from Europe. These subgenera were then raised to generic

Richard W. Baumann, Department of Entomology, National

Museum of Natural History, Smithsonian Institution 20560.

status by lilies (1966). I describe three new genera:

Indonemoura, Mesonemoura, and Illiesonemoura

and one new subfamily, Amphinemurinae, in this

study.

The Asian Nemouridae have always been placed

in one of the European genera even though they

often did not belong: Aubert (1959a, 1967),

Jewett (1958), Kimmins (1947, 1950 a and b),

Kawai (1967). My study shows that, although

many Asian species do belong in European genera,

some belong in North American genera and others

represent genera and species groups that are endemic to the Asian portion of the Palearctic

region.

The nymphs of the genera occurring in Europe

are quite well known and have been treated by

Hynes (1941), Brinck (1949), Despax (1951), and

Rauser (1956, 1963a). The Asian and North

American genera have, however, been studied very

little. I have studied all available nymphal material, written generic descriptions, constructed

nymphal keys, and used the characters found to

help understand the phylogeny.

The Nemouridae are usually the dominant

Plecoptera in mountain-river ecosystems, both in

terms of total biomass and in numbers of species

present. They are primary consumers and feed

mostly on detritus. They are distributed throughout the Northern Hemisphere and a few species do

just cross the equator in the Sunda Archipelago.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

MATERIALS AND METHODS.—During this study I

examined all of the North American species and

most of the European and Asian species. China is

the only large area from which I was unable to

study many specimens. Those species not examined are marked with an asterisk (*) where they

appear in the "Distribution and Species Lists" at

the end of each genus. This means that they were

FIGURE 1.—Habitus of Zapada haysi (Ricker), mature

female nymph.

placed in a genus based on figures or descriptions

in the literature.

Studies were carried out with a Wild M-5 stereoscopic microscope, which for most of the study was

illuminated with a dark field base illumination

system. This made it possible to study the especially

fine hairs and spines on the small structures of the

epiproct and paraprocts.

For detailed examination all small parts were

removed from the specimens using tiny dental files

or micro dissecting knives. These parts were examined in small watch glasses containing a thin

layer of beeswax and then stored in micro vials

with a cotton plug at both ends. The white-cotton

background made it much easier to see the tiny

parts.

TERMINOLOGY.—Since there are many terms in

existence for both male and female terminalia as

listed in Brinck (1955), I am defining those most

useful for the Nemouridae. I followed the work by

Crampton (1918) because it contained most of the

terms now in use.

Although some authors had recognized the different lobes of the paraprocts, the parts of the

epiproct had been essentially unstudied except for

Zwick (1973a) where the letters "a" and "b are

used for the dorsal and basal sclerites respectively.

The following list of definitions and abbreviations includes those from Crampton (1918) and

those coined in this study.

Basal cushion (be): rounded membranous area of the

epiproct, above the base of the dorsal sclerite, between the

lateral arms; found in Nemoura and closely related

Nemourinae (Figures 45, 61, 118, 133).

Basal sclerite (bs): angular sclerites located on both sides

at the base of the epiproct, between the tenth tergum and

paraprocts, small and triangular or rectangular (Figures

9, 17, 25, 49, 57, 105, 137).

Dorsal sclerite (ds): sclerotized dorsal portion of the epiproct,

with broad base and paired anterolateral extensions, covering all or part of the dorsal and lateral surfaces.

Epiproct (ep): unpaired process in the tergal region of the

eleventh abdominal segment, composed of dorsal, ventral,

and basal sclerites; also called supra-anal process.

Epiproctal flagellum (ef): long thin sclerotized process extending forward from ventral sclerite, at tip of epiproct;

as in Mesonemoura (Figures 28-30).

Hypoproct (hp): produced portion of the male ninth

sternum, containing the gonopore; also called hypandrium

or subgenital plate.

Inner lobe (il): paraproctal lobes located nearest midline,

usually small and inconspicuous, often hidden by hypoproct, lacking spines.
NUMBER 211

Lateral arms (la): thin darkly sclerotized anterolateral extensions of dorsal sclerite, usually connected to dorsal sclerite throughout length.

Lateral cervical sclerite (lcs): long, thin sclerotized bars located near lateral margins of cervical region (Figures 140—

147).

Lateral knobs (Ik): rounded hingelike structures located at

basal corners of epiproct, formed from base of ventral sclerite; found in Nemoura and closely related Nemourinae

(Figures 45, 61, 117, 133).

Outer lobe (ol): outer lobe of paraprocts, located nearest

cerci, long thin and darkly sclerotized in Amphinemurinae,

broad, round and with a membranous portion in Nemourinae; also called cereal lobe.

Median lobe (ml): middle lobe of paraprocts in Amphinemurinae, large and usually both sclerotized and membranous, often bearing spines.

Paraproct (pp): platelike structures situated behind the tenth

tergum, located in the regions homologous with the lateral

portions of the eleventh abdominal segment, may have one,

two, or three lobes; also called subanal lobe.

Pregenital plate (pgp): produced portion of seventh abdominal sternum of female (Figures 51, 67, 75, 123, 131, 139).

Subgenital plate (sgp): modified area of eighth abdominal

sternum of female, located near gonopore; often as broadly

rounded lobe or as one or more sclerotized plates.

Ventral sclerite (vs): ventral sclerotized portion of epiproct,

large, flat or keel-shaped and usually bearing spines, often

extending upward dorsally and modified into single or

paired structures.

Vesicle (v): lobed structure arising from anterior margin of

ninth sternum or hypoproct.

ACKNOWLEDGMENTS.—I heartily thank the following people for helping me with this study by

providing specimens or sharing their time and

ideas: Dr. J. Aubert, Musee Zoologique, Lausanne,

Switzerland; Dr. P. Brinck and Dr. C. H. Lindroth,

Lund University, Lund, Sweden; Dr. P. Berthelemy, Universite de Toulouse, Toulouse, France;

Dr. A. R. Gaufin, University of Utah, Salt Lake

City, Utah, USA; Dr. D. C. Geijskes, Rijksmuseum

van Natuurlijke Historic Leiden, Holland; Dr. J.

lilies, Max-Planck-Institut fur Limnologie, Schlitz,

Germany; Mr. S. G. Jewett, Jr., West Linn, Oregon, USA; Dr. T. Kawai, Nara Women's University, Nara, Japan; Dr. D. E. Kimmins and Mr. P.

Ward, British Museum (Natural History), London, England; Dr. M. Kohno, Aizu-Wakamatsu,

Japan; Dr. A. Lillehammar, Zoological Museum,

University of Oslo, Oslo, Norway; Dr. M. Meinander, Zoological Museum, University of Helsinki,

Helsinki, Finland; Dr. W. E. Ricker, Fisheries Research Board of Canada, Biological Station, Nanaimo, British Columbia, Canada; Dr. B. P. Stark,

Irving, Texas, USA; Dr. K. W. Stewart, North

Texas State University, Denton, Texas, USA; Dr.

L. A. Zhiltzova, Zoological Institute, Academy of

Sciences, Leningrad, USSR; Dr. P. Zwick, MaxPlanck-Institut fur Limnologie, Schlitz, Germany.

I am very grateful to Dr. Joachim lilies, who

made it possible for me to spend a year of postdoctoral study at his institute in Schlitz, Germany,

and I am grateful especially to Dr. P. Zwick for his

friendship, help, and continued encouragement.

The drawings were made by Ms. Elizabeth K.

Meyers, Mr. Michael L. Druckenbrod, Ms. Debra

R. Homer, and Ms. Rebecca F. Surdick. Ms. Hollis

B. Williams provided valuable support in organizing the plates and in the preparation of the

phylogenetic diagram. Special thanks are given to

Mrs. Joan B. Miles for typing the manuscript.

This study was supported in part by a Sigma Xi

Grant-in-Aid of Research and a research fellowship

from the Max-Planck Society.

Morphology

The characters used in this study are best considered by examining body sections and systems as

separate entities. My study was supplemented by

the works of Crampton (1918), Wu (1923), Ricker

(1952), Moulins (1968), and Zwick (1973a).

HEAD.—The head region is very similar in all

nemourid genera. The modification of the last

segment of the labial palpi into a large, flat,

rounded structure is very distinctive, especially in

the adults. Since the adults feed on algae, lichens,

and other plant material, it is possible that the

palpi aid in locating food. It is also possible that

the enlarged palpal segments are sensory in function, because adult nemourids are very active and

drop immediately if the substrate upon which they

are resting is disturbed.

The mandibles and maxillae are broad and

modified for chewing plant material and do not

vary enough from one genus to another to be useful for taxonomic analysis.

Branched gills that originate between the labium and submentum are present in Visoka. These

gills are considered apomorphic because of their

placement and because they are well developed

with five definite branches. They are called submental gills in this study but should not be con-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fused with the submental gills of the Perlodidae,

which originate at the base of the submentum.

NECK OR CERVIX.—The cervical region is mostly

membranous but has long, thin, cervical or jugular

sclerites along the lateral margins. These lateral

cervical sclerites are lightly sclerotized and fairly

distinct. They are excellent points of orientation

because cervical gills, when present, are located

just inside or outside of the sclerites.

Cervical gills are simple extensions of the ventral

tracheae leading to the head. They do not have

any muscles attached but do have osmoregulatory

structures at the base, "Sensilla discoidea" (Zwick,

1973a, and Wichard and Komnick, 1974). Gills are

present in all of the primitive Plecoptera families

and are considered plesiomorphic in the simple

unbranched condition. Modifications such as

branching are considered apomorphic as is the

partial or complete loss of gills. Thus a single,

simple gill on each side of the lateral cervical

sclerites, as exhibited by most Zapada, is considered to be the most primitive condition in

Nemouridae.

Cervical gills are technically called cervical gill

remnants in the adult stage, but because the gills

in Nemouridae are still well preserved in the adult

stage, I am simply calling them gills in all cases.

THORAX (Nymphs).—The pronotum is always

square or rectangular in shape and very stout. The

lateral margins and much of the dorsal surface are

covered with stiff spines. The mesonotum and

metanotum are similar in shape but are slightly

more rounded and contain large wing pads in

mature specimens. The wing pads diverge outward

in a wide angle from the longitudinal body axis.

Spines are usually present along the lateral margins of the mesonotum and metanotum and also

on the dorsal surface, especially along the anterior

margins.

Presence of spines is considered an apomorphic

character state, and they are found on most of the

Nemouridae. Modification into specialized rows

or patterns is a further apomorphic condition.

WINGS.—The wings of Nemouridae are highly

uniform throughout the family. They are very

simple when compared to those of most Plecoptera

families and have very few crossveins. The most

distinctive character is the so-called nemourid "X"

that is formed in the forewings at the cord (Figures

148-157). The "X" is formed primarily by the

terminal portion of vein Sc, which slopes abruptly

to the cord, and by the presence of a costal crossvein running between C and R at or slightly beyond the cord. In Paranemoura the crossvein joins

Sc instead of R, and I consider this a derived character. Zwick (1973a) noted that the "X" in the

forewings was also found in some Capniidae and

Taeniopterygidae. Although it is plesiomorphic,

the "X" is found in all genera of Nemouridae and

is an excellent character for general field and curatorial separation. The "X" can be skewed as noted

by Lillehammer (1974) but is always present. Veins

Ax and A2 are fused near the outer margin of the

forewings in Soyedina. This is an apomorphic state

which is sometimes variable in a single population.

Wing color is not a valid criterion for separation

at the generic level because of the extreme variation possible. It is, however, useful for separating

species complexes as noted by Baumann and

Gaufin (1972).

LEGS.—The legs are generally very uniform

throughout the family. The femur and tibia are of

approximately equal length and tarsal segment

two is much smaller than segment one.

The nymphs can often be distinguished at the

generic level by the kind and pattern of spines on

the legs. All genera have some spines, but they become useful as taxonomic characters when they

are arranged in definite patterns. Zapada, for example, has very specialized whorls of large spines

on all femora.

ABDOMEN (nymphs).—All genera have fringes of

spines along the posterior margins of the abdominal terga. Some species have specialized stout or

long spines arranged in narrow longitudinal rows,

but these arrangements of spines do not help in

separating genera.

ABDOMEN (adults).—The anterior abdominal

segments are seldom modified. Soyedina does, however, contain two species with modified tergites in

the male sex: S. vallicularia Wu has tergites six to

eight elevated, and S. producta Claassen has knobs

on numbers two to four. The males of many species in the Nemoura species complexes Cercispinosa and Ovocercia exhibit modifications of tergites eight and nine. Some Amphinemura and

Mesoncmoura species have greatly modified ninth

terga, such as A. mirabilis Martynov and A. zimmermani Joost, which are also asymmetrical.

Protonemura contains many species that have
NUMBER 211

produced areas covered with spines on terga seven,

eight, and nine.

In most genera tergum ten is sclerotized and

formed into a flat or concave base under the epiproct. It also bears large spines or prongs in some

species of Ostrocerca and the Cercispinosa and

Ovocercia complexes. Thin paired sclerotized bars

present in Nemurella, Ostrocerca, Podmosta, and

Shipsa run laterally and attach to the bases of the

cerci instead of the usual single, wide, median bar.

In Podmosta, Paranemoura, and Lednia the lateral

margins are enlarged and modified into specialized

structures that are covered with spines or hairs.

Shipsa has even larger lateral projections which

extend backward beyond the tips of the cerci.

TERMINALIA OF MALES

HYPOPROCT.—The hypoproct is usually formed

into a broad, flat plate which is wide at the base

and tapers to a narrow tip. It is greatly enlarged

and very elongate in Soyedina, Nemurella, and

some species of Ostrocerca. The apex is enlarged

and terminates in a truncate tip in other species

of Ostrocerca. The tip bears the genital opening,

and the length of the apex is dependent on the

structure of the paraprocts and epiproct because

they usually function together to transport sperm

during mating (Zwick, 1973a).

A vesicle is present at the anterior margin of the

hypoproct or ninth sternum in all genera except

Paranemoura and Lednia. The vesicle is a sensory organ as noted by Rupprecht and Gnatzy

(1974). I consider the absence of a vesicle to be

apomorphic.

EPIPROCT.—The epiproct is actually composed of

four sclerotized structures: dorsal sclerite, ventral

sclerite, and two basal sclerites.

The basal sclerites are paired angular sclerotized plates located at the basolateral corners.

Zwick (1973a) calls these the "b" sclerites. They

are present in all genera and vary in shape from

small and triangular to large and rectangular or

ovoid.

The dorsal sclerite is composed of a large darkly

sclerotized horseshoe-shaped base, lateral sclerotized arms, and a large lightly sclerotized dorsolateral area made up of many tiny scales or plates.

Zwick refers to the horseshoe-shaped base and the

lateral arms as part of sclerite "a." Nemoura and

closely allied species of Nemourinae have a large

membranous dorsal area located directly ahead of

the horseshoe-shaped base of the dorsal sclerite. I

call this area the basal cushion.

The ventral sclerite is the flat, convex or keelshaped ventral portion of the epiproct. It is connected to the tenth tergum by a single or paired

sclerotized bars. The ventral sclerite usually bears

spines or prongs somewhere along the ventral margin. It is often highly modified dorsally and extends upward inside the lateral folds of the dorsal

sclerite. The dorsal portion can be single but is

usually paired. It is often visible on the dorsal surface of the epiproct and extends forward as a flagellum in Mesonemoura and rarely in other genera

of Amphinemurinae. In Nemoura, Illiesonemoura,

Zapada, and Soyedina the lateral basal corners are

formed into distinct lateral knobs resembling ball

and socket hinges. This derived character often

obscures the lateral arms of the dorsal sclerite as

they extend forward from the horseshoe-shaped

base.

I consider the plesiomorphic state of the epiproct as being similar to some species of Amphinemura where the dorsal and ventral sclerites are

similar in size but are little modified. They bear

spines somewhere ventrally and are connected to

the tenth tergum by a single wide sclerotized bar.

PARAPROCTS.—These structures are very stable

and provide excellent taxonomic characters. They

are located ventrally in the region of the eleventh

tergum. They are paired and separated by the midline into bilateral structures. They can be one-,

two-, or three-lobed.

The subfamily Amphinemurinae has paraprocts

with three lobes, and the subfamily Nemourinae

has paraprocts with one or two lobes. Those with

three lobes have small sclerotized inner lobes which

are naked of spines. The inner lobe has a muscle

connected along the inner margin. The median

lobes are large, well developed, and have a sclerotized base and membranous apex which often bears

a few spines or prongs. There is a muscle that connects to the lobes near the apex and extends

throughout the length. The outer or cereal lobes

are usually small, thin and are located near the

base of the cerci. They are darkly sclerotized and

sometimes bear spines or prongs. These lobes have

a muscle attached at the base.

In the Nemourinae, where only two lobes are
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

present, the inner lobes are narrow and often difficult to see because they are located on the inner

margins under the apex of the hypoproct. Some

genera like Nemurella, Ostrocerca, Visoka, and

Lednia have well-developed inner lobes which extend to the base of the epiproct and possibly aid in

sperm transport as noted by Zwick (1973a). The

outer lobes are usually broad and membranous but

sometimes have a sclerotized base. Where a single

lobe is present, the inner lobe has been lost and

the outer lobes cover the complete ventral region.

The primitive Plecoptera families like Austroperlidae, Pteronarcyidae, and Scopuridae have

single large paraproctal lobes. The apomorphic

condition is a divided paraproct, but in the Nemouridae the condition of a large outer lobe and

tiny inner lobe like in Nemoura is plesiomorphic

and the three-lobed condition is derived.

TERMINALIA OF FEMALES

PREGENITAL PLATE.—The seventh sternum is

often enlarged and modified into a pregenital

plate. The median posterior margin is produced

and lightly sclerotized and extends onto sternum

eight. Many species in the Nemourinae have a

highly developed pregenital plate which covers

the genital opening and much of the eighth sternum. Modification of sternum eight is the usual

condition in Plecoptera, so formation of a pregenital plate is taken to be apomorphic.

SUBGENITAL PLATE.—The eighth sternum is usually modified into some sort of subgenital plate.

The Amphinemurinae are characterized by having

paired vaginal lobes on the posterior margin.

Protonemura, Indonemoura, and Mesonemoura

also have a large median plate, and Malenka and

Amphinemura usually have a distinct median

notch.

The Nemourinae show less development of

sternum eight. It is usually narrow and covered by

sternum seven. It is developed laterally in Nemurella and also in some Ostrocerca species. Lednia

and Shipsa have small platelike areas medially and

Podmosta exhibits distinctive sclerotized patterns.

In Prostoia, segments seven and eight are essentially unmodified.

CERCI.—Nymphs: The cerci of the nymphs are

quite long and covered with spines. They are approximately the length of the complete abdomen.

Whorls of spines are found at the distal margins of

all segments. The length of the spines varies even

between species and sometimes the ventral spines

or bristles are slightly longer as shown by Harper

and Hynes (1971). Intermediate spines are often

present on the segments between the joints. No

good generic characters have been found using the

cerci, but they are useful at the specific level in

small genera and when considering species in

limited geographical areas.

Adults: The adult cerci are reduced to single

segments. They are usually simple, membranous,

and covered by short hairs. The only departures

from this pleisomorphic state in the Amphinemurinae are the lengthening of the segments as in

Indonemoura and the development of a mesobasal

sclerotized, lobe by the males of Malenka. The

mesobasal lobe is located at the base of the cerci

on the dorsolateral margin.

Most of the Nemourinae also have simple cerci,

but Nemoura males have a distinctive lateral

sclerotized portion which usually ends in a multiforked tip at the apex. In the Cercispinosa complex large spines may occur anywhere throughout

the length of the cerci. The Ovocercia complex

has enlarged cerci that are lightly sclerotized and

modified in shape but do not bear spines. Visoka

also has sclerotized cerci that end in single projections.

