WILLIAM L. PETE A Revision of the

V Leptophlebiidae of the

¥ West Indies

(Ephemeroptera)

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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 62

William L. Peters A Revision of the

Leptophlebiidae of the

West Indies

(Ephemeroptera)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON
ABSTRACT

Peters, William L. A. Revision of the Leptophlebiidae of the West Indies

(Ephemeroptera). Smithsonian Contributions to Zoology, 62:1-48, 1971.—This

study is a revision of the West Indian Leptophlebiidae and is based on reared or

associated nymphs and adults. Three new genera, Farrodes, Careospina, and

Traverina are established; the genera Neohagenulus, Hagenulus, and Borinquena

are considered valid. Two subgenera, Borinquena sensu stricto and Australphlebia, are established for Borinquena. Eleven new species are described, Farrodes hyalinus, F. grenadae, F. bimaculatus, Careospina hespera, C. minuta, C.

annulata, Hagenulus jamaicensis, H. morrisonae, H. rangelae, Traverina cubensis, and Borinquena (A.) traverae. The phylogeny and relationships of the West

Indian genera are discussed.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

UNITED STATES GOVERNMENT PRINTING OFFICE

WASHINGTON : 1971

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington, D.C. 20402 - Price 55 cents
William L. Peters A Revision of the

Leptophlebiidae of the

West Indies

(Ephemeroptera)

Introduction

Sufficient collections of Leptophlebiidae are not

available from most parts of the world to contemplate species revisions of a genus or group of related genera. Especially, reared or associated

nymphs and adults are not available and are difficult to obtain. This study is rather unique as a

good quantity of leptophlebiid material is available

from the West Indies, including many associated or

reared nymphs and adults. While undoubtedly

more species of the West Indian Leptophlebiidae

are yet to be found and described, the present study

does provide an addition to the species revision of

the Neotropical Leptophlebiidae.

The term "West Indies" as used herein includes

all islands in the Caribbean Archipelago extending

from Florida in North America and Yucatan in

Central America to Venezuela in South America.

The term "Greater Antilles" includes Cuba, Hispaniola (Haiti and Dominican Republic), Jamaica,

and Puerto Rico, and the term "Lesser Antilles"

consists of an irregular chain of islands including

Trinidad between the Greater Antilles and South

America. Although the Bahamas are usually included as the third group of islands in the West

Indies, no mayflies were available from the

Bahamas for study.

Traver (1938) published the only detailed study

of the Leptophlebiidae from any part of the West

Indies. From Puerto Rico she established two new

genera of the Leptophlebiidae—Neohagenulus with

three new species and Borinquena with two new

William L. Peters, Entomology, Florida Agricultural and

Mechanical University, Tallahassee, Florida 32307.

species. The only other known leptophlebiid genus

from the West Indies is Hagenulus, which was

originally established by Eaton (1882) from female

imagos and a male subimago of a new species collected in Cuba. Morrison (1919) questionably described the nymphs of Hagenulus from Cuba and

later Traver (1938) discussed the possibility that

these nymphs were not Hagenulus. Until recently

Hagenulus was considered a wide-ranging genus

occurring in the West Indies and the Eastern

Hemisphere. As pointed out herein, Hagenulus is

restricted to the Neotropical region.

The following terms and procedures used in the

generic descriptions of the imagos and nymphs require further explanation. The lengths of the body

and fore wings of the male and female imagos are

given as the total observed variation within a genus.

Venational terminology used is as indicated in Figures 4 and 5. Each segment of the fore legs of the

male imagos is compared to the length of the fore

tibiae and expressed as a ratio; the average length

in millimeters of the fore tibiae is given in parentheses. In the figures of the labia of the genera, the

ventral surface is shown on the right-hand side of

the drawing, and the dorsal surface is shown on the

left.

The research on which this report is based was

supported by grants from the National Science

Foundation to the University of Utah, Salt Lake

City, and Florida Agricultural and Mechanical

University, Tallahassee, William L. Peters, Principal Investigator. Specimens collected by the author in Jamaica were from a field trip supported by

a Grant-in-Aid of Research from the Society of the

Sigma Xi.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Acknowledgments

I offer my sincere appreciation to Dr. Thomas H.

Farr, Science Museum, Institute of Jamaica, Kingston, who greatly encouraged my work on this

paper, and to Dr. C. Bernard Lewis, Director, Institute of Jamaica, and Dr. Farr for making my

collecting trip to Jamaica a successful and enjoyable experience. Special thanks are given to Dr.

P. Alayo D., Institute of Biology, Academy of

Sciences of the Republic of Cuba, Habana, for

permission to study his excellent collection of

Cuban Leptophlebiidae and for writing the Spanish translations included herein. Further, Dr. Alayo

wrote ecological notes on several species described

in this paper.

I thank Dr. George F. Edmunds, Jr., University

of Utah, for permitting me to use his collections

and facilities during a part of this study and for

information on types at the Museum of Comparative Zoology. I wish to thank Dr. Jay R. Traver,

Amherst, Massachusetts, for continuous help during this study and for permission to republish some

of her illustrations. The completion of this work

was dependent upon the following persons for the

loan or gift of specimens: Dr. Alayo; Dr. P. J. Darlington, Jr., and Dr. Howard E. Evans, Museum of

Comparative Zoology, Cambridge; Dr. Thomas H.

Farr; Dr. Oliver S. Flint, Jr., National Museum of

Natural History; Dr. V. Landa, Institute of Entomology, Czechoslovakian Academy of Science,

Prague; Dr. L. L. Pechuman, Cornell University,

Ithaca; and Dr. Jay R. Traver. I wish to thank Drs.

Traver, Edmunds, Alayo, Farr, and Flint for critical reading of part or all of the manuscript. Thanks

are expressed to Mrs. William L. Peters and Mr.

Steven L. Jensen, University of Utah, for preparation of the illustrations under my supervision.

TABLE 1.—Verification of Mature Nymphs

Genus

Farrodes

Traverina

Careospina

Neohagenulus ....

Hagenulus

Borinquena

B. (Borinquena) ..

B.(Australphlebia)

Distribution

Jamaica,

Cuba,

Grenada

Cuba

Cuba

Puerto Rico

Cuba, Haiti,

Jamaica,

Puerto Rico

Puerto Rico,

Dominica,

St. Lucia

Puerto Rico

Dominica,

St. Lucia

Denticles

on

labrum

5 small,

= sized

4-5 small,

= sized

4-5 small,

= sized

4-5 small,

= sized

3-5 small.