Nemurella and Ostrocerca males have greatly

enlarged sclerotized cerci. They are elongate and

globose in Nemurella and naked of projections

but they have a greatly enlarged base. In Ostrocerca they are thin, elongate, and terminate in

sharp points. They are bent in an S-shape in lateral aspect in some Ostrocerca species.

The females of most genera have simple, unsclerotized cerci, but those of Nemoura, Nemurella, and Visoka are lightly sclerotized laterally

and slightly modified in shape.

I submit that the occurrence of sclerotized, modified cerci in scattered genera of Nemourinae is the

result of convergence.
NUMBER 211

Key to the Subfamilies and Genera of Nemouridae

MALES

1. Paraprocts divided into three.lobes; with spines or prongs on middle or outer lobes

(Figures 7, 15, 23, 31, 39) (Amphinemurinae, new subfamily) 2

Paraprocts single or divided into two lobes; lacking spines on outer lobes (Figures 63,

71, 95, 119, 127) (Nemourinae) 6

2. (1) Cervical gills as two highly branched structures on each side of midline, one inside and

one outside of lateral cervical sclerites (Figures 140, 141) 3

Cervical gills simple, reduced or partially absent, those outside of lateral cervical

sclerites may be forked (Figures 142, 143) 4

3. (2) Mesobasal lobe present at base of cerci in dorsal aspect (Figure 24); gill branches not

all extending to gill base (Figure 141) (Nearctic) Malenka

Mesobasal lobe absent (Figure 8); gill branches all extending to gill base (Figure 140)

(Holarctic, Oriental) Amphinemura

4. (3) Cervical gills simple inside of lateral cervical sclerites and forked outside (Figure 143)

(Palearctic) ProUmemura

Cervical gills absent inside of lateral cervical sclerites and reduced to short stubby

knobs outside (Figure 142) 5

5. (4) Paraprocts with three well-defined lobes, median lobes greatly enlarged with large apical

prongs (Figure 15); epiproct without epiproctal flagellum or only very small process

present (Figure 13) (Oriental) Indonemoura, new genus

Paraprocts with lobes generally fused near base, median lobes relatively small, with

hairs or spines apically (Figure 31); epiproct with terminal flagellum present (Figure

29) (Oriental, Palearctic) Mesonemoura, new genus

6. (1) Lateral knobs present at basal corners of epiproct; basal cushion present at dorsal base

of epiproct (Figures 45, 61, 117, 133) 7

Lateral knobs absent from basal corners of epiproct; basal cushion absent from dorsal

base of epiproct (Figures 69, 77, 93, 101, 125) 10

7. (6) Cervical gills present 8

Cervical gills absent 9

8. (7) Simple cervical gills present both inside and outside of lateral cervical sclerites (Figure

147); paraprocts large but without inward directed projection near apex (Figure 135)

(Nearctic) Zapada

Simple cervical gills only present outside of lateral cervical sclerites (Figure 144);

paraprocts large and with an inward directed process near apex (Figure 47) (Palearctic, Oriental) IUiesonemoura, new genus

9.(7) Paraprocts elongate and greatly enlarged (Figure 119); epiproct bilaterally asymmetrical

(Figures 116-118); cerci small, unsderotized and unmodified (Figure 120); veins A,

and A, joined near margin of forewings (Figure 152) (Nearctic) Soyedhta

Paraprocts large and rectangular (Figure 63); epiproct bilaterally symmetrical (Figures

60-62); cerci large, mostly sclerotized, with large hooks or spines or bulbous at base

(Figure 64); veins A, and A, not joined in forewings (Figure 154) (Holarctic,

Oriental) Nemoura

10. (6) Dorsal sclerite of epiproct large and lateral arms well developed (Figures 68, 78, 92,

108) H

Dorsal sclerite of epiproct reduced in size and lateral arms poorly developed (Figures

52, 84, 100, 124) H

11.(10) Cerci greatly enlarged and modified (Figures 72, 80); paraprocts bilobed, inner lobe

large and heavily sclerotized and outer lobe mostly membranous (Figures 71, 79) ...12

Cerci small and unmodified; paraprocts with single large lightly sclerotized lobe (Figures 95, 111) 13

12.(11) Cerci long, narrow throughout length and pointed at apex (Figures 80-82) (Nearctic)

Ostrocerca

Cerci stout, enlarged at base and with broadly rounded apex (Figures 72-74) (Palearctic) Nemurella
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

13.(11) Tenth tergum enlarged at lateral proximal corners into long sclerotized projections

(Figure 113) (Nearctic) Shipsa

Tenth tergum with only stout lateral produced areas (Figure 97) (Holarctic) Podmosta

14. (10) Epiproct composed almost completely of ventral sclerite; dorsal sclerite reduced to small

projections, connected at base (Figures 100-102) (Nearctic) Prostoia

Epiproct composed of both dorsal and ventral sclerites, with dorsal sclerite slightly

reduced in size (Figures 53, 85, 125) 15

15.(14) Paraprocts with long thin inner and mostly membranous outer lobe (Figures 55, 127);

terminal costal crossvein of forewing joining R (Figure 154) 16

Paraprocts with single large lightly sclerotized lobe (Figure 87); terminal costal crossvein of forewing joining Sc (Figure 153) (Nearctic) Paranemoura

16.(15) Submental gills present and highly branched (Figure 146); vesicle present on ninth

sternum (Figure 130) (Nearctic) Visoka

Submental gills absent; vesicle absent from ninth sternum (Figure 58) (Nearctic)

Lednia

FEMALES

1. Cervical gills highly branched (Figures 140, 141), outer gills forked (Figure 143) or

outer gills short and stubby with inner gills absent (Figure 142); submental gills

absent; seventh sternum usually poorly developed; eighth sternum well developed,

sclerotized and often notched (Figures 11, 19, 27, 35, 43) (Amphinemurinae, new

subfamily) 2

Cervical gills usually simple or absent (Figures 144, 145, 147) (if branched, seventh

sternum well developed and completely covering poorly developed eighth sternum)

or submental gills present (Figure 146); seventh sternum usually more well developed

than eighth sternum (Figures 51, 67, 75, 83, 91, 123, 131, 139) or neither segment

well developed (Figures 59, 99, 107, 115) (Nemourinae) 6

2. (1) Cervical gills highly branched both outside and inside of lateral cervical sclerites; eighth

sternum without large median lobe, with sclerotized vaginal lobes or sclerotization

restricted to margins of narrow median notch (Figures 11, 27) 3

Cervical gills simple inside and forked outside of lateral cervical sclerites or simple outside and absent inside of sclerites; eighth sternum well developed into large median

lobe or subgenital plate, sclerotized vaginal lobes present (Figures 19, 35, 43) 4

3.(2) Gill branches not all extending to gill base (Figure 141); eighth sternum sclerotized

only along median notch (Figure 27) (Nearctic) Malenka

Gill branches all extending to gill base (Figure 140); eighth sternum with sclerotized

vaginal lobes (Figure 11) (Holarctic, Oriental) Atnphinemura

4.(2) Cervical gills simple inside of lateral cervical sclerites and forked outside (Figure 143)

(Palearctic) Protonemura

Cervical gills absent inside of lateral cervical sclerites and reduced to short stubby projections outside (Figure 142) 5

5.(4) Subgenital plate large and extending onto ninth sternum, usually darkly sclerotized

(Figure 19) (Oriental) Indoncmoura, new genus

Subgenital plate small and not extending onto ninth sternum, usually only lightly

sclerotized (Figure 35) (Oriental, Palearctic) Mesonemoura, new genus

6. (1) Seventh sternum enlarged into pregenital plate which covers poorly developed eighth

sternum (Figure 51); with cervical gills or veins Ax and A2 joined in forewings near

margin (Figure 152) or cerci enlarged, lightly sclerotized and truncate at apex (Figure

67); submental gills absent 7

Seventh sternum enlarged or reduced; without cervical gills, without veins At and A2

joined in forewings, without cerci elongate and sclerotized (except in Nemurella

which has large sclerotized patches on eighth sternum) or with submental gills

present 10

7. (6) Cervical gills present 8

Cervical gills absent 9

8.(7) Cervical gills present both inside and outside of lateral cervical sclerites (Figure 147)

(Nearctic) Zapada
NUMBER 211

Cervical gills present only on outside of lateral cervical sclerites (Figure 144) (Palearctic, Oriental) llliesonemoura, new genus

9. (7) Veins A, and A2 joined near margin of forewings (Figure 152); cerci small, membranous

and round at apex (Figure 120) (Nearctic) Soyedina

Veins Aj and A2 not joined in forewings (Figure 154); cerci enlarged, lightly sclerotized

and truncate at apex (Figure 67) (Holarctic, Oriental) Nemoura

10.(6) Submental gills present (Figure 146) (Nearctic) Visoka

Submental gills absent 11

11.(10) Terminal costal crossvein of forewings joining Sc (Figure 153) or with seventh sternum

not modified but eighth sternum developed into a distinctive median platelike patch

bordered by lateral sclerotized patches (Figure 59) 12

Terminal costal crossvein joining R (Figure 149); with both seventh and eighth sterna

enlarged and modified (Figure 75) or both simple and only slightly modified (Figures

99, 107) 13

12.(11) Terminal costal crossveins joining Sc (Figure 153); seventh sternum enlarged and eighth

sternum small and unmodified (Figure 91) (Nearctic) Paranemoura

Terminal costal crossveins joining R (Figure 154); seventh sternum not enlarged but

sternum modified into median platelike and lateral sclerotized patches (Figure 59)

(Nearctic) .....Lednia

13. (10) Eighth sternum enlarged and modified; seventh sternum usually enlarged and modified

(Figures 75, 83) 14

Eighth sternum not enlarged but sometimes with distinct sclerotized pattern or small

median lobe; seventh sternum sometimes slightly produced (Figures 99, 107, 115) ... 15

14.(13) Seventh sternum enlarged into pregenital plate which extends over most of eighth;

eighth sternum with enlarged, sclerotized vaginal lobes (Figure 75) (Palearctic)

Nemurella

Seventh sternum sometimes enlarged and formed into median nipplelike projection;

eighth sternum enlarged and modified into subgenital plate (Figure 83) (Nearctic) ....

Ostrocerca

15.(13) Eighth sternum with small median subgenital plate (Figure 115) (Nearctic) Shipsa

Eighth sternum sclerotized medially but not formed as-subgenital plate (Figures 99,

107) 16

16.(15) Eighth sternum with distinct median sclerotized pattern (Figure 99); forewings mostly

clear with small dark area near veins at cord (Holarctic) Podmosta

Eighth sternum with indistinct, median sclerotized area near posterior margin (Figure

107); forewings mostly fumose with narrow light band running across outer field

(Figure 148) (Nearctic) ProOoia

MATURE NYMPHS

{Lednia unknown)

1. Cervical gills present (Figures 140-143); submental gills absent; whorls of spines absent

from forelegs (Figures 158-161) (Amphinemurinae, new subfamily) 2

Cervical gills usually absent (Figure 145), if present whorls of spines present on forelegs

(Figures 169, 172) or submental gills present (Figure 146) (Nemourinae) 5

2. (1) Cervical gills as two highly branched gills on each side of midline, one inside and one

outside of lateral cervical sclerites (Figures 140, 141) S

Cervical gills simple, reduced or partially absent, gills outside of lateral cervical sclerites

may be forked (Figures 142, 143) 4

3.(2) Gill branches not all extending to gill base (Figure 141) (Nearctic) Malenka

Gill branches all extending to gill base (Figure 140) (Holarctic, Oriental) Amphmemura

4. (3) Cervical gills simple inside of lateral cervical sclerites and forked outside (Figure 143)

(Palearctic) Protonemura

Cervical gills absent inside of lateral cervical sclerites and reduced to short stubby

projections outside (Figure 142) (Oriental, Palearctic)

Mesonemoura, new genus, and (Oriental) Indonemoura, new genus

5. (1) Pronotum with a definite fringe of spines on lateral margins, spines in single row and

of equal length or with gradual shift in length toward corners (Figure 1); mesonotum

and metanotum with fringes of spines along lateral margins (Figure 1) 6
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pronotum without a fringe of spines, or if present not arranged in a single row or of

unequal lengths (Figure 146); mesonotum and metanotum without fringes of spines

or with very sparse fringes mostly visible near posterolateral margins 9

6.(5) Cervical gills present (Figures 144, 147); whorls of spines present on forelegs (Figures

169, 172) 7

Cervical gills absent (Figure 145); whorls of spines usually absent from forelegs (Figures

170, 171) 8

7. (6) Two cervical gills present on each side of midline one inside and one outside of lateral

cervical sclerites (Figure 147); whorls of femoral spines present on all leg pairs (Figure 1) (Nearctic) Zapada

One cervical gill present on each side of midline, located outside of lateral cervical

sclerites (Figure 144); whorls of femoral spines present only on forelegs (Figure 169)

(Palearctic, Oriental) Illiesonemoura

8. (6) Pronotum rounded at corners and without a definite notch on lateral margins (Holarctic, Oriental) Nemoura

Pronotum angular at corners and with a definite notch on lateral margins (Nearctic) ....

Soyedina

9. (5) Foreleg with numerous large spines on femur and tibia, outer margin of tibia with row

of large spines and often with fringe of long hairs (Figures 162, 163, 164, 166, 167);

submental gills absent 10

Foreleg with few large spines on femur and tibia, outer margin of tibia with spines of

similar length to those on surface and without fringe of long hairs (Figure 165) or

foreleg with many spines and submental gills present (Figure 168) 14

10.(9) Pronotum with irregular fringe of moderate to large spines 11

Pronotum without fringe of moderate to large spines 12

11.(10) Pronotum wider than long, with irregular fringe of large spines; tibiae with only scattered large spines along outer margins (Figure 167) (Palearctic) Nemurella

Pronotum as long as wide, with irregular fringe of moderate spines; tibiae with two

definite rows of large spines along outer margins (Figure 162) (Nearctic) ... Ostrocerca

12.(10) Foretibiae with fringe of long hairs along outer margin (Figures 164, 166) 13

Foretibiae without fringe of long hairs along outer margin but sometimes with a few

scattered long hairs (Figure 163) (Holarctic) Podmosta

13.(12) Tibiae with two rows of stout spines along outer margins (Figure 164) (Nearctic)

Prostoia

Tibiae without rows of stout spines along outer margins (Figure 166) (Nearctic) ... Shipsa

14. (9) Submental gills present (Figure 146); foreleg with numerous large spines and long hairs

(Figure 168) (Nearctic) Visoka

Submental gills absent; foreleg with few large spines or long hairs (Figure 165) (Nearctic) Paranemoura

AMPHINEMURINAE, new subfamily

Amphinemura is the type-genus for this subfamily, which also includes the genera Indonemoura, Malenka, Mesonemoura, and Protonemura.

The subfamily is recognized by the following characters:

1. Paraprocts divided into three lobes.

2. Middle and outer lobes of paraprocts usually

bearing spines or prongs.

S. Epiproct elongate, generally laterally compressed and with ventral sclerite forming a

rounded or triangular keel.

4. Cervical gills present.

5. Subgenital plate well developed in females

but pregenital plate usually poorly developed.

The Amphinemurinae are well represented in

the Palearctic and Nearctic regions and also extend

south into the Oriental region. Amphinemura is

found in the Nearctic, Palearctic, and Oriental regions (Figure 173); Mesonemoura occurs in the

Oriental and Palearctic regions (Figure 175); Protonemura (Figure 176) and Indonemoura (Figure

174) are found only in the Palearctic and Oriental

regions respectively; and Malenka is restricted to

the western part of the Nearctic region (Figure

174).
NUMBER 211 11

Amphinemura Ris

FICURES 4-11, 140, 155, 159, 173

Nemura (Amphinemura) Ris, 1902:384. [Type-species:

Nemoura cinerea Olivier = Amphinemura sulcicollis

Stephens.]

Amphinemura.—Claassen, 1940:47.

ADULT.—Length: small to medium (5-10 mm).

Wings (Figure 155): macropterous, venation typical for family, clear or fumose to dark and mottled,

veins dark and distinct, sometimes with light spots

in cells or light bands on dark background. Gills

(Figure 140): 2 branched cervical gills on each

side of midline, 1 arising inside and 1 outside of

lateral cervical sclerites, minimum branches 5,

maximum observed 16, all branches extending to

gill base.

MALE TERMINALIA.—Hypoproct (Figure 10):

broad at base, tapering to narrow apex, extending

distally over inner lobes of paraprocts, tip extending to base of epiproct and sometimes recurved dorsally; vesicle present. Paraprocts (Figure

7): divided into 3 lobes; inner lobes lightly sclerotized, naked, short and closely applied at midline,

often completely hidden by hypoproct; median

lobes mostly sclerotized but with membranous

inner portion, bearing few to many hairs or spines,

lobes large and elongate, lying alongside hypoproct

and recurved dorsally parallel to epiproct; outer

lobes mostly sclerotized, sometimes with small

membranous portion, shape variable, often recurved dorsally alongside cerci, usually with a few

hairs or spines. Cerci (Figures 8-10): membranous,

short and unmodified. Epiproct (Figures 4-6):

short and usually fairly broad in dorsal aspect and

narrow with ventral projections in lateral aspect,

completely recurved, bilaterally symmetrical; dorsal sclerite, large and broad at base of epiproct,

extending dorsolaterally toward apex, darkly sclerotized lateral arms on lateral margins, variously

modified, apical portion large and extending laterally over ventral sclerite, bearing small spines or

sclerotized scales; basal sclerites, as 2 broad triangular patches located at basolateral margins of

epiproct; ventral sclerite, heavily sclerotized, broad

at base and becoming narrower toward apex, forming median keel-shaped ridge, apical portion inserted between folds of dorsal sclerite and variously modified, often extending to dorsal margin,

sclerite bearing few to many spines which are often

very large. Tenth tergum (Figure 8) all or mostly

sclerotized, forming a large flat area anterior to

base of epiproct, usually bare but sometimes with

spines or projections. Ninth tergum sclerotized

and usually produced along distal margin, produced portion bearing spines or hairs, produced

area sometimes large and bizarre and can even be

asymmetrical.

FEMALE TERMINALIA (Figure 11).—Seventh sternum produced and extended at distal margin

forming small pregenital plate, covering part of

sternum eight, produced area lightly sclerotized.

Eighth sternum forming subgenital plate, size of

plate variable but usually quite small and bifid,

divided at genital opening into symmetrical sclerotized vaginal lobes which are often quite distinctive, lobes usually restricted to eighth sternum

but occasionally extend over anterior margin of

ninth sternum.

NYMPH.—Gills (Figure 140): 2 branched cervical gills on each side of midline, 1 arising inside

and 1 outside of lateral cervical sclerites, minimum

branches 5, maximum observed 16, all branches

originating at same place near base. Forelegs

(Figure 159): femur and tibia of approximately

equal length, whole leg covered with fine short

hairs; femur with row of long spines along posterior margin and often with a few long hairs

intermingled, dorsal surface with few to numerous

long spines on distal half; tibia with rows of

medium-sized spines along anterior and posterior

surfaces, few to numerous long hairs intermingled

with spines on posterior margin. Pronotum with

fringe of spines along lateral margins. Abdominal

terga with fringes of spines along distal margins.

Cerci with whorls of large spines at distal margins

of segments, segments covered randomly with

numerous small intermediate spines.

DISCUSSION.—Amphinemura is the most widespread genus in the family Nemouridae, ranging

from the Arctic in the north to North Africa, the

Sunda Archipelago and southern Mexico respectivly in the south. It contains 90 known species

and surely many more that are unstudied. Members of the genus are easily recognized in both the

nymphal and adult stages by their highly branched

cervical gills (Figure 140).

It is found in a wide variety of freshwater habitats but is most common in cold, clear mountain

streams that run throughout the year.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DISTRIBUTION AND SPECIES LIST.—Holarctic and

Oriental (Europe, Asia, North Africa, North

America, Orient).

amatulai Aubert 1967, Assam

apache Baumann & Gaufin 1972, Arizona

arcadia (Aubert) 1956a, Greece

banksi Baumann & Gaufin 1972, Rocky Mountains

bilolai Aubert 1967, Assam

bomdilai Aubert 1967, Assam

borealis (Morton) 1894, Europe, northern Asia, Mongolia

chiffensis (Aubert) 1956c, Morocco

•chui (Wu) 1935, China

•claassenia (Wu) 1935, China

•claviloba (Wu) 1973, n. comb., China

coreana Zwick 1973c, Korea

"cornuloba (Wu) 1973, n. comb., China

* cryptoctrcia (Wu) 1938, n. comb., China

*curvispina (Wu) 1973, n. comb., China

*decemseta Okamoto 1922, Japan

delosa (Ricker) 1952, eastern United States

•dentiloba (Wu) 1973, n. comb., China

dichotoma (Kawai) 1954, Japan

•falciloba (Wu) 1973, n. comb., China

•fililoba (Wu) 1973, n. comb., China

•flavicollis Klapalek 1912b, Taiwan

*flavostigma Okamoto 1922, Japan

•fleurdelia (Wu) 1949, n. comb., China

•forcipiloba (Wu) 1962, n. comb., China

•furcospinata (Wu) 1949, n. comb., China

*furcostyla (Wu) 1973, n. comb., China

gressitti Kawai 1969, Viet Nam

gritsayae Zhiltzova 1971, central Asia

handschini (Geijskes) 1937, Java, Sumatra, Viet Nam

*hastata (Wu) 1973, n. comb., China

•kiangsiensis (Wu) 1935, China

*lagnucula Harper 1975, Nepal

•liccnti (Wu) 1938, China

linda (Ricker) 1952, eastern and northern North America

lithami Aubert 1967, Assam

*longispina (Okamoto) 1922, Japan

luteipes Kimmins 1947, Himalaya, Assam

"macrolamellata (Wu) 1935, n. comb., China

manipurensis Aubert 1967, Assam

*maoi (Wu) 1938, China

maracandica (McLachlan) 1875, central Asia

megaloba (Kawai) 1960, Japan

mexicana Baumann 1972, Mexico

minuta Kawai 1969, Borneo

mirabilis (Martynov) 1928, western Asi.i

mockfordi (Ricker) 1952, Tennessee

mogollonica Baumann and Gaufin 1972, southwestern United

States

•mokanshenensis (Wu) 1938, China

monotuberculata (Kawai) 1956, Japan

moshingi Aubert 1967, Assam

*multispina (Wu) 1973, n. comb., China

"nepalemis Harper 1975, Nepal

nigritta (Provancher) 1876, eastern North America

•nigritula Navas 1932, Formosa

nongrimi Aubert 1967, Assam

nowegica Tobias 1973, Norway

nubila Kimmins 1950a, southern India

okinawaensis Kawai 1968a, Okinawa

paraluteipes Aubert 1967, Assam, Nepal

*pentagona (Okamoto) 1922, Japan

•pieli (Wu) 1938, n. comb., China

pseudoluteipes Aubert 1967, Assam

puebla Baumann 1972, Mexico

rahungi Aubert 1967, Assam

reinerti Baumann 1976, Mexico

renata Kimmins 1950b, Assam

*rostroloba Wu 1962, n. comb., China

*sagittata (Okamoto) 1922, Japan

schmidi (Aubert) 1959a, Pakistan

•sinensis (Wu) 1926, China

sperchiana Berthelemy 1971, Greece

'spinata (Wu) 1949, n. comb., China

standjussi (Ris) 1902, Europe

steinmanni Zwick 1973c, Korea

sulcicollis (Stephens) 1835, Europe

talungdzongi Aubert 1967, Assam

tetraspinosa (Kawai) 1960, Japan

thienemanni (Geijskes) 1952, Java

tragula Kimmins 1950a, Turkestan, Tadzhikistan

•trassaerti (Wu) 1938, China

trialetica Zhiltzova 1957, Caucasus, Asiatic Turkey

triangularis Ris 1902, Europe

tricantha (Jewett) 1958, Himalaya

•triramia (Wu) 1962, n. comb., China

*unihamata (Wu) 1973, n. comb., China

varshava (Ricker) 1952, eastern United States

venusta (Banks) 1911, Arizona, Mexico

verrucosa Zwick 1973c, Korea

wui (Claassen) 1936, eastern North America

zimmermanni Joost 1970b, Tien Shan

*zonata Okamoto 1922, Japan

Indonemoura, new genus

FIGURES 12-19, 174

TYPE-SPECIES.—Protonemura indica Kimmins,

1947:727, herein designated.

ADULT.—Length: medium to large (7-15 mm).

Wings: macropterous, venation typical for family,

clear or fumose, veins dark and distinct. Gills: single short oval gills on each side of midline, located

on outside of lateral cervical sclerites.

MALE TERMINALIA.—Hypoproct (Figure 18):

broad at base, tapering to narrow apex, extending

to base of epiproct, often covering inner lobes-of

paraprocts; vesicle present. Paraprocts (Figure

15): divided into 3 lobes; inner lobes small and

lightly sclerotized, often closely affixed to large
NUMBER 211 13

median lobes; median lobes mostly sclerotized,

broad at base, basal half lightly sclerotized and

covered with hairs, apical half darkly sclerotized,

long narrow and formed into one or more long

prongs or projections, usually with a small inner

membranous portion; outer lobes well developed,

darkly sclerotized, long narrow and almost completely encircling cerci, tip often terminating in

one or more sharp prongs. Cerci (Figures 16-18):

membranous, very long and thin, extending beyond tip of abdomen, covered with thin hairs.

Epiproct (Figures 12-14): usually long and narrow

in dorsal aspect, lateral aspect narrow near base

but expanded near apex, completely recurved, bilaterally symmetrical; dorsal sclerite, large and

fairly broad at base of epiproct, extending dorsolaterally toward apex, sometimes enlarged at bend,

apical portion usually greatly enlarged, especially

near tip; basal sclerites as 2 broad triangular

patches at basolateral margins of epiproct; ventral

sclerite, heavily sclerotized, broad at base and becoming narrower toward apex, forming median

keel-shaped ridge, usually with few to many spines

in small rounded area near apex, tip inserted in

membranous folds of dorsal sclerite and rarely

with tubelike structure extending from tip. Tenth

tergum (Figure 16) all or mostly sclerotized but

with large membranous area under apex of epiproct, usually forming a flat or concave plate but

sometimes with 1 or 2 large upwardly directed protuberances. Ninth tergum sclerotized and produced

along distal margin, produced portion bearing

spines or hairs, and sometimes large and bizarre

in shape.

FEMALE TERMINALLY (Figure 19).—Seventh sternum somewhat produced at distal margin, extending slightly onto eighth sternum medially, produced area lightly sclerotized. Eighth sternum

forming subgenital plate, median area enlarged

and heavily sclerotized, forming distinctive plate

which covers genital opening and extends over

part of vaginal lobes, shape of plate highly angular and darkly sclerotized. Cerci long, thin, membranous, and unmodified.

NYMPH.—Unknown (included in nymphal key

using adult gill characters).

DISCUSSION.—Indonemoura includes the Indica

group of Aubert (1967) and two large species from

the Sunda Archipelago. It probably occurs in other

parts of Asia, especially Burma and Thailand. It

is similar to Mesonemoura, regarding the type of

cervical gills present, but can be separated in the

male by the large, highly developed and ornamented paraprocts (Figure 15) and in the female

by the large darkly sclerotized subgenital plate

(Figure 19) which nearly covers the very small

vaginal lobes.

The generic name Indonemoura is taken from

the stem "Indo-" referring to India or the East

Indies, where the genus occurs.

DISTRIBUTION AND SPECIES LIST.—Oriental (south

central Asia). The following species are assigned to

the genus Indonemoura:

assami (Aubert) 1967, n. comb.

dirangdzongi (Aubert) 1967, n. comb., Assam

gigaoni (Aubert) 1967, n. comb., Assam

indica (Kimmins) 1947, n. comb., Assam, India

jacobsoni (Klapalek) 1912a, n. comb., Bali, Java, Malaya,

Sumatra

javanica (Banks) 1920, n. comb., Java

kamengi (Aubert) 1967, n. comb., Assam

maclachlani (Kimmins) 1950b, n. comb., Assam

manipuri (Aubert) 1967, n. comb., Assam

nahkui (Aubert) 1967, n. comb., Assam

nyukmadongi (Aubert) 1967, n. comb., Assam

quadridentata (Kimmins) 1950b, n. comb., Assam

sangtii (Aubert) 1967, new comb., Assam

shergaoni (Aubert) 1967, n. comb., Assam

Malenka Ricker

FIGURES 20-27, 141, 151, 158, 174

Nemoura (Malenka) Ricker, 1952:29. [Type-species: Nemoura

cornuta Claasen.]

Malenka.—lilies, 1966:190.

ADULT.—Length: small to medium (5-10 mm).

Wings (Figure 151): macropterous, venation typical for family, clear or slightly fumose at veins

which are dark and distinct. Gills (Figure 141):

2 branched cervical gills on each side of midline,

1 arising inside and 1 outside of lateral cervical

sclerites, minimum branches 5, maximum observed

8, some branches arising above gill base.

MALE TERMINALIA.—Hypoproct (Figure 26):

broad at base, tapering to narrow apex, extending

distally over inner lobes of paraprocts, tip extending nearly to base of epiproct; vesicle present. Paraprocts (Figure 23): divided into 3 lobes; inner

lobes lightly sclerotized, naked, short and closely

applied to midline, often completely hidden by

hypoproct; median lobes lightly sclerotized api-
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

cally, naked or with small hairs, base broad and

membranous, apical half tapering to a narrow tip

that terminates in a sclerotized point or prongs,

lying alongside hypoproct and extending to or

beyond tips of cerci; outer lobes sclerotized, shape

variable, recurved alongside cerci, often curving in

front of median lobes dorsally, nearly fused to

median lobes throughout length, naked except in

M. marionae (Hitchcock) which has numerous

short dark spines at apex. Cerci (Figures 24-26:

with distinctive mesobasal lobes which may be

membranous or heavily sclerotized, with single or

bifurcate apex, main lobe of cerci membranous

and unmodified. Epiproct (Figures 20-22): usually

short and broad in dorsal aspect and narrow in

lateral aspect, completely recurved, lightly sclerotized, bilaterally symmetrical; dorsal sclerite, fairly

large and broad at base of epiproct, extending

dorsolaterally toward apex, apical portion modified in structure, usually enlarged and extending

over ventral sclerite laterally, covered with many

tiny spines or scales; basal sderites, as 2 broad triangular plates located at basolateral margins of

epiproct; ventral sclerite, heavily sclerotized,

broadest at base, narrower toward apex, forming

median keel-shaped ridge which is naked of spines,

apical portion inverted between lateral folds of

dorsal sclerite, extending to apex where it forms a

small rounded tip. Tenth tergum (Figure 24) all

or mostly sclerotized, forming a large flat sclerotized area anterior to base of epiproct, mostly bare

but often with a few spines along the anterior median margin. Ninth tergum sclerotized and usually

slightly produced along distal margin, produced

portion bearing spines or hairs.

FEMALE TERMINALIA (Figure 27).—Seventh sternum produced at mesoposterior margin, lightly

sclerotized, bearing a nipplelike projection best

seen in lateral view. Eighth sternum forming subgenital plate which is divided at genital opening

by a median notch into 2 equal sternites or vaginal lobes, notch often lined by heavy sclerotization, lobes often swollen or produced but do not

overlap ninth sternum.

NYMPH.—Gills (Figure 141): 2 branched cervical

gills on each side of midline, 1 arising inside and 1

outside of lateral cervical sclerites, minimum

branches 5, maximum observed 8, some branches

originating some distance from base toward apex.

Forelegs (Figure 158): femur and tibia of approximately equal length, most of leg covered with

small fine hairs; femur with long hairs along posterior margin and bunched near distal end of dorsal surface; tibia with rows of long spines along

posterior margin, with many long hairs intermingled with spines, anterior margin with row of

short spines. Pronotum with fringe of spines along

lateral margins. Abdominal terga with spines along

distal margins. Cerci with whorls of large spines at

distal margins of segments, intermediate areas with

randomly arranged small spines.

DISCUSSION.—Malenka is the sister genus of

Amphinemura and is restricted to western North

America. It is found in small streams and creeks

and is also common in small springs. It usually

emerges in the late summer or fall, while most

other nemourid species emerge in the early spring.

It can be distinguished in the nymphal stage by

its highly branched cervical gills which have some

branches that do not extend to the gill base (Figure 141). The male can be recognized by the

presence of mesobasal lobes on the cerci (Figure

24), and the females have a nipplelike ventral

projection on the seventh sternum and a distinctive notch on the median posterior margin of the

eighth sternum (Figure 27).

DISTRIBUTION AND SPECIES LIST.—Nearctic (western North America). The genus Malenka contains

the following known species:

bifurcata (Claassen) 1923, northwestern United States

biloba (Claassen) 1923, California

californica (Claassen) 1923, western North America

coloradensis (Banks) 1897, Rocky Mountains

cornuta (Claassen) 1923, northwestern North America

depressa (Banks) 1898, northwestern United States

flexura (Claassen) 1923, western North America

marionae (Hitchcock) 1958, California

perplexa (Frison) 1936, northwestern United States

tina (Ricker) 1952, western United States

wenatchee (Ricker) 1965, Washington

Mesonemoura, new genus

FIGURES 28-35, 142, 161, 175

TYPE-SPECIES.—Nemoura vaillanti Navas 1922:9,

herein designated.

ADULT.—Length: medium to large (7-12 mm).

Wings: macropterous, venation typical for family,

clear or fumose, veins dark and distinct. Gills (Figure 142): single short oval gills on each side of
NUMBER 211 15

midline, located outside of lateral cervical sclerites.

MALE TERMINALIA.—Hypoproct (Figure 34):

broad at base, tapering to narrow apex, extending

to and covering part of inner lobes of paraprocts;

vesicle present. Paraprocts (Figure 31): divided

into 3 lobes; inner lobes lightly sclerotized but

sometimes with darkly sclerotized projections, lobes

well developed and fairly broad; median lobes

partially sclerotized, with large membranous inner

and apical portions, sclerotized area often bearing

spines, membranous area with many hairs, lobes

quite large and recurved toward base of epiproct;

outer lobes mostly sclerotized, long thin and curved

around cerci, often bearing a few small spines.

Cerci (Figures 32-34): membranous, long, narrow

and thin near apex, usually longer than paraprocts.

Epiproct (Figures 28—30): usually short and broad

in dorsal aspect and narrow in lateral aspect, completely recurved, mostly sclerotized, usually bilaterally symmetrical except for ventral apical tubelike structure which varies in shape from long

and thin to short and stout; dorsal sclerite, large

and broad at base of epiproct, extending dorsolaterally toward apex, apical portion enlarged and

extending over onto ventral sclerite; basal sclerites

as 2 broad triangular patches at basolateral margins of epiproct; ventral sclerite heavily sclerotized,

broad at base and becoming narrower toward apex,

forming median keel-shaped ridge, tubelike sclerotized structure extending from tip, apical portion

inserted between folds of dorsal sclerite, ventral

sclerite usually bearing few to many spines along

keel-shaped ridge. Tenth tergum (Figure 32) all

or mostly sclerotized forming a large flat area anterior to base of epiproct, usually bare but sometimes with a few spines or hairs. Ninth tergum

sclerotized and produced along distal margin,

bearing spines or hairs, produced area sometimes

large and bizarre and can even be asymmetrical.

FEMALE TERMINALIA (Figure 35).—Seventh sternum somewhat produced at distal margin, extending slightly onto eighth sternum at midline, producd area lightly sclerotized. Eighth sternum

forming subgenital plate, median area enlarged

and sclerotized, forming plate that covers most of

segment and genital opening, lateral posterior

margins of segment enlarged and lightly sclerotized, forming small vaginal lobes.

NYMPH.—Gills (Figure 142): single short oval

gills on each side of midline, arising outside of

lateral cervical sclerites, gills shaped like small

rounded knob slightly longer than wide. Forelegs

(Figure 161): femur and tibia of approximately

equal length, whole leg covered with fine short

hairs; femur with a row of very small spines along

posterior margin, with a few large long spines on

basal half on posterior margin and dorsal surface;

tibia with row of small spines near anterior margin, posterior margins with fringes of long hairs.

Pronotum with fringe of spines along lateral margins. Abdominal terga with fringes of spines along

distal margins, often also with two longitudinal

rows of single large spines. Cerci with whorls of

large spines at distal margins of segments, areas

between joints covered randomly with numerous

intermediate spines.

DISCUSSION.—Mesonemoura contains species that

were once assigned to Amphinemura, Protonemura,

and Nemoura. It is similar to Amphinemura in

many respects but is most similar to Indonemoura.

It differs from Amphinemura in only having

single, simple cervical gills on the outside of the

lateral cervical sclerites (Figure 142). The nymph

labeled Nemoura sp. (no. 102) in Kawai (1963)

seems to belong in Mesonemoura. It can be separated from Indonemoura by the smaller less

developed paraprocts (Figure 31) and by the

presence of a long thin epiproctal flagellum (Figures 28-30). Females of Mesonemoura have a

small median subgenital plate and two small vaginal lobes (Figure 35), while Indonemoura females

have a single large, darkly sclerotized subgenital

plate, which nearly obscures the vaginal lobes

(Figure 19).

Most of the species presently assigned to this

genus occur in the high mountains of Central Asia.

DISTRIBUTION AND SPECIES LIST.—Oriental and

Palearctic (south central Eurasia).

brachyfiligera (Aubert) 1967, n. comb., Assam

falcata (Kimmins) 1950b, n. comb., Assam

filigera (Kimmins) 1947, n. comb., Himalaya, Nepal

•flagellata (Wu) 1935, n. comb., China

*hamistyla (Wu) 1962, n. comb., China

*longicercia (Okamoto) 1922, n. comb., Japan

metafiligera (Aubert) 1967, n. comb., Assam

mishmica (Kimmins) 1950b, n. comb., Assam

*multispira (Wu) 1973, n. comb., China

parafiligera (Aubert) 1967, n. comb., Assam

paraproctalis (Aubert) 1967, n. comb., Assam

pseudofiligera (Aubert) 1967, n. comb., Assam

skardui (Aubert) 1959a, n. comb., Pakistan, Afghanistan
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

•spiroflagellata (Wu) 1973, n. comb., China

tianschanica (Zhiltzova) 1971, n. comb., central Asia

vaillanti (Navas) 1922, n. comb., China, Afghanistan, Pakistan

Protonemura Kempny

FIGURES 36-43, 143, 157, 160, 176

Nemura (Protonemura) Kempny, 1898:51. [Type-species:

moura meyeri Pictet = Protonemura meyeri (Pictet).]

Protonemura.—Claassen, 1940:68.

ADULT.—Length: medium to large (7-15

Wings (Figure 157): macropterous, venation typical for family, clear, fumose or mottled, veins dark

and distinct. Gills (Figure (143): 2 cervical gills,

1 on each side of lateral cervical sclerites, outer

gills with single branch at base forming 2 l°ng

thin nearly equal parts, inner gills long, thin

 and

single.