= sized

2 large,

= sized &

1-3 small,

T^ sized

Maxillary palpi Labial palpi

Length Length Length Length

segment segment segment segment

2 to 1 3 to 2 2 to 1 3 to 2

!% MrM>

 J% MrM>

1% % little %-#

longer

114 % little yt

longer

— 1/2-% little %~Vi

longer

-!%

= or % little %

11&-2 longer

-less

than 2

1%-1% Vs^k little %-%

longer

Denticles

on claws

Progressively

larger apically,

apical denticle

larger

Progressively

larger

apically

Progressively

larger

apically

Progressively

larger

apically

Progressively

larger

apically

Progressively

larger apically,

apical denticle

larger

Abdominal

gills

Long & slender

(2 lamellae)

Platelike,

terminated with

3 filaments

(2 lamellae)

Long & slender

(deeply

forked)

Long & slender

(deeply

forked)

Long & slender

(deeply

forked)

Long & slender

(1 lamella or

deeply

forked

Long & slender

(1 lamella)

Long & slender

(deeply forked)

Posterolateral

spines on

abdominal

segments

8-9

6-9

5 or 6-9

3-9

5-9

6 or 7-9
NUMBER 62

Keys to the Genera of the Leptophlebiidae from the West Indies

IMAGOS

1. Apical two-thirds of penes of male genitalia divided, a ventral appendage arising from

apex of each penis lobe (Figures 73-78); ninth sternum of female entire (Figure 104).

Farrodes, new genus

Penes of male genitalia not as above (Figures 79-96); ninth sternum of female cleft (Figures

105-110) 2

2. Terminal filament greatly reduced, with 6-7 segments (Figure 121); penes of male genitalia

without subapical, ventral spines (Figures 80-83) Careospina, new genus

Terminal filament longer than cerci; penes of male genitalia with subapical, ventral spines

(Figures 79, 84-87, 89-96), except for Hagenulus morrisonae (Figure 88) 3

3. Penes of male genitalia divided, basal half of penis lobes connected by a sclerotized membrane (Figures 84-86); apex of costal projection of hind wings extending past apex of

wings (Figures 34-35, 37-42) Neohagenulus

Penes of male genitalia divided, no sclerotized membrane connecting penis lobes (Figures

79, 87-96); costal projection of hind wings, if present, not extending past apex of wings

(Figures 16-17, 19-20, 44-45, 47-48, 50-51, 53-54, 56-57, 59-60, 62-63, 65-66, 68-71) ... . 4

4. Apex of hind wings broad and angularly truncated (Figures 16-17, 19-20); male genitalia

as in Figure 79; female with a short ovipositor or egg guide extending to anterior margin

of abdominal segment 8 (Figure 111) Traverina, new genus

Apex of hind wings, if present, rounded (Figures 44-45, 47-48, 50-51, 53-54, 56-57, 59-60,

62-63, 65-66, 68-71); male genitalia as in Figures 87-96; female with a well-developed

ovipositor or egg guide extending to at least anterior margin of abdominal segment 10

(Figures 114-120) 5

5. Cross veins in fore wings of female surrounded with dark clouds (Figures 46, 52, 55, 61); few

to many cross veins in fore wings of male surrounded with dark clouds (Figures 43, 49,

64) or fringed with darker color (Figure 58) Hagenulus

Cross veins in fore wings of male and female not surrounded with dark clouds (Figures

67, 72) (Borinquena) 6

6. Hind wings present (Figures 68-71) B. (Borinquena, sensu stricto)

Hind wings absent B. (Australphlebia), new subgenus

MATURE NYMPHS

1. Abdominal gills on segments 1-7 alike, dorsal and ventral lamellae platelike and terminated

in three processes, with median projection longer than laterals (Figure 205).

Traverina, new genus

Abdominal gills not as above (Figures 204, 206-210); each lamella slender or deeply

forked 2

2. Anterior margin of labrum fringed with an even or uneven band of hair (Figures 186,

188, 190) Hagenulus

Two even rows of hair on anterior margin of labrum (Figures 176, 181, 183, 192, 195) ... . S

3. Posterolateral spines on abdominal segments 3-9, spines progressively larger posteriorly.

Neohagenulus

Posterolateral spines on abdominal segments 5 or 6 to 9, spines progressively larger

posteriorly 4

4. Two large, equal-sized denticles and sometimes 1 to 3 small unequal-sized denticles on

anteromedian emargination of labrum (Figures 193-194, 196) (Borinquena) 5

Four to five small, equal-sized denticles on anteromedian emargination of labrum (Figures

177, 182) 6

5. Abdominal gills on segments 1-7 alike, each gill consists of 1 long, slender portion (Figure

209) B. (Borinquena, sensu stricto)

Abdominal gills on segments 1-7 alike, each gill deeply forked (Figure 210).

B. (Australphlebia), new subgenus
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Posterolateral spines on abdominal segments 8 and 9 (Figure 211); denticles on claws progressively larger apically except apical denticle much larger (Figure 197).

Farrodes, new genus

Posterolateral spines on abdominal segments 5 or 6 to 9 (Figure 212); denticles on claws

progressively larger apically (Figure 199) Careospina, new genus

La clave los generos de Leptophlebiidae de las Antillas

IMAGOS

Machos con los penes divididos en sus y3 apicales, y con un apendice ventral naciendo del

apice de cada lobulo penial (Figuras 73-78); hembras con el noveno esternito abdominal

entero (Figura 104) Farrodes, nuevo genero

Machos con los penes en otra forma (Figuras 79-96); hembras con el noveno esternito

abdominal hendido (Figuras 105-110) 2

Filamento terminal muy reducido, con 6-7 segmentos (Figura 121); machos con los penes

desprovistos de espinas ventrales subapicales (Figuras 80-83) . .. Careospina, nuevo genero

Filamento terminal mas largo que los ceros caudales; machos con los penes provistos de

espinas ventrales subapicales (Figuras 79, 84-87, 89-96), excepto en Hagenulus morrisonae

(Figura 88) 3

Machos con los penes divididos, mitad basal de los 16bulos peniales conectada por una

membrana esclerotizada (Figuras 84-86); apice de la proyeccidn costal de las alas

posteriores prolongada mas alia del apice del ala (Figuras 34-35, 37-42). Neohagenulus

Machos con los penes divididos, 16bulos peniales no conectados por una membrana

esclerotizada (Figuras 79, 87-96); la proyecci6n costal de las alas posteriores, si existe, no

sobrepasa el apice del ala (Figuras 16-17, 19-20, 44-45, 47-48, 50-51, 53-54, 56-57, 59-60,

62-63, 65-66, 68-71) 4

Apice de las alas posteriores amplia y angularmente truncado (Figuras 16-17, 19-20);

genitalia de los machos como en la Figura 79; hembras con un corto ovipositor que llega

hasta el margen anterior del octavo segmento abdominal (Figura 111).