MALE TERMINALLY.—Hypoproct (Figure 42):

broad at base, tapering to narrow apex, extending

distally over inner lobes of paraprocts, tip extending to base of epiproct; vesicle present. Paraprocts

(Figure 39): divided into 3 lobes; inner lobes

lightly sclerotized, naked, short, and closely

applied to midline, often completely hidden by

hypoproct; median lobes heavily sclerotized but

with large membranous apical portion, sclerotized

portion broad at base, tapering to apex, which is

often branched, inner branch darkly sclerotized

and bearing one or more prongs or spines near

apex, membranous portion bulbous and covered

with small hairs; outer lobes mostly sclerotized,

elongate and recurved dorsally alongside cerci,

usually with one or more spines at apex. Cerci

(Figures 40-42: membranous, short and Unmodified. Epiproct (Figures 36-38): usually long and

thin in dorsal aspect and thin but enlarged apically in lateral aspect, completely recurved, mostly

sclerotized, bilaterally symmetrical, occasionally

with an apical tubelike projection arising froitf the

ventral sclerite; dorsal sclerite, large and broad at

base of epiproct, extending dorsolaterally to apex,

apical portion enlarged covering dorsolateral margins, heavily sclerotized lateral arms variously

modified, apex of sclerite round, covered with

small spines or scales; basal sclerites shaped like

broadly rounded triangles, located at basolateral

margins of epiproct; ventral sclerite, broadest at

base, becoming narrower toward apex, apical Portion inserted between folds of dorsal sclerite, often

extending to dorsal margin, ventral margin of sclerite bearing spines along complete length or in a

tuft near apex. Tenth tergum (Figure 40) all or

mostly sclerotized, forming a large flat area anterior to base of epiproct, usually bearing small

spines or hairs. Ninth tergum sclerotized and usually produced, especially along distal margin,

bearing spines and hairs and often distinctly

shaped. Eighth tergum usually enlarged, bearing

spines and hairs. Seventh tergum usually normal

and bare but sometimes with a few spines and

hairs.

FEIVIALE TERMINALIA (Figure 43).—Seventh sternum slightly produced at distal margin, seldom extending over segment eight, produced area lightly

sclerotized. Eighth sternum forming subgenital

plate, central area enlarged and sclerotized forming plate that covers most of segment and genital

opening, lateral posterior margins of segment produced and sclerotized, forming vaginal lobes.

NYMPH.—Gills (Figure 143): 2 cervical gills on

each side of midline, 1 arising inside and 1 outside

of lateral cervical sclerites, outer gills forked at base

into 2 nearly equal branches, inner gills single,

gills long thin and often constricted one or more

times. Forelegs (Figure 160): femur and tibia of

approximately equal length, femur with spines

along posterior margin, extending to dorsal surface on distal half; tibia with rows of spines along

posterior margins, intermingled with scattered

small spines and hairs, anterior half covered with

random spines that become most dense at distal

end covering entire surface. Pronotum with fringe

of spines along lateral margins. Abdominal terga

with fringes of spines along lateral margins, often

with 2 longitudinal rows of single large spines.

Cerci with whorls of large spines at distal margins

of segments, areas between joints covered randomly

with small intermediate spines.

DISCUSSION.—Protonemura is very common in

southern Europe and seems to fill the niche occupied by Amphinemura in North America and most

of Asia. It is a large genus containing over 80

species.

The distinctive features of the genus are the

cervical gills and the well developed female subgenital plate. The outer cervical gills have a single

deep fork and the inner gills are long and simple

(Figure 143). The female eighth sternum is well
NUMBER 211 17

developed into a large median subgenital plate

and large sclerotized vaginal lobes (Figure 43).

DISTRIBUTION.—Palearctic (Europe, Asia, North

Africa).

abchasica Zhiltzova 1964, Caucasus

aestiva Kis 1965a, Balkans, Carpathians

albanica Rauser 1963b, Albania

alcazaba (Aubert) 1954b, Spain

algirica (Aubert) 1956c, Algeria, Morocco

algovia Mendl 1968a, Alps

alticola Zhiltzova 1958, Caucasus

angelieri Berthelemy 1963, Pyrenees

auberti lilies 1954, central Europe

ausonia (Consiglio) 1955, Italy

autumnalis Rauser 1956, Balkans, Carpathians

bacurianica Zhiltzova 1957, Caucasus, Asia Minor

beatensis (Despax) 1929, Pyrenees, Italy

beautnonti (Aubert) 1956a, Balkans

besucheti Zwick 1971, Turkey

bifida Martynov 1928, Caucasus, Iran

bipartita Consiglio 1962, Italy

bithynica Aubert 1964b, Turkey

*brevilobata Klapalek 1912b, Taiwan

brevistyla (Ris) 1902, Alps, Carpathians

bucolica (Consiglio) 1957b, Corsica

capitata Martynov 1928, Caucasus, Armenia

caprai (Aubert) 1954c, Italy

consiglioi (Aubert) 1953b, Italy

corsicana (Morton) 1930, Corsica

costai (Aubert) 1953a, Italy

dilatata Martynov 1928, Caucasus

elbourzi Aubert 1964a, Iran

eumontana Zhiltzova 1957, Caucasus, Asia Minor

gladifera Balinsky 1950, Caucasus

hassankifi Aubert 1964a, Iran

hiberiaca Aubert 1963a, Spain

hirpina (Consiglio) 1958, Italy

hispanica (Aubert) 1956b, Spain

•hotakana Ueno 1931, Japan

hrabei Rauser 1956, central Europe

ichnusae (Consiglio) 1957a, Sardinia

illiesi Kis 1963, Balkans, Carpathians

intricata (Ris) 1902, Europe

italica (Aubert) 1954a, Italy

•jezoensis Okamoto 1922, Japan

lagrecai (Aubert) 1954a, Italy

later alls (Pictet) 1836, Europe

libanica Aubert 1964b, Lebanon

macrura (Aubert) 1953b, Italy

mattheyi (Aubert) 1956a, Greece

meyeri (Pictet) 1841, Europe

microstyla Martynov 1928, Caucasus, Armenia

montana Kimmins 1941, Europe

navacerrada (Aubert) 1954b, Spain

nimborella Mosely 1930, Alps

nimborum (Ris) 1902, central Europe

nitida (Pictet) 1835, Europe

•nohirae Okamoto 1922, Japan

oitica Aubert 1963c, Greece

orcas Martynov 1928, Caucasus

praecox (Morton) 1894, Europe

pseudonimborum Kis 1965, Carpathians

pyrenaica Mosely 1930, Pyrenees

ressli Zwick 1971, Asia Minor

risi (Jacobson & Bianchi) 1902, central Europe

ruffoi Consiglio 1961, Italy

salfii (Aubert) 1954a, Italy

sicula Consiglio 1961, Sicily

spinulata Martynov 1928, Caucasus

strandschaensis Braasch & Joost 1972, Bulgaria

talboti (Navas) 1929, North Africa

taygetiana (Aubert) 1956a, Greece

teberdensis Zhiltzova 1958, Caucasus, Asia Minor

* towadensis Kawai 1954, Japan

•triangulata Martynov 1928, Caucasus

tuberculata (Despax) 1929, Pyrenees

tyrrhena (Festa) 1938, Italy, Morocco

vandeli Berthelemy 1963, Pyrenees

vercingetorix Aubert 1963a, France

vernalis Zhiltzova 1958, Caucasus, Armenia

viridis Balinsky 1950, Caucasus

waliabadi Aubert 1964a, Iran

zernyi Aubert 1964b, Lebanon, Israel

NEMOURINAE Newman, 1853

Nemoura is the type-genus for this large subfamily, which also contains the following genera:

Illiesonemoura, Lednia, Nemurella, Ostrocerca,

Paranemoura, Podmosta, Prostoia, Shipsa, Soyedina, Visoka, and Zapada.

The subfamily can be distinguished by the following characters:

1. Paraprocts with a thin inner lobe and a broad

outer lobe or just a single broad lobe.

2. Paraprocts lacking spines or prongs.

3. Epiproct usually short and dorsoventrally

compressed, with the ventral sclerite enlarged,

especially at base.

4. Cervical gills usually absent.

5. Pregenital plate well developed in females or

neither pregenital plate nor subgenital plate

especially well developed.

Nemourinae is a very diverse grouping that contains one large genus, Nemoura, and many small

genera, four of which are monotypic. Nemoura is

found throughout the Holarctic and also extends

to the Oriental region (Figure 178). Illiesonemoura is Palearctic and Oriental, and Nemurella

is Palearctic (Figures 177, 179). Podmosta extends
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

from the Nearctic into the eastern part of the

Palearctic (Figure 181); Lednia, Ostrocerca, Paranemoura, Prostoia, Shipsa, Soyedina, Visoka, and

Zapada have been found only in the Nearctic

region (Figures 177, 179-186).

This subfamily may prove to be a polyphyletic

lineage, but it is treated in this study as a raono

phyletic lineage based on the shape and structure

of the ventral sclerite of the epiproct.

Illiesonemoura, new genus

FIGURES 44-51, 144, 156, 169, 177

TYPE-SPECIES.—Nemoura punctata Jewett, 1958:

324, herein designated.

ADULT.—Length: small to large (5-12 mm).

Wings (Figure 156): macropterous, venation typical for family, fumose, mottled or with light spots

in cells on dark background, veins dark and distinct. Gills (Figure 144): 1 unbranched cervical

gill on each side of midline, arising on outside of

lateral cervical sclerites, gills triangular-shaped

and of medium length, small gill-like nubs on inside of lateral cervical sclerites.

MALE TERMINALIA.—Hypoproct (Figure 50):

broad at base, tapering to narrow apex, extending

distally over base of paraprocts, often covering inner lobes of paraprocts; vesicle present. Paraprocts

(Figure 47): consisting of 2 lobes; inner lobes

sclerotized, short and narrow, closely applied to

midline and often turned inward, sometimes completely hidden by hypoproct; outer lobes sclerotized ventrally and membranous dorsally, very

large and generally triangular in shape, with inward directed sclerotized projection from outer

margin near cerci. Cerci (Figures 48-50): mostly

membranous, long thin and often bent. Epiproct

(Figures 44-46): short and broad in dorsal aspect,

short and varying from broad to rather thin in lateral aspect, apex rounded, completely recurved,

bilaterally symmetrical; dorsal sclerite large and

broad at base of epiproct, extending dorsolaterally, becoming narrow around lateral knobs and

then very large and triangular, often completely

covering lateral aspects of epiproct and much of

ventral aspect, most darkly sclerotized areas at base

and near posterior end immediately ahead of basal

cushion, anterior area lightly sclerotized; basal

sclerites as 2 broad triangular or rectangular

patches located near basolateral corners of epiproct; ventral sclerite heavily sclerotized, broad at

base and with lateral knobs at basolateral corners,

becoming narrower toward apex, forming parallel

ridges, one on each side of midline, each bearing a

row of spines, usually covered by dorsal sclerite

near tip of epiproct, extending inward and upward

to dorsal surface, extended portion paired and

highly variable in shape, usually visible dorsally

and can even extend out of epiproct near apex.

Tenth tergum (Figure 48) mostly sclerotized, forming a large flat or concave area anterior to base of

epiproct, sometimes produced at lateral basal margins, usually bearing thin hairs. Ninth tergum

sclerotized but not produced, most heavily sclerotized along distal margin, bearing thin hairs.

FEMALE TERMINALIA (Figure 51).—Seventh sternum enlarged and extended distally forming pregenital plate, covering most or all of eighth

sternum, produced area lightly sclerotized. Eighth

sternum narrow, with sclerotized patch over genital opening. Cerci medium sized, membranous and

simple.

NYMPH.—Gills (Figure 144): 1 cervical gill on

each side of midline, arising outside of lateral

cervical sclerites, gills short and triangular, small

gill-like nubs on inside of lateral cervical sclerites.

Forelegs (Figure 169): femur and tibia of approximately equal length; femur with a whorl of long

spines near distal end, spines in loosely arranged

row, scattered spines along dorsal posterior margin; tibia with a row of long spines along dorsal

posterior margin and with occasional long hairs

intermingled with spines, with a row of spines near

dorsoanterior margin, ventral surface with rows of

spines along anterior and posterior margins. Middle and hind legs without whorls of spines on

femur. Pronotum with fringe of long spines on

lateral margins. Mesonotum with fringe of long

spines. Abdominal terga with fringe of long spines

along distal margins. Cerci with whorls of large

spines at distal margins of segments, intermediate

segmental spines usually absent.

DISCUSSION.—Illiesonemoura is made up mostly

of the species of the Pakistani and Polystigma

groups of Aubert (1959a) and a cluster of species

studied by Jewett in (1958). Nemoura sp. (no.

102) in Kawai (1966a) appears to fit in this genus.

Illiesonemoura is very similar to Zapada in most

respects but can be most easily separated by the

structure of the cervical gills, which in Illieso-
NUMBER 211 19

nemoura are only present outside of the lateral

cervical sclerites (Figure 144). The nymph of

Illiesonemoura can also be separated by the absence of whorls of spines on the femora of the middle and hind legs.

Illiesonemoura is named in honor of Professor

Dr. Joachim lilies of Schlitz, Germany, for his

many contributions to the study of Plecoptera and

especially for his help with this revision.

DISTRIBUTION AND SPECIES LIST.—Palearctic and

Oriental (central Asia). The following species are

placed in the genus Illiesonemoura:

*alabeli (Zhiltzova) 1971, n. comb., central Asia

ampula (Jewett) 1958, n. comb., Himalaya

atripes (Aubert) 1959a, n. comb., Pakistan

battakundi (Aubert) 1959a, n. comb., Pakistan

besali (Aubert) 1959a, n. comb., Pakistan

*bispinosa (Kawai) 1968b, n. comb., Taiwan

cor data (Jewett) 1958, n. comb., Himalaya

•falcifera (Harper) 1975, n. comb., Nepal

"gosainkundensis (Harper) 1975, n. comb., Nepal

lilami (Aubert) 1959a, n. comb., Pakistan

maluksari (Aubert) 1959a, n. comb., Pakistan

Pakistani (Aubert) 1959a, n. comb., Pakistan

polystigma (Aubert) 1959a, n. comb., Pakistan

punctata (Jewett) 1958, n. comb., Himalaya

punjabensis (Jewett) 1958, n. comb., Himalaya

ornata (McLachlan) 1875, n. comb., central Asia

•tuberostyla (Wu) 1962, n. comb., China

*verrucosa (Harper) 1975, n. comb., Nepal

Lednia Ricker

FICURES 52-59, 177

Nemoura (Lednia) Ricker, 1952:27. [Type-species: Nemoura

tumana Ricker.]

Lednia.—lilies, 1966:190.

ADULT.—Length: small (4-6 mm). Wings: macropterous, venation typical for family, hyaline

but with small dark area at cord, veins dark brown

and distinct. Gills: absent but with small single

membranous gill-like nubs on outside of lateral

cervical sclerites, base of lateral cervical sclerites

very broad.

MALE TERMINALIA.—Hypoproct (Figure 58):

sclerotized, elongate, broad at base, becoming

gradually narrower toward pointed apex, extending over paraprocts to base of epiproct; vesicle absent. Paraprocts (Figure 55): composed of 2 lobes;

inner lobes long, thin and heavily sclerotized, with

base at lateral basal margin of cerci and extending

to near midline, apical half extending backward

parallel to hypoproct, long thin and lancet-shaped,

reaching to base of epiproct; outer lobes broad

rounded and membranous, extending from cerci to

midline. Cerci (Figures 56-58): medium, membranous and unmodified. Epiproct (Figures 52-

54): broad, flat and mostly sclerotized, bilaterally

symmetrical; dorsal sclerite with broad flat base,

lateral corners broad and darkly sclerotized, extending forward near apex, replaced at apex by 2

long narrow patches of small spines or overlapping

plates, very lightly sclerotized; basal sclerites as

large broad triangles, located at basolateral corners

of epiproct; ventral sclerite large and broad at

base, thin in cross section and trough-shaped, becoming gradually narrower toward pointed apex,

sclerite smooth and naked. Tenth tergum (Figure

56) highly modified, median basal area deeply concave, sclerotized and forming a large smooth surface under epiproct, posterior lateral areas greatly

produced into long narrow prongs, with rounded

heavily sclerotized tips, bearing many small stout

spines. Ninth tergum lightly sclerotized, not produced or modified, bearing a few small spines and

hairs.

FEMALE TERMINALIA (Figure 59).—Seventh sternum slightly produced and enlarged at midline,

produced area lightly sclerotized. Eighth sternum

mostly sclerotized, median area sclerotized and

formed into broad somewhat triangular subgenital

plate, apex rounded or slightly bilobed and covering genital opening, lateral posterior areas with

large, slightly triangular sclerotized patches. Cerci

small, membranous and unmodified. Paraprocts

large, triangular and with rounded corners.

NYMPH.—Unknown.

DISCUSSION.—This genus is monotypic and has

only been collected from two localities in Glacier

National Park (Ricker, 1952; Gaufin et al., 1972).

It lacks a vesicle on the ninth sternum of the male

(Figure 58) like Paranemoura perfecta but otherwise is most similar to Visoka cataractae in the

details of the epiproct and paraprocts. Visoka is,

however, very distinctive because it has submental

gills and also has sclerotized, modified cerci.

Further collecting in small streams at high elevations in the Northern Rocky Mountains should

add to our knowledge of this rare genus.

DISTRIBUTION.—Nearctic (western North America). The genus Lednia contains only one species:

tumana (Ricker) 1952, Montana
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Nemoura Latreille

FIGURES 60-67, 145, 154, 171, 178

Nemoura Latreille, 1796:101.—International Commission on

Zoological Nomenclature, 1963:29-30. [Type-species: Perla

cinerea Retzius = Nemoura cinerea (Retzius).]

ADULT.—Length: small to large (5-15 mm).

Wings (Figure 154): macropterous, venation typical for family, hyaline, fumose or mottled, veins

dark and distinct. Gills (Figure 145): absent but

with single small membranous gill-like nubs outside of lateral cervical sclerites.

MALE TERMINALIA.—Hypoproct (Figure 66):

broad at base, tapering to narrow apex, extending

distally to base of paraprocts, often covering inner

lobes; vesicle present. Paraprocts (Figure 63):

consisting of 2 lobes; inner lobes sclerotized, short

and narrow, closely applied to midline and usually

turned inward, sometimes completely hidden by

hypoproct; outer lobes sclerotized ventrally and

membranous dorsally, very large and triangular or

elongate in shape. Cerci (Figures 64-66): mostly

sclerotized, lateral portion as sclerotized strip that

usually terminates at apex into 1 to 3 spines or

hooks, but spines may occur anywhere or be entirely absent, body of cercus usually elongate and

of moderate width but sometimes greatly enlarged

at base. Epiproct (Figures 60-62): generally short

and broad in dorsal aspect, short and varying from

broad to rather thin in lateral aspect, apex

rounded, completely recurved, sclerotized and

membranous, bilaterally symmetrical; dorsal sclerite large and broad at base of epiproct, extending

dorsolaterally, becoming narrow around lateral

knobs and then very large and triangular, usually

completely covering lateral aspects of epiproct and

part of ventral aspect, most darkly sclerotized areas

at base and near dorsoposterior margin immediately ahead of basal cushion, anterior area usually

lightly sclerotized but bearing spines in some species; basal sclerites as 2 large, broad, triangular or

rectangular patches located near basolateral corners of epiproct; ventral sclerite darkly sclerotized,

broad at base, with lateral knobs at basolateral

corners, becoming narrower toward apex, forming

parallel ridges, 1 on each side of midline, each

bearing a row of spines, usually covered by dorsal

sclerite near tip of epiproct, extending inward and

upward to dorsal surface, visible portion paired

and quite variable in shape, usually semicircular

near apex, often bearing spines or hooks. Tenth

tergum (Figure 64) mostly sclerotized, forming a

large flat or concave area anterior to base of epiproct, usually bearing only thin hairs and small

spines but with large spines or projections in some

exotic species. Ninth tergum sclerotized, but not

produced, bearing thin hairs and spines. Eighth

tergum produced and modified in some species.

FEMALE TERMINALIA (Figure 67).—Seventh sternum enlarged and extended distally, covering most

or all of eighth sternum, produced area sclerotized.

Eighth sternum narrow and mostly membranous,

with small sclerotized area at genital opening.

Cerci mostly sclerotized, quite angular and truncate at apex.