Traverina, nuevo gdnero

Apice de las alas posteriores (si dstas existen), redondeado (Figuras 44-45, 47-48, 50-51, 53-54,

56-57, 59-60, 62-63, 65-66, 68-71); genitalia de los machos como en las Figuras 87-96;

hembras con un bien desarrollado ovipositor que llega por lo menos hasta el margen

anterior del decirno segmento abdominal (Figuras 114-120) 5

Hembras con las nervuras transversales de las alas anteriores rodeadas por una manchita

oscura (Figuras 46, 52, 55, 61); machos con pocas o muchas nervuras transversales de las

alas anteriores rodeadas por manchitas oscuras (Figuras 43, 49, 64) a bordeadas por un

color mas oscuro (Figura 58) Hagenulus

Machos y hembras con las nervuras transversales de las alas anteriores no rodeadas por

manchitas oscuras (Figuras 67, 72) (Borinquena) 6

Alas posteriores presentes (Figuras 68-71) B. (Borinquena, sensu stricto)

Alas posteriores ausentes B. (AustralphUbia), nuevo subgenero

NlNFAS MADURAS

Branquias abdominales de los segmentos 1-7 iguales, con sus lamelas dorsal y ventral en

forma de laminas lanceoladas, y terminadas en tres procesos, el del medio mas largo

que los laterales (Figura 205) Traverina, nuevo genero

Branquias abdominales de otra forma (Figuras 204, 206-210); cada larnela muy delgada u

hondamente bifurcada 2

Margen anterior del labro provisto de una franja de pelos pareja o dispareja (Figuras 186,

188, 190) Hagenulus

Margen anterior del labro con dos hileras parejas de pelos (Figuras 176, 181, 183, 192, 195).

S
NUMBER 62

Segmentos abdominales 3-9 con espinas posterolaterales, cstas progresivamente mayores.

Neohagenulus

Segmentos abdominales 5 6 6 hasta el 9 con espinas posterolaterales, &tas progresivamente

mayores 4

Dos denticulos grandes de tamano igual y algunas veces de uno a tres mas, pequenos y

disparejos, en la emarginacidn antero-media del labro (Figuras 193-194, 196).

(Borinquena) 5

De 4 a 5 denticulos pequenos e iguales en la emarginacion antero-media del labro (Figuras

177, 182) 6

Branquias abdominales de los segmentos 1-7 iguales, cada branquia de forma alargada y

delgada, no bifurcada (Figura 209) B. (Borinquena, sensu stricto)

Branquias abdominales de los segmentos 1-7 iguales, cada una hondamente bifurcada

(Figura 210) B. (Australphlebia), nuevo subgenero

Segmentos abdominales 8 y 9 con espinas posterolaterales (Figura 211); denticulos de las

garras progresivamente mayores hacia el apice, excepto el dentkulo apical que es mucho

mayor (Figura 197) Farrodes, nuevo ge'nero

Segmentos abdominales 5 6 6 hasta el 9 con espinas posterolaterales (Figura 212); denticuios

de las garras progresivamente mayores hacia el apice (Figura 199).

Careospina, nuevo ge'nero

Farrodes Peters, new genus

FICURES 6-14, 73-78, 97, 104, 122-124, 139, 148, 157, 166, 176-

177, 197,204,211

IMAGO.—Length of male: body, 4.2-5.5 mm.; fore

wings, 4.0-6.2 mm. Length of female: body, 4.2-5.8

mm.; fore wings, 5.2-5.6 mm. Eyes of male meet on

meson of head, upper portion of eyes on short stalk,

lower portion of eyes two-thirds lengih of upper

portion; eyes of female separated on meson of head

by a length 4 times maximum width of an eye.

Wings (Figures 6-14) : vein Rg of fore wings forked

one-third of distance from base to margin; vein

MA forked more than half of distance from base to

margin, fork asymmetrical; vein MP forked less than

half of distance from base to margin, fork symmetrical; cubital area as in Figures 6, 9, and 12;

cross veins few. Costal projection of hind wings

acute, short, and well developed (Figures 7-8, 10-

11, 13-14), apex located half to more than half the

distance from base; apex of wings broad and angularly truncated (Figures 7-8, 10-11, 13-14) ; cross

veins few. Legs: ratios of segments in male fore legs,

0.45: 1.00 (2.30 mm.): 0.05: 0.36: 0.27: 0.11: 0.07.

Claws of a pair dissimilar, one apically hooked

(Figure 97), other obtuse, padlike. Male genitalia

(Figures 73-78): segments 2 and 3 of forceps short,

basal segment of forceps narrow, strongly bowed;

styliger plate deeply cleft apically and extends posteriorly dorsal to forceps (Figures 73-78); apical

two-thirds of penes divided, a ventral appendage

arising from apex of each penis lobe as in Figures

73-78. Ninth sternum of female entire (Figure

104). Terminal filament longer than cerci.

MATURE NYMPH.—Head prognathous. Antennae

2i/2 times maximum length of head. Mouthparts

(Figures 139, 148, 157, 166, 176-177) : dorsal hair

on labrum as in Figure 176; submedian and anterolateral areas of hair ventrally; 5 equal-sized denticles on anteromedian emargination (Figure 177).

Clypeus as in Figure 176. Left mandible as in Figure 166. Lingua of hypopharynx with welldeveloped lateral processes (Figure 157), anterior

margin deeply cleft; superlingua of hypopharynx

as in Figure 157, with a row of hair along anterior

margin. Segment 2 of maxillary palpi one and onethird length of segment 1; segment 3 one-third to

half the length of segment 2, triangular; a V-shaped

ridge near the ventral, inner anterolateral margin

of maxillae; hair on maxillae as in Figure 148.