NYMPH.—Gills (Figure 145): absent but with

very small nubs outside of lateral cervical sclerites

on each side of midline. Forelegs (Figure 171):

femur and tibia of approximately equal length,

whole leg usually covered with small spines or

hairs; femur with randomly scattered large spines

along posterior margin and near distal end (N.

mortoni has whorls of large spines), with a few

long hairs often intermingled with spines; tibia

with rows of small spines along dorsal anterior and

posterior margins, ventral surface without rows of

spines or with very short spines. Pronotum with

fringe of short spines on lateral margins. Mesonotum and metanotum with sparse fringe of small

spines along lateral margins. Abdominal terga with

fringes of short spines along distal margins. Cerci

with whorls of spines at distal margins of segments,

intermediate segmental spines very small and

sparse.

DISCUSSION.—Nemoura, the type-genus, received

many species over the years that belonged in

Nemouridae but could not be easily assigned to

any of the known genera. Most of these misplaced

species have been removed and placed in other

genera in this paper. The genus is, however, still

not a completely homogeneous unit, as evidenced

by the included species groups.

As a result of this study, I feel that the typical

Nemoura species are primarily found in more

northern areas. The species figured in this paper

belong to the typical concept of Nemoura. Nemoura males have distinctive sclerotized

cerci with terminal hooks (Figures 64—66). The

females have a large pregenital plate and lightly

sclerotized truncate cerci (Figure 67).
NUMBER 211 21

I did, however, delineate two species complexes

from exotic areas in Asia that, although they are

here placed in Nemoura, exhibit some variation

in the structure of the male terminalia. These species complexes can be distinguished by the following characteristics:

Cercispinosa Complex: Cerci enlarged and

quite thick, bearing one or more spines at apex,

near base or somewhere along the length. Tenth

tergum with large spines or protuberances. Ninth

tergum bearing large spines and somewhat produced. Aubert (1967) in his study of the Nemouridae of Assam provides excellent figures of many

species in this complex.

Ovocercia Complex: Cerci enlarged laterally

and very broad at base, ending in a rounded tip,

body of cerci naked of spines. Tenth tergum produced and often with large humps or knobs.

Ninth tergum usually produced and bearing

spines. Eighth tergum sometimes produced posteriorly. Kawai (1967) and Zwick (1973b) give

figures of species that belong to this complex.

The distribution lists of the species included in

the above complexes are separated from the rest of

the Nemoura species for convenience.

DISTRIBUTION AND SPECIES LIST.—Holarctic and

Oriental (Europe, Asia, North Africa, North

America).

arctica Esben-Petersen 1910, circumpolar

avicularis Morton 1894, Europe, Siberia

babiagorensis Sowa 1964, Poland

braaschi Joost 1970a, Bulgaria

brevipennis Martynov 1928, Caucasus, Armenia, Asia Minor

bulgarica Rauser 1962, Bulgaria

cambrica Stephens 1835, Europe

carpathica lilies 1963, Carpathians

caspica Aubert 1964a, Iran

ceciliae Aubert 1956b, Spain

cinerea (Retzius) 1783, Europe, central Asia

confusa Zwick 1970, France

dubitans Morton 1894, central and northern Europe

elegantula Martynov 1928, Caucasus

erratica Claassen 1936, western Europe, Britain

flaviscapa Aubert 1956a, Greece

flexuosa Aubert 1949, Europe, Asia Minor

fuhiceps Klapalek 1902, Europe

fusca Kis 1963, Romania

•gladiala U£no 1929, Japan

hamata Kis 1965b, Romania

hamulata Zhiltzova 1971, Tien Shan

hesperiae Consiglio 1958, Italy

illiesi Mendl 1968b, Austria

irani Aubert 1964a, Iran

kownackorum Sowa 1970, Bulgaria

kuwayamai Kawai 1966b, Japan

lacustris Pictet 1865, Pyrenees

lepnevae Zhiltzova 1971, Tien Shan

levanidovae Zwick 1974, Siberia

lingulata Navas 1918, Pyrenees

longicauda Kis 1964, Carpathians

marginata Pictet 1835, central Europe

martynovia Claassen 1936, Caucasus, Asia Minor

*tnatangshanensis Wu 1935, China

minima Aubert 1946, Alps

monticola Rauser 1965, Czechoslovakia

mortoni Ris 1902, central Europe

moselyi Despax 1934, Pyrenees

naraiensis Kawai 1954, Japan

navasi Aubert 1953c, Spain, Sicily

normani Ricker 1952, Alaska

obtusa Ris 1902, central Europe

ovoidalis Kis 1965a, Romania

palliventris Aubert 1953b, Italy

*papilla Okamoto 1922, Japan

peristeri Aubert 1963b, Yugoslavia

pirinensis Rauser 1962, Bulgaria

pygmaea Braasch & Joost 1972, Bulgaria

rickeri Jewett 1971, Alaska

rifensis Aubert 1960, Morocco, Spain

sachaliensis Matsumura 1911, Japan, Sachalin

sahlbergi Morton 1896, Arctic Eurasia, Mongolia, Korea

sciurus Aubert 1949, central Europe

'securigera Klapalek 1907, China

sinuata Ris 1902, Alps

subtilis Klapalek 1895, Carpathians, Balkans, Asia Minor

taurica Zhiltzova 1967, Crimea, Asia Minor

transsylvanica Kis 1963, Romania

trispinosa Claassen 1923, eastern North America

*uenoi Kawai 1954, Japan

undulata Ris 1902, Alps

viki Lillehammer 1972, Norway

CERCISPINOSA COMPLEX

*arlingtoni Wu 1940, China

*bispinosa Kawai 1968b, Taiwan

bokhari Aubert 1967, Assam

•cercispinosa Kawai 1960, Japan

chattriki Aubert 1967, Assam

chugi Aubert 1967, Assam

•fulva Samal 1921, Japan

•furcocauda (Wu) 1973, China

•janeti Wu 1938, China

khasii Aubert 1967, Assam

kuhleni Aubert 1967, Assam

•manchuriana U6no 1941, Manchuria

mawlangi Aubert 1967, Assam

•nankinensis Wu 1926, China

•needhamia Wu 1927, China

*pseudofluva Kawai 1954, Japan

schmidi Aubert 1967, Assam

serrarimi Aubert 1967, Assam
22

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

spiniloba Jewett 1954, California

spinosa Wu 1940, China, Assam

•yunnanensis Wu 1940, China

OVOCERCIA COMPLEX

*akagii Kawai 1960, Japan

•cochleocercia Wu 1962, China

•geei Wu 1929, China

lahkipuri Aubert 1967, Assam

ovocercia Kawai 1960, Japan

redimiculum Kawai 1966c, Japan

tau Zwick 1973b, Korea

Nemurella Kempny

FIGURES 68-75, 167, 179

Nemura (Nemurella) Kempny, 1898:59. [Type-species: Nemura inconspicua Kempny = Nemurella pictetii Klapalek.]

Nemurella.—Klapalek, 1900:30.

ADULT.—Length: medium (5-10 mm). Wings:

macropterous, venation typical for family, hyaline

to light brown, veins dark and distinct. Gills: absent but with single membranous gill-like nubs on

outside of lateral cervical sclerites.

MALE TERMINALIA.—Hypoproct (Figure 74):

sclerotized, long and drawn out, broadest near

base, becoming abruptly narrower at posterior

margin of ninth sternum, apical half very thin and

extending to bases of outer lobes of paraprocts,

apex thin, rounded and lightly colored; vesicle

present. Paraprocts (Figure 71): consisting of 2

lobes; inner lobes with bases extending to outer

margin near base of cerci, lobes extremely long,

slender and heavily sclerotized, basal third running from point of attachment laterally to apex of

hypoproct, basal portion dorsoventrally flattened,

apical two-thirds twisted so that thin lateral margins are seen in ventral view, extending well beyond tip of abdomen and slightly beyond epiproct,

tips narrow and pointed; outer lobes extending

beyond tip of epiproct, broadest at base, tapering

gradually to narrowly rounded apex, lobes quite

long and thin, round in cross section, most heavily

sclerotized on outer surface, bearing many long

thin hairs. Cerci (Figures 72-74): very large, sclerotized and almost completely covering epiproct in

lateral view, broad and bulbous at base, becoming

long and parallel sided, ending in broadly rounded

apex, most darkly sclerotized laterally, inner surface and apex light in color, outer surfaces bearing

many small fine hairs. Epiproct (Figures 68-70):

completely sclerotized, short broad and bilaterally

symmetrical, not completely recurved but directed

backward slightly from base; dorsal sclerite divided

into 3 darkly sclerotized parts, heavy basal part

shifted forward into a dorsal position, short broad

and concave, forming an open-sided funnel, ending in a narrow apex with a small rounded opening, lateral arms long thin and lying between dorsal part and ventral sclerite on lateral margins,

broadest at base, apex shaped like blunt forcep

which forms half of forceps as lateral pieces come

together at apex; basal sclerites as 2 broad triangular shaped sclerotized patches located at basolateral

corners of epiproct; ventral sclerite flat, broad and

darkly sclerotized, base very broad, tapering to

broadly rounded apex, sclerite slightly concave

dorsally, apex bifurcate with narrow groove, lateral

apical areas rounded and curled under ventrally,

ventral surface of sclerite flat smooth and naked

except for single rows of very small spines along

lateral margins. Tenth tergum (Figure 72) mostly

membranous medially, paired sclerotized lateral

bars extending to base of epiproct, with enlarged

sclerotized areas at bases of cerci, tergum bearing a

long thin patch of hairs in a lightly sclerotized area

below tip of epiproct. Ninth tergum broad, heavily

sclerotized and slightly produced, bearing many

small thin hairs.

FEMALE TERMINALIA (Figure 75).—Seventh sternum forming pregenital plate, greatly enlarged

and expanded medially over eighth sternum, produced area darkly sclerotized, triangular in shape

with rounded apex which covers genital opening.

Eighth sternum narrow, membranous medially, lateral portions produced as darkly sclerotized vaginal

lobes. Cerci lightly sclerotized but otherwise

simple and unmodified. Paraprocts large and triangular in shape, apex narrowly pointed.

NYMPH.—Gills: absent but with single membranous gill-like nubs on outside of lateral cervical sclerites. Forelegs (Figure 167): femur slightly

shorter than tibia, whole leg covered with many

small spines; femur with a few large spines near

distal margin, occasional long hairs along posterior

margin; tibia with rows of small spines along ventral margins, with an occasional long hair intermingled. Pronotum with irregular fringe of long

thin spines along lateral margins, sides somewhat

rounded in dorsal view. Abdominal terga with
NUMBER 211 23

sparse fringes of spines along distal margins, with

2 longitudinal rows of single large spines. Cerci

with whorls of spines at distal margins of segments,

areas between segmental joints with a few small

spines.

DISCUSSION.—Although Nemurella is monotypic

it has been recognized as a separate subgenus since

1898. It has very distinctive bilobed paraprocts

that make it possible to separate it from all of the

European nemourids (Figure 71). The epiproct

is also highly modified with the horseshoe-shaped

base of the dorsal sclerite shifted forward into a

dorsal position (Figures 68-70). The females have

a well-developed pregenital plate and large vaginal

lobes on sternum eight (Figure 75).

It is very common throughout Europe and is

often extremely abundant in spring-fed habitats.

DISTRIBUTION AND SPECIES LIST.—Palearctic (Europe). The records from the Altai Mountains

and Siberia (Zapekina-Dulkeit, 1957, 1961) are

unconfirmed and highly questionable. This genus

contains only one known species:

pictetii Klapdlek 1900, Europe

Ostrocerca Ricker

FIGURES 76-83, 162, 179

Nemoura (Ostrocerca) Ricker, 1952:38. [Type-species: Nemoura foersteri Ricker.]

Ostrocerco.—lilies, 1966:217.

ADULT.—Length: small (4-8 mm). Wings: macropterous, venation typical for family, hyaline

and sometimes slightly darkened around veins near

cord, veins dark and distinct. Gills: absent but with

single membranous gill-like nubs outside of lateral

cervical sclerites.

MALE TERMINALIA.—Hypoproct (Figure 82):

broad at base, large sclerotized and extending back

over base of epiproct, with a bulbous membranous

inner portion, shape varying from long and narrow

to very broad and truncate or flat at apex; vesicle

present. Paraprocts (Figure 79): consisting of 1

darkly sclerotized lobe and 1 membranous lobe on

each side of midline; inner lobes sclerotized, with

bases at lateral margins near base of cerci, lobes

large and quite angular in shape, sometimes with

lateral projections near bend, tip pointed and directed outward; outer lobes rounded membranous

and covered with hairs. Cerci (Figures 80-82):

large, sclerotized and terminating in a pointed tip,

sometimes with a second subapical point, extending laterally beyond tip of epiproct, most heavily

sclerotized laterally and at apex, covered with

many small hairs, sometimes distinctly bent in lateral aspect. Epiproct (Figures 76-78): mostly

sclerotized, short broad and bilaterally symmetrical, not completely recurved but directed at an

oblique angle upward and backward from longitudinal axis of abdomen; dorsal sclerite large and

broad at base, shifted forward dorsally, extending

forward on dorsal and lateral aspects, lateral arms

running from near base to apex along lateral surfaces, often covered by large dorsal area which is

lightly sclerotized and covered by spines or scalelike plates; basal sclerites as 2 long very thin sclerotized patches, curved around basolateral corners

of epiproct; ventral sclerite heavily sclerotized,

broad at base and throughout most of length, apex

pointed or narrowly rounded, usually with one or

more prongs directed downward from or near the

tip, best seen in lateral view, margins bearing small

spines or scales. Tenth tergum (Figure 80) with

paired sclerotized bars leading to base of epiproct,

generally broad and flat or concave but sometimes

with thin paired projections, located 1 on each

side of apex of epiproct, extreme lateral margins

below cerci enlarged and sclerotized, tergum bearing small hairs or spines. Ninth tergum sclerotized

but not produced or modified, bearing many small

hairs.

FEMALE TERMINALIA (Figure 83).—Seventh sternum usually enlarged and expanded distally at

midline, enlarged area sclerotized and varying in

shape from large slightly rounded area to narrow

elongate nipplelike process similar to male hypoproct. Eighth sternum broad and heavily sclerotized, forming subgenital plate, covering genital

opening, truncate, indented or incised on distal

margin at midline. Paraprocts, triangular and

pointed at apex. Cerci short, membranous and

unmodified.

NYMPH.—Gills: absent but with single membranous gill-like nubs outside of lateral cervical

sclerites. Forelegs (Figure 162): femur and tibia of

approximately equal length; femur with large

spines along dorsoposterior margin, randomly

placed and covering entire surface near distal margin, occasionally with one or more long hairs inter-
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mingled at posterior edge; tibia with rows of small

spines along dorsal margins and rows of long spines

along ventral margins, often with very small spines

scattered over surface between margins, usually

with a few long hairs along posterodorsal margin.

Pronotum with irregular fringe of small spines

along lateral margins. Mesonotum and metanotum

with patches of short heavy spines at anterior margin, lateral margins of wing pads with a sparse

fringe of thin spines. Abdominal terga with sparse

fringes of small spines along distal margins, also

with spines covering distal half of abdominal segments. Cerci with whorls of small spines at distal

margins of segments, areas between segmental

joints elongate and naked.

DISCUSSION.—Ostrocerca is a very distinctive genus that is very similar to Nemurella. The epiproct is directed backward at an angle (Figures

80-82) and the dorsal sclerite is moved forward

(Figures 77-78). The paraprocts are modified with

large darkly sclerotized inner lobes (Figure 79)

and the cerci are large, sclerotized and have a

pointed apex (Figures 80-82).

Most of the species are extremely rare in collections, but more collecting in pristine-spring

areas in extreme eastern and western North America should add to the present knowledge.

DISTRIBUTION AND SPECIES LIST.—Nearctic (North

America). The following species are presently

recognized in the genus Ostrocerca:

albidipennis (Walker) 1852, eastern North America

complexa (Claassen) 1937, eastern North America

dimicki (Frison) 1936, northwestern North America

foersteri (Ricker) 1943, northwestern North America

prolongata (Claassen) 1923, eastern North America

truncate (Claassen) 1923, eastern North America

Paranemoura Needham and Claassen

FIGURES 84-91, 153, 165, 180

Nemoura (Paranemoura) Needham and Claassen, 1925:288.

[Type-species: Nemoura perfecta Walker.]

Paranemoura.—Claassen, 1940:50.

ADULT.—Length: medium (5-8 mm). Wings

(Figure 153): macropterous, terminal costal crossvein connected to Sc instead of R as is typical for

the family, hyaline with fumose areas at and near

the cord region, veins dark brown and distinct.

Gills: absent but with small single membranous

gill-like nubs on outside of lateral cervical sclerites.

MALE TERMINALIA.—Hypoproct (Figure 90):

sclerotized, broad at base, tapering to narrow

rounded apex, extending to and between paraprocts; vesicle absent. Paraprocts (Figure 87):

consisting of 1 large sclerotized lobe on each side

of midline, lobes shaped like broad triangles,

covered with short thin hairs. Cerci (Figures 88-

90): short, membranous and unmodified. Epiproct

(Figures 84-86): mostly sclerotized, composed

primarily of ventral sclerite, broad at base with

long thin apex, bilaterally symmetrical, completely recurved; dorsal sclerite with thin base at

posterior basal end, lateral portions of sclerite

extending upward and forward as thin paired arms

which fit into large groove formed by ventral sclerite, apex of paired arms becoming abruptly large

and sickle-shaped, pointed tips directed backward

and protruding slightly from dorsoposterior margins of epiproct; basal sclerites as large broad triangles, located at basolateral corners of epiproct;

ventral sclerite very large, and broad at base, convex ventrally and curving upward laterally to enclose dorsal sclerite, dorsolateral margins heavily

sclerotized and bearing small stout spines, apical

third narrow and elongate, forming naked pointed

apex. Tenth tergum (Figure 88) almost completely

bisected at midline, median membranous area very

broad and concave, with sclerotized areas near base

of epiproct and at base of segment directly under

tip of epiproct, lateral areas produced, heavily

sclerotized and bearing many small spines. Ninth

tergum heavily sclerotized, not produced but constricted and quite narrow at midline, bearing a few

small hairs and spines.

FEMALE TERMINALIA (Figure 91).—Seventh sternum slightly produced at midline, produced area

lightly sclerotized, extending over anterior margin

of eighth sternum. Eighth sternum narrow, posterior median margin indented toward genital

opening, dark triangular sclerotized area over

genital opening. Cerci small membranous and unmodified. Paraprocts large, triangular and with

rounded corners.

NYMPH.—Gills: absent but small membranous

gill-like nubs on outside of lateral cervical sclerites.

Forelegs (Figure 165): tibia slightly longer than

femur, whole leg covered with many small spines;

femur with a few long thin spines along posterior

margin, and on surface near distal end, occasional
NUMBER 211 25

long hairs extending from posterior margin; tibia

with fringes of long thin spines along anterior margins, with an occasional long hair along posterior

margin. Pronotum with many small spines covering entire surface, without definite lateral fringe

of spines, corners rounded, segment longer than

wide. Mesonotum and metanotum with a few stout

spines along- anterior margins. Abdominal terga

sparsely covered with small spines, fringes of spines

along distal margins sparse, with longer spines

near lateral margins. Cerci with whorls of spines

along distal margins of segments, areas between

segmental joints with a few small spines.

DISCUSSION.—Paranemoura is presently monotypic, but I have seen specimens of another species

belonging to this genus. The wings do not exhibit

a perfect "X" because the terminal costal crossvein that usually runs from C to R runs from C to

Sc at the cord (Figure 153). The vesicle is absent

from the ninth sternum (Figure 90) as in Lednia,

but the paraprocts in Paranemoura are single,

short, and membranous (Figure 87). The epiproct

is specialized with the dorsal sclerite being enclosed in groove of the ventral sclerite and located

near the base of the epiproct (Figures 84-85).

This genus is uncommon and known only from

isolated localities in northeastern North America.

DISTRIBUTION AND SPECIES LIST.—Nearctic (North

America). Paranemoura contains one known

species:

perfecta (Walker) 1852, eastern North America

Podmosta Bicker

FIGURES 92-99, 163, 181

Nemoura (Podmosta) Ricker, 1952:42. [Type-species: Nemoura decepta Frison.]