Labium as in Figure 139; segment 2 of palpi equal

to one and one-third length of segment 1; segment

3 one-third to half the length of segment 2, triangular; paraglossae ventral to glossae. Fine hair on

anterolateral margins of prothorax only. Legs (Figure 197) : claws apically hooked, denticles on claws

progressively larger apically except apical denticle

much larger. Gills (Figure 204): gills on segments

1-7 alike; each portion of gills long and slender.

Posterolateral spines on abdominal segments 8-9,

spines progressively larger posteriorly. Terminal

filament slightly longer than cerci.

ETYMOLOGY.—Genus is named for Dr. Thomas

H. Farr, Institute of Jamaica, Kingston, who has
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

greatly contributed to and encouraged my work on

the West Indian Leptophlebiidae.

TYPE SPECIES.—Farrodes hyalinus Peters, new

species.

DISCUSSION.—Farrodes, new genus, can be distinguished from other West Indian genera of the

Leptophlebiidae by the following combination of

characters. In the imagos (1) a ventral appendage

arises from the apex of each penis lobe of the males

(Figures 73-78), (2) the base of vein IC^ of the

fore wings does not connect with vein CuP (Figures

6, 9, 12), and (3) the hind wings are well developed

and each has a short, well-developed costal projection (Figures 7-8, 10-11, 13-14). In the nymphs

(1) the denticles on the claws are progressively

larger apically, except the apical denticle is much

larger (Figure 197), (2) 5 equal-sized denticles are

present on the anteromedian emargination of the

labrum (Figure 177), and (3) posterolateral spines

are present on abdominal segments 8-9 (Figure

211). Farrodes can be distinguished from all genera of the Leptophlebiidae by the following combination of characters. In the imagos (1) the fork

of vein MP of the fore wings is symmetrical (Figures 6, 9, 12), (2) the base of vein ICii! of the fore

wings does not attach to vein CuA or CuP (Figures

6, 9, 12), (3) the tarsal claws of a pair are dissimilar, one is apically hooked while the other is

obtuse, padlike (Figure 97), and (4) the styliger

plate of the male imagos is deeply cleft apically and

extends posteriorly dorsal to the forceps (Figures

73-78). In the nymphs (1) abdominal gills on

segments 1-7 are alike and each portion of the gills

is long and slender (Figure 204), (2) the denticles

on the claws are progressively larger apically except the apical denticle is much larger (Figure

197), (3) 5 equal-sized denticles are present on the

anteromedian emargination of the labrum (Figure

177), and (4) a V-shaped ridge is present near the

ventral, inner anterolateral margin of the maxillae

(Figure 148).

Key to the Species of Farrodes Peters, new genus

IMAGOS

1. Thorax of male buff, sutures lighter; a large, longitudinal, dark brown band on pleura of

thorax of female subimago; ventral appendages on penes of male as in Figures 75-76;

Grenada F. grenadae, new species

Thorax of male brown, sutures lighter; no darker bands on pleura of thorax of female;

appendages on penes of male as in Figures 73-74, 77-78 2

2. Submedian, dark brown maculae near anterior margin on terga 2-6 of male; femora of legs

of female uniformly pale; ventral appendages on penes of male as in Figures 77-78; Cuba.

F. bimaculatus, new species

Terga 2-6 of male without submedian maculae; apex of femora of legs of female darker

brown than remaining portion of femora; ventral appendages on penes of male as in

Figures 73-74; Jamaica F. hyalinus, new species

MATURE NYMPHS

1. Posterior margin of abdominal terga 1-8 with a narrow darker band (Figure 211); Jamaica.

F. hyalinus, new species

Abdominal terga 1-8 uniformly dark brown; Grenada F. grenadae, new species

Clave para las Especies de Farrodes Peters, nuevo genero

IMAGOS

1. Machos con el t6rax de color amarillo pajizo, con las suturas mas claras; subimagos hembras

con una faja grande y longitudinal, pardo-oscura, en las pleuras del t6rax; apendices

vcntrales en los penes del macho como en la Figuras 75-76; Grenada.

F. grenadae, nuevo especie

Machos con el t6rax de color pardo, con las suturas mas claras; hembras sin fajas oscuras

pleurales; apendices en los penes del macho como en las Figuras 73-74, 77-78 2
NUMBER 62

2. Macho con manchas pardo-oscuras submedianas cercanas al margen anterior de los tergitos

2-6; hembra con los femures uniformemente palidos; apendices ventrales en los penes del

macho como en las Figuras 77-78; Cuba F. bimaculatus, nuevo especie

Macho sin manchas submedianas en los tergitos 2-6; hembra con el apice de los femures de

un color pardo mas oscuro que el resto del segmento; apendices venirales en los penes del

macho como en las Figuras 73-74; Jamaica F. hyalinus, nuevo especie

NlNFAS MADURAS

1. Margen posterior de los tergitos abdominales 1-8 con una estrecha banda oscura; Jamaica.

F. hyalinus, nuevo especie

Tergitos abdominales 1-8 uniformemente pardo-oscuros; Grenada.

F. grenadae, nuevo especie

Farrodes hyalinus Peters, new species

FICURES 6-8, 73-74, 97, 104, 139, 148, 157, 166, 176-177, 197,

204, 211

MALE IMAGO (in alcohol).—Length: body, 4.4-

5.0 mm.; fore wings, 4.6-5.0 mm. Upper portion of

eyes and stalks brown, lower portion black. Head

light brown. Base of ocelli black, remainder white.

Thorax and coxae of legs brown, sutures paler.

Prothoracic and mesothoracic legs pale, apex of

femora light brown; metathoracic legs pale, apex

of femora brown. Wings (Figures 6-8): longitudinal veins of fore and hind wings light brown, cross

veins paler; membrane of fore and hind wings

hyaline, except distal third of cells C and Sc of fore

wings translucent, and base of wings brown. Abdomen: segments 1-6 hyaline washed lightly with

brown, posterior margin of terga 1-6 with a narrow

black band, terga 3-6 with a narrow, black, longitudinal streak; spiracles and tracheae black on segments 1-8; segments 7-9 opaque, terga 7-9 washed

with dark brown, sterna 7-9 paler. Genitalia (Figures 73-74) : basal half of forceps segment 1 brown,

remainder of forceps and penes paler. Caudal filaments pale.