Podmosta.—lilies, 1966:219.

ADULT.—Length: small (3-6 mm). Wings: macropterous, venation typical for family, hyaline

but with dark areas around veins in area of cord,

veins dark brown and distinct. Gills: absent but

with small single membranous gill-like nubs on

outside of lateral cervical sclerites, base of lateral

cervical sclerites very broad.

MALE TERMINALIA.—Hypoproct (Figure 98):

lightly sclerotized medially, heavily sclerotized

around margins and at apex, base broad, tapering

abruptly to narrow pointed apex which reaches to

base of epiproct; vesicle present. Paraprocts (Figure 95): consisting of one large lightly sclerotized

lobe on each side of midline, lobes shaped like

short triangles, covered with many small hairs.

Cerci (Figures 96-98): short membranous and unmodified. Epiproct (Figures 92-94): complex in

structure, often with dorsal and ventral sclerites

separated at apex and throughout length but

joined at base, mostly sclerotized, short broad and

bilaterally symmetrical, not completely recurved

but directed upward at an angle to the body axis;

dorsal sclerite with broad lightly sclerotized base,

often bearing long hairs, base shifted forward to

base of dorsal surface, median area very large and

produced, with a lightly sclerotized dorsal band or

patch, lateral arms of sclerite darkly sclerotized,

long and thin, extending laterally from base to

near apex of epiproct, tips modified as hooks or

prongs; basal sclerites as long, broad triangles,

located at basolateral corners of epiproct; ventral

sclerite long, sclerotized and flat, apex often enlarged and bearing small paired sclerotized patches

ventrally, dorsal surface usually with paired sclerotized bars, curved at apex. Tenth tergum (Figure

96) greatly produced laterally, usually darkly

sclerotized and with many small hairs or spines,

anterior median area concave and membranous,

tergum often nearly completely bisected, lateral

posterior corners enlarged and sclerotized, basal

concave area mostly membranous but with paired

sclerotized bars which attach to base of epiproct.

Ninth tergum not produced or greatly modified,

sometimes with small incised area along anterior

margin, bearing hairs or spines.

FEMALE TERMINALIA (Figure 99).—Seventh sternum usually slightly produced and lightly sclerotized at posterior margin. Eighth sternum with

sclerotized median area that terminates at genital

opening, sometimes overlapping into opening, in

one case formed into a small subgenital plate,

sometimes with small lightly sclerotized vaginal

lobes. Cerci small membranous and unmodified.

Paraprocts small and triangular, with rounded

corners.

NYMPH.—Gills: absent but with small single

membranous gill-like nubs on outside of lateral

cervical sclerites, base of cervical sclerites very

broad. Forelegs (Figure 163): tibia and femur of

approximately equal length; femur with very

small spines and very large spines mixed along
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

posterior margin and on distal third; tibia with

fringe of medium, stout spines along posterior margin, sparse fringe of long hairs also along posterior

margin, whole segment with numerous small

spines. Pronotum with sparse areas of small spines

along lateral margins but without a definite lateral

fringe, corners rounded producing an almost oval

shape. Mesonotum and metanotum with a few

stout spines at anterior margins. Abdominal terga

with fringes of moderate spines along distal margins. Cerci with whorls of spines at distal margins

of segments, areas between segmental joints naked

near base of cerci but with occasional spines near

apex.

DISCUSSION.—Podmosta is a small genus and is

very distinctive in both the male and female. The

males have an epiproct which is directed upward

at an angle almost perpendicular to the body axis

(Figure 97). Both the dorsal and ventral sclerites

are highly developed and often nearly separate

except at the base (Figures 92-94). The females

have a distinct darkly sclerotized patch along the

midline of the eighth sternum (Figure 99).

This genus is common in western North America

in small cold streams at high elevations. It also

occurs in northeastern Canada and one species has

recently been collected by Dr. L. A. Zhiltzova in

the Soviet Far East (pers. coram.).

DISTRIBUTION AND SPECIES LIST.—Holarctic

(North America, Asia). The genus Podmosta contains the following species:

decepta (Frison) 1942, western North America

delicatula (Claassen) 1923, western North America

macdunnoughi (Ricker) 1947, eastern Canada

obscura (Frison) 1936, northwestern United States

weberi (Ricker) 1952, Alaska, northeastern Asia

Prostoia Ricker

FIGURES 100-107, 148, 164, 182

Xemoura (Prostoia) Ricker, 1952:47. [Type-species: Nemoura

completa Walker.]

Prostoia.—lilies, 1966:220.

ADULT.—Length: small (4-7 mm). Wings (Figure 148): macropterous, venation typical for

family, hyaline with scattered fumose areas, large

dark areas near apex and beginning at cord and

extending to posterior margin of forewing, distinctive narrow light vertical stripe formed between fumose areas, veins dark brown and distinct.

Gills: absent but with small single membranous

gill-like nubs on outside of lateral cervical sclerites.

MALE TERMINALIA.—Hypoproct (Figure 106):

sclerotized, broad at base and throughout most of

length, apex tapering abruptly to narrow pointed

tip, extending between paraprocts; vesicle present.

Paraprocts (Figure 103): single lightly sclerotized

lobe on each side of midline, lobes .shaped like

small short equilateral triangles, bearing many

thin hairs. Cerci (Figures 104-106): moderately

long, membranous and unmodified. Epiproct (Figures 100-102): completely sclerotized, dorsal sclerite and ventral sclerite almost completely separated, ventral sclerite comprising most of epiproct

which is long, narrow and completely recurved

dorsally; dorsal sclerite with long, moderately thick

base, running from one basolateral corner to the

other, not extending above base of epiproct, lateral arms beginning at lateral corners, fused to

base of sclerite, hooking forward and outward

nearly parallel to ventral sclerite, usually very

small but nearly three-fourths length of epiproct

in one species; basal sclerites as long broad triangles, located at basolateral corners of epiproct

below base of dorsal sclerite; ventral sclerite

heavily sclerotized, lateral aspect long and narrow,

curving upward laterally to form dorsal surfaces,

dorsal aspect long but quite broad, bilaterally symmetrical, with a definite dorsal groove, whole sclerite smooth and naked. Tenth tergum (Figure

104) little produced but extending forward and

nearly dividing ninth tergum at midline, mostly

sclerotized, flat and broad, almost completely bisected at midline by narrow membranous band

that lies directly under ventral sclerite, sclerotized

portions covered with many small hairs or spines.

Ninth tergum broad laterally but very narrow and

almost divided at midline, not produced but

covered with many small hairs or spines.

FEMALE TERMINALIA (Figure 107).—Seventh sternum slightly produced, broadly rounded and often

appearing connected to eighth sternum at midline,

produced area often lightly sclerotized. Eighth

sternum broad and little developed, sclerotized at

posterior margin near genital opening, subgenital

plate poorly developed except in one species where

the posterolateral margins are extended as short

flat platelike lobes. Cerci small membranous and

unmodified. Paraprocts small and triangular, with

rounded corners.
NUMBER 211 27

NYMPH.—Gills: absent but with small single

membranous gill-like nubs on outside of lateral

cervical sclerites. Foreleg (Figure 164): tibia and

tarsus of approximately equal length, most of leg

covered with many small spines; femur with few

very large spines found along posterior margin on

distal half; tibia with fringe of large spines along

posterior margin, fringe of long hairs intermingled

along posterior margin. Pronotum with sparse

areas of small spines on lateral margins but without definite lateral fringe, corners rounded producing an almost oval shape. Mesonotum and

metanotum with a few stout spines at anterior margins. Abdominal terga with fringes of spines along

distal margins, sometimes with two longitudinal

rows of single or double long spines. Cerci with

whorls of spines at distal margins, areas between

segmental joints naked except for occasional spines

near apex of cerci.

DISCUSSION.—Prostoia is highly modified and is

very similar in the shape of the epiproct to some

species of Capniidae, especially the genus Mesocapnia. The males have a very small dorsal sclerite

located at the basolateral margins of the epiproct.

The ventral sclerite is very large and completely

sclerotized both dorsally and ventrally (Figures

100-102). The females have no definite pregenital

or subgenital plate but only a small sclerotized area

along the posterior margin of the eighth sternum

(Figure 107).

This genus can be very common in the early

spring and is often collected on bridges along with

species of Capniidae and Taeniopterygidae.

DISTRIBUTION AND SPECIES LIST.—Nearctic (North

America). The genus Prostoia includes three

known species:

besametsa (Ricker) 1952, western North America

completa (Walker) 1852, eastern North America

similis (Hagen) 1861, eastern North America

Shipsa Ricker

FIGURES 108-115, 149, 166, 183

Nemoura (Shipsa) Ricker, 1952:49. [Type-species: Nemoura

rotunda Claassen.]

Shipsa.—lilies, 1966:247.

ADULT.—Length: medium (5-8 mm). Wings

(Figure 149): macropterous, venation typical for

family, basically hyaline but with scattered dark

areas in forewing, large dark area at apex and wide

dark stripe running downward from cord, separated by narrow light strip running the width of

forewing, veins dark and distinct. Gills: absent but

with small single membranous gill-like nubs on

outside of lateral cervical sclerites, base of lateral

cervical sclerites very broad.

MALE TERMINALIA.—Hypoproct (Figure 114):

sclerotized, short and broad, broadest at base,

tapering abruptly at apex to short narrow rounded

tip, extending to base of epiproct; vesicle present.

Paraprocts (Figure 111): consisting of 1 large

heavily sclerotized lobe on each side of midline,

lobes shaped like short broad triangles. Cerci (Figures 112-114): medium, membranous and unmodified. Epiproct (Figures 108-110): mostly sclerotized, short, broad and bilaterally symmetrical,

not completely recurved but directed upward

slightly at base; dorsal sclerite divided into 3

heavily sclerotized parts, heavy basal part shifted

somewhat forward and extending onto dorsal surface at lateral margins, base broad and heavily

sclerotized, apical portions broad and ending in

slanted points, lateral arms arising below apices of

basal part, long thin and horn-shaped, curving

outward and upward at narrow apex; basal sclerites large and broad, shaped like short broad triangles, located at basolateral corners of epiproct, usually covered by paraprocts; ventral

sclerite flat and very broad ventrally, naked except

for single small stout spines on anterolateral margins, paired lightly sclerotized bands extending

outward from apex, bands curving backward onto

epiproct between dorsal sclerite and extending upward dorsally where they become very large, appearing as two thick sclerotized forceplike plates

on dorsal surface. Tenth tergum (Figure 112) bisected medially, paired lateral sclerotized bars

leading to base of epiproct, lateral distal portions

large and produced into long inward curved

processes, outer margin of processes heavily sclerotized, inner margin membranous, inner and dorsal

surfaces covered with many small dark spines.

Ninth tergum incised along basal margin forming

light membranous hemispherical area, median

sclerotized portion narrow. Terga 8 to 5 with

hemispherical membranous areas at base, membranous areas becoming smaller toward anterior

portion of abdomen.

FEMALE TERMINALIA (Figure 115).—Seventh sternum little produced, extending only slightly onto
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

eighth sternum, with broad lightly sclerotized area

along posterior margin. Eighth sternum broad,

mostly sclerotized, median area developed into narrow sclerotized subgenital plate which reaches to

posterior margin of sternum, small dark triangular

patch located near genital opening, large triangular sclerotized patches extending from basolateral

margins of sternum to midline, broad sclerotized

bands on posterolateral margins. Cerci small,

membranous and unmodified. Paraprocts small

and triangular, with rounded corners.

NYMPH.—Gills: absent but with small single

membranous gill-like nubs on outside of lateral

cervical sclerites, base of cervical sderites very

broad. Forelegs (Figure 166): tibia slightly longer

than femur, whole leg covered with many mediumsized spines; femur with numerous long spines

scattered along posterior margin; tibia with fringe

of long thin hairs along posterior margin. Pronotum with numerous small spines scattered over

surface but without a definite lateral fringe. Mesonotum and metanotum with scattered small spines.

Abdominal terga covered with numerous small

spines, with fringes of small spines along distal

margins. Cerci with whorls of spines at distal margins of segments, areas between segmental joints

with a few large spines, number of spines increasing from base to apex of cerci.

DISCUSSION.—Shipsa is a very distinctive genus,

especially in the male terminalia. The tenth tergum is produced into long terminal projections,

1 on each side of the epiproct (Figures 112-113).

The epiproct is modified with the ventral sclerite

extending to the dorsal surface and terminating in

a prominent forcep-shaped structure (Figures 108-

110). The nymphs have a definite fringe of long

hairs along the posterior margins of the tibiae

(Figure 166).

It is common in eastern and northern North

America in the early spring.

DISTRIBUTION AND SPECIES LIST.—Nearctic (North

America). This genus contains a single known

species:

rotunda (Claassen) 1923, eastern and northern North America

Soyedina Ricker

FIGURES 116-123, 152, 170, 184

Nemoura (Soyedina) Ricker, 1952:50. [Type-species: Nemoura

vallicularia Wu.]

Soyedina.—lilies, 1966:247.

ADULT.—Length: medium (6-12 mm). Wings

(Figure 152): macropterous, venation altered so

that Ai and A2 of forewings merge slightly before

wing margin, generally hyaline, with scattered

fumose areas, without definite mottled pattern,

veins dark and distinct. Gills: absent but with

single membranous gill-like nubs outside of lateral

cervical sclerites. {

MALE TERMINALIA.—Hypoproct (figure 122):

broadest at base, quite long and thin, extending

distally to base of paraprocts, apical third bent

abruptly upward, terminating in a very narrow tip;

vesicle present. Paraprocts (Figure 119): consisting

of 2 lobes (inner lobe sometimes difficult to observe); inner lobes lightly sclerotized, short and

narrow, closely applied to inner margin of outer

lobes, often completely hidden by hyr)6proct; outer

lobes large and darkly sclerotized, with small membranous area along inner margin and near apex,

very broad at base and extending around cerci,

apical half elongate and rectangular in shape,

sometimes with serrated margin or blunt process

near apex, usually completely enclosing epiproct

laterally. Cerci (Figures 120-122): small, simple

and membranous. Epiproct (Figures 116-118):

mostly sclerotized, short thin and asymmetrical in

both dorsal and lateral views, not completely recurved, but directed at an oblique angle upward

and backward from longitudinal axis of abdomen,

left half similar but smaller than right half; dorsal

sclerite broad at base of epiproct, extending toward

apex, becoming abruptly very narrow near base

and then broad again in apical two-thirds, covering

most of dorsal and lateral aspects, faost heavily

sclerotized areas at base and directly ahead of basal

cushion, apical area lightly sclerotized but covered

with numerous small spines or scalelike plates;

basal sclerites as 2 large elongate rectangular

patches located near basolateral corners of epiproct; ventral sclerite heavily sclerotized, broad at

base, with small lateral knobs at basolateral corners, becoming narrower toward apex, forming 2

parallel bars, 1 on each side of midline, each bearing a row of spines, narrow projection extending

inward and upward to dorsal surface near .apex,

projection ending in single tube-shaped process or

double asymmetrical processes. Tenth tergum (Figure 120) moderately produced, mostly sclerotized,

forming a large flat area antepor to base of epiproct, surface naked or with a few very small hairs
NUMBER 211 29

or spines. Ninth tergum sclerotized but not produced, sclerotized portion very narrow near midline, sometimes with very small hairs or spines.

Anterior abdominal tergites sometimes modified as

elevated knobs or projections.

FEMALE TERM IN ALIA (Figure 123).—Seventh sternum enlarged and extended distally, covering part

or all of eighth sternum, produced area lightly

sclerotized. Eighth sternum narrow and excavated

at midline, mostly membranous but with a distinct

darkly sclerotized patch at genital opening. Paraprocts triangular and pointed at apex. Cerci short,

simple and membranous.

NYMPH.—Gills: absent but with very small single

nubs outside of lateral cervical sclerites on

each side of midline. Forelegs (Figure 170): femur

and tibia of approximately equal length, whole

leg usually covered with small spines or hairs; femur with randomly scattered large spines dorsally

on distal portion, extending from anterior to posterior margin, with a few long hairs intermingled

along posterior margin; tibia with rows of large

spines somewhat randomly distributed on anterior,

posterior, and dorsal surfaces, ventral inner surface

naked or with a few small spines. Pronotum not

completely square or rectangular, with definite

notch on lateral margins, basal third somewhat

narrower, fringe of short stubby spines on all margins. Mesonotum and metanotum with sparse

fringe of small spines on lateral margins of wingpads. Abdominal terga with fringe of short spines

along distal margins, often with lateral rows of

single long hairs. Cerci with whorls of spines at

distal margins of segments, intermediate segmental

spines very small, sparse and located near base of

cerci.

DISCUSSION.—Soyedina is the only genus in Nemouridae where the shape of the epiproct is always

asymmetrical. The epiproct is bisected down the

middle and the right half is larger than the left

half -(Figures 116-118). The paraprocts are also

greatty enlarged and highly modified into ventrolateral structures which often nearly enclose the

epiproct (Figures 119, 122). The character that

veins Ax and A2 of the forewings are joined (Figure 152) does not always hold true. Sometimes

only one wing exhibits this character and rarely

neither wing.

This genus is usually found in clear cold streams

at high elevations and emerges in the spring or

early summer.

DISTRIBUTION AND SPECIES LIST.—Nearctic (North

America). The following species are known in the

genus Soyedina:

carolinensis (Claassen) 1923, southeastern United States

interrupta (Claassen) 1923, northwestern North America

nevadensis (Claassen) 1923, California, Nevada

potteri (Baumann and Gaufin) 1971, Idaho, Montana

producta (Claassen) 1923, northwestern North America

vallicularia (Wu) 1923, eastern North America

Washington! (Claassen) 1923, northeastern North America

Visoka Ricker

FIGURES 124-131, 146, 168, 185

Nemoura (Visoka) Ricker, 1952:53. [Type-species: Nemoura

cataractae Neave.]

Visoka.—lilies, 1966:249.

ADULT.—Length: small to medium (5-8 mm).

Wings: macropterous, venation typical for family,

clear or slightly cloudy around veins, which are

dark and distinct. Gills (Figure 146): 1 branched

gill on each side of midline, between labium and

submentum, gill branches 5, divided nearly to base.

MALE TERM IN ALIA.—Hypoproct (Figure 130):

broad at base, tapering to narrow apex, extending

distally over inner lobes of paraprocts and sometimes to base of epiproct; vesicle present. Paraprocts (Figure 127): divided into 2 lobes; inner

lobes darkly sclerotized, naked, narrow and sickleshaped, closely applied to midline, sometimes completely hidden by hypoproct; outer lobes darkly

sclerotized at broad triangular base, apical half

bluntly rounded and membranous, covered with

small hairs. Cerci (Figures 128-130): heavily

sclerotized dorsally and with a large distinct spine

at the inner apical margin, lightly sclerotized

laterally, membranous ventrally and covered with

small hairs, tip with a small light area which encircles the small brown segment remnant. Epiproct (Figures 124-126): elongate and narrow in

dorsal view, bilaterally symmetrical, elongate and

recurved in lateral view, with widest area at

bend; dorsal sclerite broad but very short at base,

apical portion narrow, lateral arms extending

dorsolaterally toward apex, arms thin and narrow

extending under large lightly sclerotized area near

base; basal sclerites, as 2 thin triangular plates,
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

located at basolateral margins of epiproct; ventral

sclerite heavily sclerotized, broadest at base, tapering to narrow apex, forming total epiproct at apex

(apical third), ventral keel-shaped ridge naked of

conspicuous spines. Tenth tergum (Figure 128)

mostly sclerotized, forming a large flat dorsal area,

with rounded membranous area under apex of epiproct, median sclerotized patch located beyond

tip of epiproct, lateral margins produced and

slightly angular. Ninth tergum mostly sclerotized,

narrow at midline, slightly produced, bearing small

hairs.