FEMALE IMAGO (in alcohol).—Length: body, 4.2-

4.4 mm.; fore wings, 5.2-5.6 mm. Eyes black. Head

brown. Color of thorax and legs as in male imago,

except apex of metathoracic femora light brown.

Wings: longitudinal and cross veins of fore and

hind wings brown, membrane hyaline. Abdomen:

brown, posterior margin of segments 1-8, spiracles

and tracheae darker. Caudal filaments light brown.

MATURE NYMPH (in alcohol).—Head brown,

venter paler. Thorax brown, sterna paler, lateral

margins of terga black. Legs pale, markings as in

male and female imagos. Abdomen brown, sterna

paler, markings as in Figure 211. Gills gray, tracheae

black. Caudal filaments pale.

Holotype male imago, Jamaica: Trelawny Parish,

Martha Brae River, Good Hope, 23-24 September

1963; allotype female imago: same data as for holotypes; paratopotypes: 33 male imagos, 16 female

imagos, 49 male subimagos, 38 female subimagos,

20 nymphs. All types were collected by W. L. Peters

and T. H. Farr and are in alcohol. Four male imaginal, two female imaginal, six male subimaginal,

five female subimaginal, and four nymphal paratopotypes were deposited in each of the collections

of National Museum of Natural History, University

of Utah, and Institute of Jamaica. Holotype, allotype, and remaining paratopotypes have been

deposited in the collections of Florida A & M

University.

ETYMOLOGY.—Latin hyalinus, meaning transparent.

DISCUSSION.—Farrodes hyalinus can be distinguished from the other two described species of

Farrodes by the following characters. In the imagos

(1) the shape of the ventral appendages on the

penes of the male imagos is as given in Figures 73-

74, (2) the basal half of forceps segment 1 of the

male genitalia is brown, remainder of forceps is

paler, and (3) the color pattern on the abdominal

terga of the male and female imagos is as given in

the descriptions. In the nymphs (1) a narrow dark

brown band is present on the posterior margin of

abdominal terga 1-8 (Figure 211), and (2) the

apex of the femora of the prothoracic and mesothoracic legs is dark brown. The nymphs and adults

of F. hyalinus are associated by rearing.

Mature nymphs are found among submerged

plants near the banks of the river. The nymphs of

F. hyalinus emerge at dusk. Subimagos molt at sun-
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

rise and mating occurs in the morning in full sunlight.

Farrodes grenadae Peters, new species

FIGURES 9-11, 75-76, 122-123

MALE IMAGO (in alcohol).—Length: body, 4.2

mm.; fore wings, 4.0-4.2 mm. Upper portion of

eyes, stalks, and head buff, lower portion of eyes

black. Base of ocelli black, remainder white. Thorax

and coxae of legs buff, sutures paler. Remainder of

legs pale, except apex of metathoracic femora

brown, and a brown macula on outer surface of

metathoracic femora. Wings (Figures 9-11): longitudinal veins of fore and hind wings buff, cross

veins paler; membrane of fore and hind wings

hyaline, except distal third of cells C and Sc of fore

wings translucent. Abdomen: segments 1-7 hyaline,

terga 1-7 washed with dark brown as in Figures

122-123, remainder of terga 1-7 and sterna 1-7

washed with light buff; spiracles and tracheae dark

brown on segments 1-8; segments 8 and 9 opaque,

terga 8 and 9 uniformly washed with buff, sterna

8 and 9 paler. Genitalia (Figures 75-76): buff.

Caudal filaments pale.

FEMALE IMAGO.—Unknown.

FEMALE SUBIMAGO (in alcohol).—Length: body,

3.8-4.5 mm.; fore wings, 4.5-4.7 mm. Eyes black.

Head light buff. Thorax buff, sutures, sterna, and

prothorax paler; submedian, oblique, dark brown

bands on notum of prothorax, lateral margins of

prothoracic notum dark brown; a large, longitudinal dark brown band on pleura of thorax. Legs:

buff, color markings as in male imago, except the

brown macula on outer surface of metathoracic

femora smaller. Wings: color as in male imago.

Abdomen: buff, terga 1-9 washed with dark brown

as terga 1—7 of male imago. Caudal filaments pale.

MATURE NYMPH (in alcohol).—Head brown,

venter paler. Thorax brown, venter paler, lateral

margins of prothoracic notum darker. Legs pale,

color markings as in male imago. Abdomen brown,

sterna paler. Gills gray, tracheae black. Caudal filaments pale.

Holotype male imago, Grenada: Balthazar, 7

August 1963; paratopotypes: 10 male imagos, 107

male subimagos, 89 female subimagos; paratypes: 1

nymph, Grenada: Great River, Balthazar, 8 August

1963; 1 male imago, 6 male subimagos, 2 female

subimagos, Grenada: 2 miles east of Lago Grand

Etang, 8 August 1963. All types were collected by

O. S. Flint, Jr., and are in alcohol. Three male

imaginal, 20 male subimaginal, and 15 female subimaginal paratopotypes were deposited in each of

the collections of Florida A & M University and the

University of Utah; the remaining types were deposited in the collections of the National Museum

of Natural History.

ETYMOLOGY.—Species is named for the island of

Grenada.

DISCUSSION.—Farrodes grenadae can be distinguished from the other two described species of

Farrodes by the following characters. In the imagos

(1) the shape of the ventral appendages on the

penes of the male imagos is as given in Figures 75-

76, (2) forceps segment 1 of the male genitalia is

entirely buff, and (3) the color pattern on the abdominal terga of the male and female imagos is as

given in the descriptions and Figures 122-123. In

the nymphs (1) abdominal terga 1-8 are uniformly

brown, and (2) the prothoracic and mesothoracic

legs are entirely pale. The nymphs and adults of

F. grenadae are associated by the developing wing

pads and color markings on the metathoracic legs

of the nymphs collected from the same locality as

the adults.

Farrodes bimaculatus Peters and Alayo, new

species

FICURES 12-14, 77-78, 124

MALE IMAGO (in alcohol).—Length: body, 5.5

mm.; fore wings, 6.2 mm. Upper portion of eyes

and stalks brown, lower portion black. Head brown,

base of ocelli brown, remainder white. Thorax and

coxae of legs brown, sutures paler, carinae darker.