FEMALE TERMINALIA (Figure 131).—Seventh sternum produced and lightly sclerotized, extending

over half or more of segment eight and covering

the genital opening, forming a large nipplelike

structure in lateral aspect. Eighth sternum narrow

and with a distinct darkly sclerotized patch over

genital opening. Ninth sternum wide and only

lightly sclerotized but with more darkly sclerotized areas near lateral margins. Cerci lightly sclerotized, truncate at apex, with a small projection

on inner apical margins.

NYMPH.—Gills (Figure 146): 1 branched gill on

each side of midline between labium and submentum, gill branches 5, all divided nearly to base.

Forelegs (Figure 168): femur and tibia of approximately equal length, most of leg covered with small

fine hairs; femur with fringe of small hairs along

posterior margin, with a diagonal whorl of long

irregular spines on distal half; tibia with row

of very long spines along posterior margin, with

many long hairs intermingled, a row of small

spines along anterior margin, with an occasional

large spine in between. Pronotum with irregular

fringe of spines along lateral margins (Figure 146).

Abdominal terga with very sparse fringes of hairs

along distal margins, composed mostly of long

hairs in longitudinal rows. Cerci with whorls of

spines at distal margins of segments, areas between

joints with occasional small spines.

DISCUSSION.—Visoka is the only genus in the

Nemouridae that has submental gills (Figure 146).

The structure of the cerci and epiproct is similar

to some species in the Cercispinosa Complex of

Nemoura, but those species have simple bulbous

paraprocts and Visoka has a narrow sclerotized

inner lobe and a broad outer lobe, which has a

membranous apex (Figure 127).

Visoka is known only from scattered localities in

western North America.

DISTRIBUTION AND SPECIES LIST.—Nearctic (North

America). The genus Visoka contains one species:

cataractae (Neave) 1933, western North America

Zapada Ricker

FIGURES 1, 132-139, 147, 150, 172, 186

Nemoura (Zapada) Ricker, 1952:54. [Type-species: Nemoura

haysi Ricker.]

Zapada— lilies, 1966:250.

ADULT.—Length: small to large (5-14 mm). Wings

(Figure 150): macropterous or brachypterous, venation typical for family in macropterous species,

fumose or mottled, veins dark and distinct. Gills

(Figure 147): 2 simple cervical gills on each side

of midline, 1 arising inside and 1 outside of lateral

cervical sclerites, usually unbranched but with 3 or

4 branches in Z. cinctipes (branches arising beyond

gill base).

MALE TERMINALIA.—Hypoproct (Figure 138):

broad at base, tapering to pointed apex, extending

distally over inner lobes of paraprocts, tip extending nearly to base of epiproct; vesicle present.

Paraprocts (Figure 135): consisting of 2 lobes;

inner lobes sclerotized, long narrow and closely

applied at midline, usually completely hidden by

hypoproct; outer lobes sclerotized ventrally and

membranous dorsally, very large and almost square

or rectangular-shaped. Cerci (Figures 136-138):

membranous, short and unmodified. Epiproct

(Figures 132-134): completely recurved, mostly

sclerotized, bilaterally symmetrical, short and

broad in dorsal aspect, short and broad to narrow

laterally, apex narrow; dorsal sclerite very large

and broad at base of epiproct, extending dorsolaterally, becoming narrow around lateral knobs

and then very large and triangular, often completely covering lateral portions of epiproct, most

darkly sclerotized areas at base and near posterior

end immediately ahead of basal cushion, anterior area lightly sclerotized and usually composed

of many small overlapping scales or plates; basal

sclerites as 2 broad triangular or rectangular

patches located near basolateral corners of epiproct; ventral sclerite heavily sclerotized, broad

at base, with lateral knobs at basolateral cor-
NUMBER 211 31

ners, becoming gradually narrower toward

apex, forming a flat or convex base, bearing rows

of long spines along lateral ridges, often covered

by dorsal sclerite near apex, sometimes extending

inside epiproct to dorsal margin, extended portion paired and sometimes visible in lateral view.

Tenth tergum (Figure 136) mostly sclerotized,

forming a large flat or concave area anterior to

base of epiproct, bearing hairs and small spines,

especially near proximal margin. Ninth tergum

sclerotized but not produced, sclerotized portion

narrow and with a few hairs and small spines.

FEMALE TERMINALIA (Figure 139).—Seventh

sternum enlarged and extended at distal margin

forming large pregenital plate, covering most or all

of eighth sternum, produced area heavily sclerotized. Eighth sternum narrow, with a dark sclerotized patch over genital opening. Cerci short

membranous and unmodified. Paraprocts broad

and somewhat triangular.

NYMPH (Figure 1).—Gills (Figure 147): 2 cervical gills on each side of midline, 1 arising inside

and 1 outside of lateral cervical sclerites, usually

single and elongate, sometimes constricted but

with 3 or 4 branches arising beyond gill base in

Z. cinctipes. Forelegs (Figure 172): femur and

tibia of approximately equal length; femur with

a whorl of long spines near distal end, spines arranged in rows, with small space between dorsal

and ventral arcs, with a few scattered spines along

dorsoposterior margin; tibia with row of long

spines along dorsoposterior margin; often with

long hairs intermingled, ventral surface with rows

of long spines along anterior and posterior margins. Whorls of spines present on femora of middle and hindlegs. Pronotum (Figure 1) with

fringe of large stout spines on lateral margins

which often completely encircle the outer margins.

Mesonotum and metanotum (Figure 1) with

fringes of long stout spines along lateral margins.

Abdominal terga (Figure 1) with fringes of spines

along distal margins, often with some enlarged and

arranged in longitudinal rows. Cerci (Figure 1)

with whorls of large spines at distal margins of

segments, intermediate segmental spines absent or

rare near apex of cercus.

DISCUSSION.—Zapada is the most common genus

of Nemouridae in western North America. Zapada

cinctipes Banks is found in almost every flowingwater habitat and sometimes emerges throughout

the year as recorded by Hales and Gaufin (1971).

The genus is not uncommon in Alaska and could

also be present in eastern Asia. Most species emerge

very early in the year and are collected along with

the winter Capniidae.

Zapada is a very distincitve genus in both the

adult and nymphal stage. The combination of a

single, simple gill on each side of the lateral cervical sclerites (Figure 147), large angular outer paraproctal lobes (Figures 135) and a short, broad epiproct with the dorsal sclerite well developed (Figures 132-134) serves to separate this from all other

Nemouridae. The nymphs have whorls of large

spines on all femora (Figure 1).

DISTRIBUTION AND SPECIES LIST.—Nearctic (North

America). The following species are in the genus

Zapada:

chila (Ricker) 1952, Tennessee

cinctipes (Banks) 1897, western North America

columbiana (Claassen) 1923, western North America

cordillera (Baumann and Gaufin) 1971, western North

America

frigida (Claassen) 1923, western North America

glacier (Baumann and Gaufin) 1971, Montana

hay si (Ricker) 1952, western North America

oregonensis (Claassen) 1923, western North America

Species Incertae Sedis

The generic placement of the following species

of the family Nemouridae is uncertain at this time.

In most cases the original description and drawings

are very poor, and fresh specimens were not available for study. Sometimes the species is only known

from the type-specimen, which is in very poor

condition.

Although some of these species have been placed

in other genera by some authors, I am simply listing them as they were originally described.

"Amphinemura microcercia Wu 1938, China

*Nemoura asakawae Kohno 1941, Japan

Nemoura babai Kawai 1966c, Japan

*Nemoura bituberculata Kimmins 1950b, Assam

Nemoura denticulata Kawai 1954, Japan

Nemoura elephas Zwick 1974, Taiwan

'Nemoura grandicauda Wu 1973, China

*Nemoura hangchowensis Chu 1928, China

Nemoura spinosa Kawai 1960, Japan

Nemoura stratum Kawai 1966c, Japan

Nemoura wahkeena Jewett 1954, Oregon

Protonemura cherrapunjii Aubert 1967, Assam

*Protonemura chinonis Okamoto 1922, Japan
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

*Protonemura filigera Kawai 1969, Thailand

•Protonemura longiplatta Wu 1949, China

•Protonemura recurvata Wu 1949, China

Protonemura scutigera Kimmins 1950b, Assam

'Protonemura trifurcata Wu 1949, China

Phylogeny

This study of the phylogeny of the Nemouridae

is based on Hennig's (1966) method of "phylogenetic systematics." Hennig's method is of the

cladistic type and is as follows: taxa to be monophyletic must be based upon synapomorphic character states. Character states are identified as either

apomorphic or plesiomorphic and a system of

monophyletic taxa is established. Within this system, two monophyletic taxa form a sister-group

pair if each taxon exhibits a different apomorphic

character state that is plesiomorphic in the other,

and if they share a third apomorphic character

state. The two taxa are then sister groups of each

other. The method is to search for the sister group

(Brundin, 1965; Munroe, 1974).

A similar phylogenetic study was done by Zwick

(1973a) for the order Plecoptera. He included

some information on the Nemouridae but refrained

from a detailed analysis because he knew that I

was revising the family Nemouridae.

The following synapomorphic character states

were cited by Zwick (1973a) for the family

Nemouridae:

1. The structure of the internal male sex organs.

The testes are large and long and radiate from the

anterior bend of the vas deferens in the shape of a

star.

2. The reduction of the abdominal ganglia to

only five.

3. The large, round flat shape of the last segment

of the labial palpi.

4. The shape and site of insertion of the coxae.

APOMORPHIC CHARACTER STATES

The following list of apomorphic character states

was used in detecting the phylogenetic relationships between the genera of Nemouridae as shown

in the Argumentation Scheme (Figure 2). After

each character state I give my reasons for deciding

that it is apomorphic. The numbers refer to those

used in the Argumentation Scheme.

TABLE 1.—Plesiomorphic (primitive) and apomorphic (derived) conditions of some characters of

Nemouridae

Character

Head

submental gills

Thorax

cervical gills

Legs (nymphs)

spines

Wings

veins At and A2

terminal costal crossveins..

Abdomen (male)

vesicle

terga4to8

tergum 9

Tergum 10

sclerotized bars

Male genitalia

epiproct

dorsal sderite ...

base

basal cushion

ventral sderite .

flagellum

lateral knobs

paraprocts

lobes

inner lobes

Female genitalia

pregenital plate

subgenital plate

Cerd

vaginal lobes

posterior margin

mesobasal lobe .

ventrobasal lobe

spines or hooks .

Plesiomorphic Apomorphic

absent

present

simple

absent

irregular

separate

joining R

present

simple

slightly

produced

symmetrical

flat or concave

naked

single

symmetrical

completely

recurved

fused throughout

large

normal

absent

narrow and

keel-shaped

absent

absent

naked

simple

short

two

small lightly

sclerotized

absent

present

lightly

sclerotized

small

straight

short

membranous

absent

absent

absent

present

absent

branched

present

whorls

fused

joining Sc

absent

enlarged or

modified

greatly enlarged

asymmetrical

enlarged and

modified

spines or prongs

double

asymmetrical

directed upward

divided longitudinally

small

dorsal

present

broad and flat

present

present

spines or

prongs

modified

elongate

three or one

large heavily

sclerotized

present

absent

heavily

sclerotized

large

bifurcate

elongate

sclerotized

present

present

present
NUMBER 211 33

At the base of the diagram, the four synapomorphies of the Nemouridae given by Zwick

(1973a) are shown but not numbered.

1. Division of male paraprocts into three lobes.

The plesiomorphic state of the male paraprocts in

Plecoptera is a single, large, lightly sclerotized lobe.

I suggest that the hypothetical stem nemourid had

already developed a small poorly developed inner

lobe like is present in most of the Nemourinae

today. The subsequent development of an outer or

cervical lobe is the apomorphic character state that

separates the Amphinemurine as a subfamily and

unites the genera in the Amphinemurinae as a

monophyletic lineage.

2. Male paraprocts armed with spines or prongs.

The majority of the Plecoptera have paraprocts

naked of spines or prongs. I conclude that spines

or prongs on the middle or outer paraproctal lobes,

found in all genera of Amphinemurinae, is an apomorphic state.

3. Ventral sclerite of epiproct enlarged and with

a very broad base. The plesiomorphic condition is

a thin, sclerotized keel-shaped sclerite that connects

at the base as a narrow sclerotized band. This derived modification unites the Nemourinae as a

monophyletic lineage.

4. Two highly branched cervical gills present,

one on each side of lateral cervical sclerites. The

plesiomorphic state of nemourid gills is the single

fingerlike condition. This synapomorphy unites the

genera Amphinemura and Malenka.

5. Female subgenital plate developed into large

sclerotized median lobe which overlaps vaginal

lobes. The plesiomorphic state in Amphinemurinae is paired vaginal lobes and no median plate.

The development of a large sclerotized median

plate is the synapomorphic condition that combines Indonemoura, Mesonemoura, and Protonemura in a monophyletic lineage.

6. Mesobasal lobe present on dorsolateral margins of male cerci. Most male nemourids have

simple membranous cerci. Formation of a specialized mesobasal lobe is the apomorphic state

that separates Malenka from its sister-genus

Amphinemura.

7. Cervical gill branches all extending to gill

base. Although branched gills represent a derived

character state, most branching is partial and incomplete. Numerous thin branches that all extend

to the gill base is an apomorphic character state

that distinguishes Amphinemura as a distinct

genus.

8. Outer cervical gills with a single deep fork.

The plesiomorphic state is a single, simple gill on

each side of the lateral cervical sclerites. This

forked condition of the outer gill represents an

apomorphic state that serves to separate the genus

Protonemura.

9. Inner cervical gills absent or reduced to small

nubs. The plesiomorphic state is a single, simple

gill on each side of the lateral cervical sclerites.

The loss of the gills inside the lateral cervical sclerites is a synapomorphic condition that unites

Mesonemoura and Indonemoura as a monophyletic

sister-group pair.

10. Male cerci elongate. Most male Nemouridae

have very short cerci. This synapomorphic state

helps unite the genera Mesonemoura and

Indonemoura.

11. Epiproct with large epiproctal flagellum. Although the other genera in this lineage occasionally

have a small epiproctal projection, the remainder

of the Nemouridae are without one. The species of

Mesonemoura all have a large well-developed epiproctal flagellum, and this distinguishes Mesonemoura as a separate genus.

12. Median lobes of male paraprocts greatly enlarged and bearing large apical prongs. The plesiomorphic condition in the Amphinemurinae is

median paraproctal lobes of moderate size with

spines or small prongs. This apomorphic state

separates the genus Indonemoura.

13. Lateral knobs present on base of ventral

sclerite of epiproct. The plesiomorphic condition

of the basolateral corners of the ventral sclerite is a

simple broadening at the junction of the sclerotized portion of the tenth tergum. Modification

into definite hingelike knobs is one synapomorphic

character state that unites the genera Zapada,

Illiesonemoura, Nemoura, and Soyedina.

14. Basal cushion present on dorsal base of epiproct. The lateral arms of the dorsal sclerite bend

inward and form a cavity where the basal cushion

forms. This structure is absent from the rest of the

Nemouridae. I conclude that the presence of this

large membranous cushionlike structure immediately in front of the horseshoe-shaped base of the

dorsal sclerite unites Nemoura and the other

genera in this monophyletic lineage.

15. Ventral sclerite forming entire ventral sur-
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

face of epiproct and extending over much of lateral

and dorsal aspects. The plesiomorphic condition

is a narrow ventral sclerite, which is slightly enlarged at the base and extends upward between the

folds of the dorsal sclerite. This enlarged modified

state unites the genera in this lineage.

(O

c

<u

to

J_

3

E

a)

c

t3

E

<o

J3

O

£E

CD a>

o

•a

• a

<TJ <d

to +->

a . to

s i

O

a.

c o

"o to

0) •!-

<T3 *O

3 O

O +J

E W

OJ O

C J -

rd Q.

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<a

Apomorphic

Plesiomorphic

Ancestral stem species

FIGURE 2.—Argumentation scheme for the hypothetical phytogeny of the family

Nemouridae.
NUMBER 211 35

16. Distinct whorls of femoral spines present on

forelegs of nymphs. Although the nymphs of most

genera have femoral spines, they are seldom arranged in definite fringes or whorls a single spine

wide. This synapormophic state unites Zapada and

Illiesonemoura as sister genera and separates them

from Nemoura and Soyedina.

17. Cervical gills absent. The primitive condition

is the presence of cervical gills. The absence of

cervical gills is an apomorphic condition that

serves as a synapomorphic character state to unite

Nemoura and Soyedina.

18. Whorls of femoral spines present on all

nymphal leg pairs. Since the presence of whorls of

femoral spines has been stated to be apomorphic,

the transformation of this character state to

all leg pairs represents a further apomorphic

modification.

19. Inner cervical gills absent. Gills both inside

and outside of the lateral cervical sclerites is considered the plesiomorphic state. The loss of the

inner gills is an apomorphic state.

20. Outer lobe of male paraprocts with large

inward-directed process near apex. The primitive

state of the male paraprocts in the Nemouridae is

a large unmodified outer lobe and a small inconspicuous inner lobe. I suggest that this addition of

an inward-directed process on the inner margin of

the outer lobe is an apomorphic character state

that serves to separate the genus Illiesonemoura.

21. Male cerci enlarged, sclerotized, and usually

bearing large terminal spines or hooks. The plesiomorphic condition in Nemouridae is for the cerci

to be simple and membranous or lightly sclerotized. The modification of the male cerci into

sclerotized, hook-bearing structures is an apomorphic character state.

22. Male paraprocts with outer lobes elongate

and greatly enlarged. The plesiomorphic state is

simple, angular outer lobes. This modification into

large modified, sclerotized structures is an apomorphic character state of Soyedina.

23. Epiproct asymmetrical. Most Nemouridae

have a bilaterally symmetrical epiproct. I deduce

that an asymmetrical condition produced by the

reduction in size of the left half represents a derived character state.

24. Veins Ax and A2 fused near margin of forewings. The plesiomorphic condition is for veins

A! and A2 to run parallel to the margin of the

forewings. This fusion of the anal veins in the forewing is an apomorphic condition.

25. Dorsal and ventral sclerites separated apically

in a longitudinal plane. The plesiomorphic structure of the epiproct is for the dorsal and ventral

sclerites to be joined throughout the length of the

epiproct. This division of the two sclerites represents a derived condition. This synapomorphic

character state combines Nemurella, Ostrocerca,

Shipsa, and Podmosta in a monophyletic lineage.

26. Male tenth tergum joined to epiproct by two

lateral sclerotized bars. The rest of the Nemouridae

have the epiproct joined to the tenth tergum by a

single, median, sclerotized bar. I suggest that two

sclerotized bars joining the epiproct is an apomorphic character state.

27. Anterior lightly sclerotized portion af dorsal

sclerite greatly reduced or absent. The plesiomorphic state in Nemouridae is for the anterior

portion of the dorsal sclerite to cover the entire

anterodorsal surface. I suggest that this reduction

in the anterior portion of the dorsal sclerite is an

apomorphic character development. It is the synapomorphic state that unites the genera Prostoia,

Paranemoura, Lednia, and Visoka.

28. Male paraprocts with enlarged darkly sclerotized inner lobes. The primitive condition is small,

lightly sclerotized inner lobes. This modification of

the inner lobes of the paraprocts is an apomorphic

character state.

29. Base of dorsal sclerite situated on top of epiproct. Since the primitive condition is for the

horseshoe-shaped base of the dorsal sclerite to be

located at the base of the epiproct, this represents

a derived condition. This apomorphic character

state helps to further unite Nemurella and

Ostrocerca.

30. Laterodistal margins of male tenth tergum

enlarged and bearing hairs or spines. The lateral

margins of the tenth tergum are slightly enlarged

in Nemurella and Ostrocerca, but this is related to

their enlarged, sclerotized cerci. This modification

of the distal margins is an apomorphic character

state that unites Shipsa and Podmosta.

31. Inner lobes of male paraprocts absent. The

primitive condition is a large outer lobe and a

small thin, sclerotized inner lobe. This loss of the

inner lobe helps to combine Shipsa and Podmosta.