Prothoracic, mesothoracic, and metathoracic legs

pale. Wings (Figures 12-14) : longitudinal veins

and membrane of fore and hind wings hyaline.

Abdomen: tergum 1 hyaline, lateral margins

washed with dark brown, terga 2-6 hyaline, posterior margin with a narrow, transverse, dark brown

band, submedian, dark brown maculae near anterior margin on each segment, spiracles and tracheae

dark brown, tergum 7 light brown, darker brown

markings similar to those of female as in Figure

124, terga 8 and 9 dark brown; sterna 1-7 hyaline,

sterna 8 and 9 dark brown. Genitalia (Figures 77-
NUMBER 62

78): forceps segment 1 brown, remainder of forceps

and penes paler. Caudal filaments pale.

FEMALE IMAGO (in alcohol).—Length: body, 5.8

mm.; fore wings, 5.6 mm. Eyes gray. Head light

brown. Thorax light brown, sutures paler, carinae

darker on pleura. Legs uniformly pale. Wings:

longitudinal veins of fore and hind wings light

brown, cross veins paler; membrane of fore and

hind wings hyaline, except distal third of cells C

and Sc of fore wings translucent. Abdomen: terga

light brown, terga 2-7 with darker color markings

as in Figure 124; sterna light brown. Caudal filaments broken off and missing.

MATURE NYMPH.—Unknown.

Holotypes male imago, Cuba: La Habana Prov.,

Giiines (field no. 43), March 1966; allotype female

imago, Cuba: La Habana Prov., Giiines (field no.

42), March 1966; paratopotypes: 3 male imagos. All

types were collected by P. Alayo D. and are in alcohol. Holotype, allotype, and 1 paratopotype were

deposited in the collection of the Institute of Biology, Academy of Science of the Republic of Cuba.

Remaining paratopotypes were deposited in the

collections of Florida A & M University.

ETYMOLOGY.—Latin bis, meaning two; Latin

macula, meaning spot.

DISCUSSION.—Farrodes bimaculatus can be distinguished from the other two described species of

Farrodes by the following characters. In the imagos

(1) the shape of the ventral appendages on the

penes of the male imagos is as given in Figures 77-

78, (2) forceps segment 1 of the male genitalia is

uniformly brown, and (3) the color pattern on the

abdominal terga of the male and female imagos is

as given in the descriptions and Figure 124.

Traverina Peters, new genus

FICURES 15-20, 79, 98, 105, 111, 125-126, 140, 149, 158, 167-

168, 178-180, 198, 205

IMAGO.—Length of male: body, 6.4 mm.; fore

wings, 6.3 mm. Length of female: body, 6.1 mm.;

fore wings, 6.9 mm. Eyes of male meet on meson of

head, upper portion of eyes on short stalk, lower

portion of eyes three-fourths length of upper portion; eyes of female separated on meson of head by

a length 3 times maximum width of an eye. Fore

wings of male (Figure 15): vein Rs of fore wings

forked one-fourth of distance from base to margin;

vein MA forked half of distance from base to margin, fork asymmetrical, distal portion of vein MA

sagged posteriorly; vein MP forked less than onethird of distance from base to margin, fork symmetrical; cubital area as in Figure 15; cross veins

few. Fore wings of female (Figure 18) : vein Rg of

fore wings forked less than one-fifth of distance

from base to margin; vein MA forked less than half

of distance from base to margin, fork symmetrical;

vein MP forked one-fifth of distance from base to

margin, fork symmetrical; fork of veins MA and

MP narrower than those forks in male wings; cubital area as in Figure 18; cross veins few. Costal

projection of hind wings of male and female acute,

short, and well developed (Figures 16-17, 19-20),

apex of projection located half of distance from

base; apex of wings broad and angularly truncated

(Figures 16-17, 19-20); cross veins few. Legs: ratios

of segments in male fore legs, 0.56: 1.00 (2.5 mm.) :

0.07: 0.36: 0.22: 0.16: 0.09. Claws of a pair dissimilar, one apically hooked (Figure 98), other

obtuse, padlike. Male genitalia (Figure 79): segments 2 and 3 of forceps short; basal segment of

forceps bowed, base of forceps broad, its inner margin forming an angular bend; length of styliger

plate along median line one-third length of maximum width; penes divided except at base, divided

portions flared laterally and tubular, a subapical,

ventral spine on each lobe. Ninth sternum of female deeply cleft (Figure 105). Female with a short

ovipositor or egg guide extending to anterior margin of segment 8 (Figure 111). Terminal filament

longer than cerci.

MATURE NYMPH.—Head prognathous. Antennae

3 times maximum length of head. Mouthparts

(Figures 140, 149, 158, 167-168, 178-180): dorsal

hair on labrum as in Figure 178; submedian and

anterolateral areas of hair ventrally; 4 to 5 small,

equal-sized denticles on anteromedian emargination (Figures 179-180). Clypeus as in Figure 178.

Left mandible as in Figures 167-168. Lingua of

hypopharynx with well-developed lateral processes

(Figure 158), anterior margin deeply cleft; superlingua of hypopharynx as in Figure 158, with a row

of hair along anterior margin. Segment 2 of maxillary palpi one and one-third length of segment 1;

segment 3 half the length of segment 2, triangular;

a V-shaped ridge near the ventral, inner anterolateral margin of maxillae; hair on maxillae as in
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Figure 149. Labium as in Figure 140; segment 2 of

palpi a little longer than length of segment 1; segment 3 a little less than one-fourth to one-fourth

length of segment 2, triangular; paraglossae ventral

to glossae. Fine hair on anterolateral margins of

prothorax only. Legs (Figure 198) : claws apically

hooked, denticles on claws progressively larger

apically. Gills (Figure 205) : gills on segments 1-7

alike; dorsal and ventral lamellae platelike and

terminated in 3 processes, with median projection

longer than laterals (Figure 205). Posterolateral

spines on abdominal segments 6-9, spines progressively larger posteriorly. Terminal filament longer

than cerci.

ETYMOLOGY.—Genus named for Dr. Jay R.

Traver, Amherst, Massachusetts, in honor of her

significant contribution in the study of Ephemeroptera.

TYPE-SPECIES.—Traverina cubensis Peters and

Alayo, new species.

DISCUSSION.—Traverina, new genus, can be distinguished from other West Indian genera of the

Leptophlebiidae by the following combination of

characters. In the imago (1) the costal projection

of the hind wings is acute, short, and well developed

(Figures 16-17, 19-20), (2) the penes of the male

genitalia are divided except at the base and the

divided portions are flared laterally (Figure 79),

and (3) the females possess a short ovipositor or egg

guide extending to the anterior margin of segment

8 (Figure 111). In the nymphs (1) the dorsal and

ventral lamellae of abdominal gills 1-7 are platelike and terminated in 3 processes, with median

projection longer than the laterals (Figure 205),

(2) posterolateral spines are present on abdominal

segments 6-9, and (3) the claws are apically hooked

and the denticles on the claws are progressively

larger apically (Figure 198). Traverina can be distinguished from all genera of the Leptophlebiidae

by the following combination of characters. In the

imagos (1) a subapical, ventral spine is present

on each penis lobe of the male genitalia (Figure

79), (2) the base of vein ICuj of the fore wings is

attached to vein CuP (Figures 15, 18), (3) the

females possess a short ovipositor or egg guide extending to the anterior margin of segment 8 (Figure 111), and (4) the tarsal claws of a pair are

dissimilar, one is apically hooked while the other

is obtuse, padlike (Figure 98). In the nymphs (1)

the dorsal and ventral lamellae of abdominal gills

1-7 are platelike and terminate in 3 processes, with

median projections longer than the laterals (Figure

205), (2) 4-5 equal-sized denticles are present on

the anteromedian emargination of the labrum (Figures 179-180), (3) the claws are apically hooked

and the denticles on the claws are progressively

larger apically (Figure 198), and (4) a V-shaped

ridge is present near the ventral, inner anterolateral

margin of the maxillae (Figure 149).

Traverina cubensis Peters and Alayo, new species

FICURES 15-20, 79, 98, 105, 111, 125-126, 167, 178-179, 205

MALE IMAGO (in alcohol).—Length: body, 6.4

mm.; fore wings, 6.3 mm. Upper portion and stalks

of eyes dark brown, lower portion black. Head light

brown, anterior margin darker. Basal half of ocelli

dark brown, apical half white. Thorax brown, sutures paler, pleura with dark brown markings, especially in diagonal streaks posteromedially from

leg bases. Legs light brown; femora and tibiae of

prothoracic legs washed lightly with dark brown,

large transverse darker brown, median and apical

maculae on femora; large transverse dark brown,

median and apical maculae on femora of mesothoracic and metathoracic legs, median macula

lighter on dorsal surface of femora. Wings (Figures

15-17): longitudinal veins of fore wings brown,

cross veins lighter except in cells C, Sc, and Rlf

membrane transparent, washed with light brown;

longitudinal veins of hind wings brown, cross veins

lighter, membrane transparent, washed with light

brown. Abdomen: terga 1-10 light brown, terga 1

and 10 uniformly washed with dark brown, color

markings as in Figures 125-126, markings lighter on

terga 7-9, spiracles and tracheae dark brown; sterna

light brown. Genitalia (Figure 79) : forceps and

penes light brown, basal segment of forceps washed

lightly in darker brown. Caudal filaments brown,

dark brown annulations at articulations, except

alternate annulations lighter.

FEMALE IMAGO (in alcohol).—Length: body, 6.1

mm.; fore wings, 6.9 mm. Eyes black. Head brown,

anterior margin darker. Color and markings of

thorax and legs as in male imago. Abdomen: terga

brown, tergum 1 uniformly washed with darker

brown, posterior margin of tergum 10 darker

brown, terga 2-9 with darker brown markings as in
NUMBER 62 11

Figures 125-126, markings lighter on terga 8 and 9,

spiracles and tracheae darker; sterna light brown.

Caudal filaments broken off and missing.

MATURE NYMPH (in alcohol).—Head brown,

venter paler. Thorax brown, sterna paler, lateral

margins of terga darker. Legs brown, markings as in

male and female imagos. Abdomen: brown, sterna

paler, markings as in male and female imagos. Gills

gray, tracheae black. Caudal filaments light brown.

Holotype male imago, Cuba: Pinar del Rio Prov.,

Rio Cuzco, near Soroa (field no. 57), March 1964;

allotype female imago, Cuba: Pinar del Rio Prov.,

Rfo Cuzco, near Soroa (field no. 25), March 1965;

paratypes: 2 nymphs, Cuba: Pinar del Rio Prov.,

Soroa (field no. 51), June 1965; 2 nymphs, Cuba:

Pinar del Rfo Prov., Rio Cuzco, near Soroa (field

no. 4), March 1965. All types were collected by P.

Alayo D. and are in alcohol. Holotype was deposited

in the collections of the Institute of Biology, Academy of Sciences of the Republic of Cuba. Two paratypes (1 from field no. 51 and 1 from field no. 4)

were deposited in the collections of the University

of Utah. Remaining types were deposited in the

collections of Florida A & M University.

ETYMOLOGY.—Species is named for the island of

Cuba.

DISCUSSION.—The nymphs and adults of T. cubensis are associated by the developing wing pads

and color markings on the legs and abdomen of the

nymphs collected from the same locality as the

adults. We have seen nymphal specimens of Traverina which appear to represent an undescribed

species. The abdominal color markings are entirely

different than those of T. cubensis. The specimens

are recorded from Cuba: Rio de San Antonio, by

Charco Azul, Trinidad Mts., 23-29 March 1939,

J. G. Needham. Four specimens were deposited in

each of the collections of the University of Utah

and Florida A & M University.

Careospina Peters, new genus

FIGURES 21-32, 80-83, 99, 106, 112, 121, 127-128, 141, 150, 159,

169, 181-182, 199, 206, 212

IMAGO.—Length of male: body, 3.2-6.5 mm.; fore

wings, 3.8-6.5 mm. Length of female: body, 5.1

mm.; fore wings, 5.2 mm. Eyes of male separated on

meson of head by a length half maximum length of

median ocellus, upper portion of eyes on short

stalk, lower portion of eyes a little more than twothirds length of upper portion; eyes of female separated on meson of head by a length 3i/2 times

maximum width of an eye. Fore wings of male

(Figures 21, 24, 27) : vein Rs of fore wings forked

less than one-fourth of distance from base to margin; vein MA forked less than to more than half of

distance from base to margin, fork asymmetrical,

distal portion of vein MA sagged posteriorly; vein

MP forked one-third to more than a third of distance from base to margin, fork symmetrical; cubital area as in Figures 21, 24, and 27; cross veins few.