32. Male cerci with large ventrobasal lobe. Primitive cerci are simple and unmodified. The cerci of
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ostrocerca are elongate and have sclerotized points

at the apex but are unmodified at the base. I suggest that this formation of a ventrobasal lobe in

Nemurella represents an apomorphic character

state that divides it from Ostrocerca.

33. Male cerci elongate, heavily sclerotized, and

pointed at apex. Modifications of the cerci represent apomorphic states in the Nemouridae. The

highly modified cerci of Ostrocerca are used in

separating it from Nemurella.

34. Male tenth tergum with very large sclerotized terminal projections. The tenth tergum of

most Nemouridae is essentially unmodified. The

sister genus in this branch, Podmosta, also has an

enlarged tenth tergum, but it is only partially modified in this direction. These enlarged, highly modified projections of the tenth tergum are an apomorphic condition in Shipsa.

35. Female seventh and eighth sterna not modified into plates. Most Plecoptera have either the

seventh or eighth sternum modified into a genital

plate. The plesiomorphic condition is for the

eighth sternum to be modified but in the Nemourinae the seventh sternum is usually modified. I submit that a reduction to essentially no genital plate

is a derived condition. This apomorphic character

state unites the species in Podmosta and separates

it from Shipsa.

36. Male paraprocts with large darkly sclerotized

inner lobes. The plesiomorphic character state is

small, lightly sclerotized inner lobes. This modification of the paraprocts is an apomorphic character state that unites Lednia and Visoka as a

monophyletic sister group.

37. Ventral sclerite curving upward laterally and

forming most of epiproct. The plesiomorphic state

is for the ventral sclerite to be in a ventral or comparable position. The dorsal sclerite is also reduced, but this occurs in other genera without a

corresponding modification of the ventral sclerite.

This implied synapomorphic state unites Paranemoura and Prostoia as a monophyletic pair.

38. Lateral margins of male tenth tergum modified as long narrow upward projections. The usual

condition in primitive Plecoptera and most Nemouridae is a broad flat tenth tergum. These distinct spine-bearing projections represent an apomorphic character state that helps to separate the

genus Lednia.

39. Vesicle absent from male ninth sternum.

Most Nemouridae have a vesicle on the hypoproct

or anterior margin of the ninth sternum. The absence of the vesicle represents an apomorphic state

that helps separate Lednia from Visoka.

40. Branched submental gills present. Submental gills are not present in any other genera of

Nemouridae. Submental gills are present in some

other Plecoptera families, but they are usually

simple and attached to the base of the submentum

as in many Perlodidae. I submit that the presence

of branched submental gills, attached between the

labium and submentum, is an apomorphic character state that delineates Visoka as a distinct

genus.

41. Male cerci sclerotized and highly modified.

The plesiomorphic condition of cerci in Nemouridae is simple, membranous, and unmodified. This

modification into short flat cerci with an apical

spine is an apomorphic character state. I submit

that this occurrence of sclerotized cerci with an

apical projection similar to those in Nemoura is a

result of convergence.

42. Terminal costal crossvein of forewings joining Sc. The plesiomorphic state is for the terminal

costal crossvein of the forewings to join R. This

apomorphic character state separates the genus

Paranemoura.

43. Vesicle absent from male ninth sternum. The

primitive condition is for the vesicle to be present

on the ninth sternum of the male. The presence of

this apomorphic condition in a second genus in

this branch of the phylogenetic scheme is explained

as an example of convergence. The genetic basis

for the loss of the vesicle must have been present

in the hypothetical ancestor and has occurred

twice in separate genera.

44. Lateral margins of ventral sclerite fused

dorsally, forming dorsal and ventral surfaces of

epiproct. The plesiomorphic state is with the ventral sclerite forming essentially only the ventral

aspect of the epiproct, although dorsal projections

are often present. This apomorphic character state

distinguishes Prostoia as a separate genus.

Zoogeography

The Nemouridae are distributed throughout

the Northern Hemisphere. The sister family, Notonemouridae, has an extreme Southern Hemisphere

or Gondwanian distribution pattern (lilies, 1965).
NUMBER 211 37

I suggest that the prenemourids or stem nemourids

were present when there was still a distributional

connection between Laurasia and Gondwanaland.

This would make them approximately 120 million

years old (Wilson, 1963).

The family Nemouridae developed on the northern landmass Laurasia. Fossil evidence in the form

of fossilized Plecoptera in Baltic amber shows that

some of the nemourid genera were present in the

lower Oligocene epoch (lilies, 1965). These fossils

indicate that the family Nemouridae is at least 25

million years old. When the family split into subfamilies is not known, but it must have been while

there was still a good connection between North

America and Eurasia, because both subfamilies are

well represented by genera in both the Palearctic

and Nearctic regions.

The Nemouridae radiated throughout the

Northern Hemisphere with evolutionary centers in

south central Asia and western North America.

These centers correspond generally with the Himalayan Mountains and the Rocky Mountains. These

areas presently contain the most extant genera

with seven and ten respectively. They also have

the most endemic genera with three each (Figure

3).

The subfamily Amphinemurinae contains four

genera besides Amphinemura, three are found in

the Oriental and Palearctic regions, and one is endemic to the Nearctic region. Protonemura is

widely distributed in the Old World, but Mesonemoura and Indonemoura are restricted to central Asia. Malenka is only found in western North

America. Amphinemura is the most widely distributed genus in the Nemouridae, it is present

throughout the Holarctic region and has also

radiated southward into the Oriental region.

The subfamily Nemourinae contains eleven genera besides Nemoura and with the exception of

Podmosta, which is present in both North America

and eastern Asia, they are all restricted to either

the Oriental and Palearctic regions or the Nearctic

region. Two genera, Nemurella and Illiesonemoura, are only found on the Eurasian continent, and their sister genera Ostrocerca and Zapada are only found in North America. The six

remaining genera—Lednia, Paranemoura, Prostoia,

Shipsa, Soyedina, and Visoka—are only known from

North America. This does, however, reflect the fact

that more research has been done on the North

American nemourid fauna. When an adequate

study of the Nemouridae of Asia is completed a

FIGURE 3.—Distribution map of the family Nemouridae. Numbers indicate the numbers of

endemic genera occurring within the outlined areas.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

better overall understanding will be possible. The

species complexes in Nemoura—Cercispinosa and

Ovocercia—may quite possibly represent distinct

genera endemic to Asia.

Some genera have very interesting distribution

patterns. Four genera in Nemourinae—Ostrocerca,

Prostoia, Soyedina, and Zapada—have a disjunct

pattern. They are found in extreme western North

America and then again in eastern North America.

They probably once ranged all across northern

North America but are now absent in the central

area. Most eastern species are rare and restricted to

high mountain areas that are known to contain

relict populations of plants and animals.

Podmosta has a similar disjunct distribution pattern in North America but has also been collected

in eastern Asia. Podmosta weberi Ricker is known

from northern Alaska and the Soviet Far East.

This can be best explained by the fact that a land

connection existed between Alaska and eastern

Asia quite recently in geologic time. Nemoura

arctica Esben-Petersen is another species that supports this fact because of its widespread holarctic

distribution. There was much faunal exchange between North America and Asia in the Pleistocene

epoch. This exchange was via the Bering Bridge

which served as a major distributional track in the

sense of Croizat et al. (1974).

Indonemoura is common in the Himalayan

Mountains and then occurs again in the Sunda

Islands. It is highly possible that collecting in

Burma and Thailand will fill in this distribution

gap.

Lednia is only known from two localities in

Glacier National Park, Montana. This area of the

northern Rocky Mountains was collected intensively over a ten-year period by Dr. Arden R.

Gaufin and others while studying the Plecoptera of

Montana (Gaufin et al., 1972), but further records

were not obtained. This simply shows that much

more collecting needs to be done before our knowledge of the Nemouridae will be complete.

It will also be extremely helpful when the species placed in incertae sedis are studied and assigned to their proper places in the phylogenetic

scheme.

Distribution maps are provided for all genera in

Figures 173-186, and detailed distribution data

are given for each species in the "Distribution and

Species List" sections under each genus.

Epilogue

The basis for this revision was to organize the

present knowledge of the family Nemouridae, analyze and define the morphological characters available for taxonomy, and construct a hypothetical

phylogeny for use in further studies. With this accomplished, the foundation has been laid for

detailed studies of the Nemouridae at the specific

level.

I plan to do complete revisions of all the genera

in Nemouridae over the next decade, including the

species incertae sedis. These revisions will include

studies of the internal anatomy and will incorporate further analysis of external features using a

scanning electron microscope. Ecology and dynamic zoogeographic factors will be studied, especially when dealing with the nymphal or aquatic

life stage.

My goal upon completion of the detailed generic

revisions is to complete a special study on the

phylogeny and zoogeography of the family

Nemouridae.

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Joost, W.

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NUMBER 211 41

Kawai, T.

1954. Studies on the Holognathous Stoneflies of Japan, I:

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42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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1943. Stoneflies of Southwestern British Columbia. Indiana University Publications, Science Series, 12:1-

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1974. Die Feinstruktur der Sinneshaare der Bauchblase

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1964. Drei Interessante Arten der ATemoura-Gattung

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NUMBER 211 43

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1974. Feinstruktureller und Histochemischer Nachweis

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1963. Continental Drift. Scientific American, 208:86-100.

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1957. Plecoptera Slovenska (Faunisticko-Systematicka

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1923. Morphology, Anatomy and Ethology of Nemoura.

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1957. Plecoptera from the Altai-Sajan Mountains. Akademia Nauk USSR Siberskii, Novosibirsk, pages 27-

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1961. Hydrobiological and Ichthyological Descriptions of

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115.

Zhiltzova, L. A.

1957. Contribution a l'£tude des Plecopteres du Caucase,

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Entomologicheskoye Obozreniye USSR, 36:659-670.

1958. Contribution a l'£tude des Plecopteres du Caucase,

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1964. On the Knowledge of Plecoptera from the Caucasus, VI: New Species of Taeniopterygidae, Nemouridae and Capniidae. Entomologicheskoye Obozreniye USSR, 43:347-362.

1967. New Genus and Three New Species of Stoneflies

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1971. On the Knowledge of Plecoptera from Middle Asia,

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50:347-365.

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1970. Was ist Nemoura marginata F. J. Pictet 1836? Bestimmung eines Neotypus und Beschreibung einer

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1971. Plecoptera aus Anatolien und Benachbarten Gebieten. Mitteilungen der Schweizerischen Entomologischen Gesellschaft, 44:233-264.

1973a. Insecta: Plecoptera, Phylogenetisches System und

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Berlin: Walter de Gruyter & Company.

1973b. On the Stoneflies from Korea (Insecta, Plecoptera).

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1973c. Plecoptera from Korea. Annales Historico-Naturales

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1974. Zwei Neue Nemouridae (Ins., Plecopteva) aus dem

Fernen Osten. Nouvelle Revue d'Entomologie, 4:75-

78.
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 4-11.—Amphinemura delosa (Ricker): 4, epiproct, dorsal; 5, epiproct, lateral; 6, epiproct,

ventral; 7, male paraproct, right; 8, male terminalia, dorsal; 9, male terminalia, lateral; 10, male

terminalia, ventral; 11, female terminalia, ventral.
NUMBER 211 45

18

19

FIGURES 12-19.—Indonemoura indica (Kimmins): 12, epiproct, dorsal; 13, epiproct, lateral; 14,

epiproct, ventral; 15, male paraproct, right; 16, male terminalia, dorsal; 17, male terminalia,

lateral; 18, male terminalia, ventral; 19, female terminalia, ventral.
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 20-27.—Malenka calijornica (Claassen): 20, epiproct, dorsal; 21, epiproct, lateral; 22,

epiproct, ventral; 23, male paraproct, right; 24, male terminalia, dorsal; 25, male terminalia,

lateral; 26, male terminalia, ventral; 27, female terminalia, ventral.
NUMBER 211 47

28

FIGURES 28-35.—Mesonemoura vaillanti (Navas): 28, epiproct, dorsal; 29, epiproct, lateral; 30,

epiproct, ventral; 31, male paraproct, right; 32, male terminalia, dorsal; 33, male terminalia,

lateral; 34, male terminalia, ventral; 35, female terminalia, ventral.
48

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

39 43

FIGURES 36-43.—Protonemura nitida (Pictet): 36, epiproct, dorsal; 37, epiproct, lateral; 38,

epiproct, ventral; 39, male paraproct, right; 40, male terminalia, dorsal; 41, male terminalia,

lateral; 42, male terminalia, ventral; 43, female terminalia, ventral.
NUMBER 211 49

47

PQP

51

FIGURES 44—51.—Illiesonemoura punctata (Jewett): 44, epiproct, dorsal; 45, epiproct, lateral; 46,

epiproct, ventral; 47, male paraproct, right. Illiesonemoura cordata (Jewett): 48, male terminalia, dorsal; 49, male terminalia, lateral; 50, male terminalia, ventral. Illiesonemoura punctata:

51, female terminalia, ventral.
50

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

vs

55

1 . 0 mm

sgp

pgp

FIGURES 52-59.—Lednia tumana (Ricker): 52, epiproct, dorsal; 53, epiproct, lateral; 54, epiproct,

ventral; 55, male paraproct, right; 56, male terminalia, dorsal; 57, male terminalia, lateral; 58,

male terminalia, ventral; 59, female terminalia, ventral.
NUMR FR

Ik

51

65

FIGURES 60-67.—Nemoura trispinosa Claassen: 60, epiproct, dorsal; 61, cpiproct, lateral; 62,

epiproct, ventral; 63, male paraproct, right; 64, male terminalia, dorsal; 65, male terminalia,

lateral; 66, male terminalia, ventral; 67, female terminalia, ventral.
52

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

la

71

75

FIGURES 68-75.—Nemurella pictetii Klapalek: 68, epiproct, dorsal; 69, epiproct, lateral; 70,

epiproct, ventral; 71, male paraproct, right; 72, male terminalia, dorsal; 73, male terminalia,

lateral; 74, male terminalia, ventral; 75, female terminalia, ventral.
NUMBER 211

79

81

83

FIGURES 76-83.—Ostrocerca truncata (Claassen): 76, epiproct, ventral; 77, epiproct, lateral; 78,

epiproct, dorsal; 79, male paraproct, right; 80, male terminalia, dorsal; 81, male terminalia,

lateral; 82, male terminalia, ventral; 83, female terminalia, ventral.
54

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

vs

86

91

FIGURES 84-91.—Paranemoura perfecta (Walker): 84, epiproct, dorsal; 85, epiproct, lateral; 86,

epiproct, ventral; 87, male paraproct, right; 88, male terminalia, dorsal; 89, male terminalia,

lateral; 90, male terminalia, ventral; 91, female terminalia, ventral.
NUMBER 211 55

la

95

FIGURES 92-99.—Podmosta delicatula (Claassen): 92, cpiproct, dorsal; 93, epiproct, lateral; 94,

epiproct, ventral; 95, male paraproct, right; 96, male terminalia, dorsal; 97, male terminalia,

lateral; 98, male terminalia, ventral; 99, female terminalia, ventral.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

vs

105

102

103

FIGURES 100-107.—Prostoia besametsa (Ricker): 100, epiproct, dorsal; 101, epiproct, lateral; 102,

epiproct, ventral; 103, male paraproct, right; 104, male terminalia, dorsal; 105, male terminalia,

lateral; 106, male terminalia, ventral; 107, female terminalia, ventral.
NUMBER 211

57

113

110

"\

FIGURES 108-115.—Shipsa rotunda (Claassen): 108, epiproct, dorsal; 109, epiproct, lateral; 110,

epiproct, ventral; 111, male paraproct, right; 112, male teminalia, dorsal; 113, male terminalia,

lateral; 114, male terminalia, ventral; 115, female terminalia, ventral.
58

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

121

122

FIGURES 116-123.—Soyedina producta (Claassen): 116, epiproct, ventral; 117, epiproct, lateral;

118, epiproct, dorsal; 119, male paraproct, right; 120, male terminalia, dorsal; 121, male terminalia, lateral; 122, male terminalia, ventral; 123, female terminalia, ventral.
NUMBER 211 59

vs

127

FIGURES 124-131.—Visoka cataractae (Neave): 124, epiproct, dorsal; 125, epiproct, lateral; 126,

epiproct, ventral; 127, male paraproct, right; 128, male terminalia, dorsal; 129, male terminalia,

lateral; 130, male terminalia, ventral; 131, female terminalia, ventral.
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

132

135 J 139

FIGURES 132-139.—Zapada oregonensis (Claassen): 132, epiproct, dorsal; 133, epiproct, lateral;

134, epiproct, ventral; 135, male paraproct, right; 136, male terminalia, dorsal; 137, male terminalia, lateral; 138, male terminalia, ventral; 139, female terminalia, ventral.
NUMBER 211

1 . 0 mm 1 . 0 mm

61

140 141

. 1. 0 mm 1 . 0 mm

142 143

FIGURES 140-143.—Cervical region of nymphs: 140, Amphinemura delosa (Ricker); 141, Malenka

californica (Claassen); 142, Mesonemoura vaillanti (Navas); 143, Protonemura sp.
62

1 . 0 mm

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

i 1. 0 mm ,

144 145

1 . 0 mm 1 . 0 mm

146 147

FIGURES 144-147.—Cervical region of nymphs: 144, Illiesonemoura sp.; 145, Nemoura cinerea

(ReUius); 146, Visoka cataractae (Neave); 147, Zapada hay si (Ricker).
NUMBER 2H 63

148

150

151

152

FIGURES 148-152.—Right forewings: 148, Prostoia besametsa (Ricker), female; 149, Shipsa rotunda

(Claassen), female; 150, Zapada columbiana (Claassen), female; 151, Malenka coloradensis

(Banks), female; 152, Soyedina producta (Claassen), female.
64

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

153

154

157

FIGURES 153-157.—Right forewings: 153, Paranemoura perfecta (Walker), male; 154, Nemoura

arctica Esben-Petersen, female; 155, Amphinemura apache Baumann and Gaufin, female; 156,

Illicsonemoura cordata (Jewett), female; 157, Protonemura meyeri (Pictet), female.
NUMBER 211 65

I

o

X

FICURES 158-161.—Right forelegs of nymphs: 158, Malenka californica (Claassen); 159,

Amphinemura wui (Claassen); 160, Protonemura sp.; 161, Mesonemoura vaillanti (Navas).
66

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

164

V^

«fpp^

167

FICURES 162-167.—Right forelegs of nymphs: 162, Ostrocerca truncate (Claassen); 163, Podmosta

decepta (Frison); 164, Prostoia besametsa (Ricker); 165, Paranemoura perfecta (Walker); 166,

Shipsa rotunda (Claassen); 167, Nemurella pictetii Klapalek.
NUMBER 211 67

FIGURES 168-172.—Right forelegs of nymphs: 168, Visoka cataractae (Neave); 169, Illiesonemoura

sp.; 170, Soyedina vallicularia (Wu); 171, Nemoura cinerea (Retzius); 172, Zapada haysi (Ricker).
68

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

60 180

180 IBO 140 120 100

FIGURE 173.—Distribution map of Amphinemura.

*0 120 IQO 6 0 60 4,0 20 0 2 0 100 120 14-0 160 180

FIGURE 174.—Distribution map of Malenka (1) and Indonemoura (2).
NUMBER 211 69

40 160 ISO

180 160 1^0 |2Q 100 BO 60 4 0 20 "5o XQ 6 0 S o Too fz o FXo f60 fao

FIGURE 175.—Distribution map of Mesonemoura.

2 ?° 0 20 »0 60 80 JOO 120 _ _ 140 160 ISO

TO i o 60 io Too fio 1*0 i«o ~iso

FIGURE 176.—Distribution map of Protonemura.
70

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

160 180

ISO 160 140 120 100 80 60 »0 20 25 «0 60 80 TOO flo Tto ltd 180

FIGURE 177.—Distribution map of Lednia (1) and llliesonemoura (2).

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72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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74 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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