Fore wings of female (Figure 30) : vein Rg of fore

wings forked less than one-fourth of distance from

base to margin; vein MA forked less than half of

distance from base to margin, fork symmetrical;

vein MP forked one-fifth of distance from base to

margin, fork symmetrical; fork of veins MA and

MP narrower than those forks in male wings; cubital area as in Figure 30; cross veins few. Costal

projection of hind wings of male and female acute,

short, and well developed (Figures 22-23, 25-26, 28-

29, 31-32), apex of projection located less than half

to half distance from base; apex of wings broad

and angularly truncated (Figures 22-23, 25-26, 28-

29, 31-32); cross veins few. Legs: ratios of segments

in male fore legs, 0.50: 1.00 (3.0 mm.) : 0.04: 0.32:

0.29: 0.18: 0.11. Claws of a pair dissimilar, one

apically hooked (Figure 99), other obtuse, padlike.

Male genitalia (Figures 80-83) : segments 2 and 3

of forceps short; base of forceps broad, its inner

margin forming an angular bend; length of styliger

plate along median line less than one-third to

greater than one-third length of maximum width;

penes divided, tubular. Ninth sternum of female

deeply cleft (Figure 106). Female with a short ovipositor or egg guide extending to anterior third of

segment 8 (Figure 112). Terminal filament greatly

reduced, with 6-7 segments, basal 3 segments short

and remaining apical segments long (Figure 121).

MATURE NYMPH.—Head prognathous. Antennae

2i/2 times maximum length of head. Mouthparts

(Figures 141, 150, 159, 169, 181-182): dorsal hair

on labrum as in Figure 181; submedian and anterolateral areas of hair ventrally; 4 to 5 equal-sized denticles on anteromedian emargination (Figure 182).

Clypeus as in Figure 181. Left mandible as in Figure 169. Lingua of hypopharynx with welldeveloped lateral processes (Figure 159), anterior
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

margin deeply cleft; superlingua of hypopharynx as

in Figure 159, with a row of hair along anterior

margin. Segment 2 of maxillary palpi I14 times

length of segment 1; segment 3 a little less than half

to half length of segment 2, triangular; a V-shaped

ridge near the ventral, inner anterolateral margin

of maxillae, hair on maxillae as in Figure 150. Labium as in Figure 141; segment 2 of palpi a little

longer than length of segment 1; segment 3 onefourth length of segment 2, triangular; paraglossae

ventral to glossae. Fine hair on anterolateral margins of prothorax only. Legs (Figure 199) : claws

apically hooked, denticles on claws progressively

larger apically. Gills (Figure 206): gills on segments 1-7 alike, each portion of gills long and slender. Posterolateral spines on abdominal segments

5 or 6 to 9, spines progressively larger posteriorly.

Terminal filament a little longer than cerci.

ETYMOLOGY.—Latin careo, meaning to be deprived of; Latin spina, meaning spine.

TYPE-SPECIES.—Careospina hespera Peters and

Alayo, new species.

DISCUSSION.—Careospina, new genus, can be distinguished from other West Indian genera of the

Leptophlebiidae by the following combination of

characters. In the imagos (1) the terminal filament

is greatly reduced and with only 6-7 segments (Figure 121), (2) the penes of the male genitalia are

divided and tubular (Figures 80-83), and (3) the

costal projection of the hind wings is acute, short,

and well developed (Figures 22-23, 25-26, 28-29,

31-32). In the nymphs (1) posterolateral spines

are present on abdominal segments 5 or 6 to 9 (Figure 212), (2) the tarsal claws are apically hooked;

the denticles on the claws are progressively larger

apically (Figure 199), and (3) 4 to 5 equal-sized

denticles are present on the anteromedian emargination of the labrum (Figure 182). Careospina can

be distinguished from all genera of the Leptophlebiidae by the following combination of characters. In the imagos (1) the base of vein ICux of

the fore wings is attached to vein CuP (Figures 21,

24, 27, 30), (2) the penes of the genitalia are divided and tubular (Figures 80-83), (3) the terminal filament is greatly reduced, and with only

6-7 segments (Figure 121), and (4) the eyes of the

males are separated on the meson of the head by a

length half the maximum width of the median

ocellus. In the nymphs (1) the abdominal gills are

alike on segments 1-7, each portion of the gills is

long and slender (Figure 206); (2) 4 to 5 equalsized denticles are present on the anteromedian

emargination of the labrum (Figure 182); (3) the

tarsal claws are apically hooked, the denticles on

the claws are progressively larger apically (Figure

199); and (4) posterolateral spines are present on

abdominal segments 5 or 6 to 9 (Figure 212).

Key to the Species of Careospina Peters, new genus

MALE IMAGOS

Forceps of genitalia 3i/£ times length of penes (Figure 83); Haiti . .. C. anmtlata, new species

Forceps of genitalia a little longer than length of penes (Figures 80, 82); Cuba 2

Basal two-thirds and apex of prothoracic femora darker in color; color markings on abdominal segments 5 and 6 as in Figure 127; Pinar del Rio Prov., Cuba.

C. hespera, new species

Apex of prothoracic femora darker in color; color markings on abdominal segments 5 and

6 as in Figure 128; Trinidad Mountains, Cuba C. minuta, new species

Clave para las Especies de Careospina Peters, nuevo g£nero

IMAGOS MACHOS

1. Pinzas de la genitalia 3i/2 veces del largo de los penes (Figura 83); Haiti.

C. annulata, nuevo especie

Pinzas de la genitalia solo algo mas largas que la longitud de los penes (Figuras 80, 82);

Cuba 2

2. Femures protoracicos con los y3 basales y el apice, de color mas oscuro que el resto; marcas

de color en los segmentos abdominales 5 y 6 como en la Figura 127; Prov. de Pinar del
NUMBER 62 13

Rfo, Cuba C. hespera, nuevo especie

Femures protoracicos unicamente con los apices de color mas oscuro que el resto; marcas

de color en los segmentos abdominales 5 y 6 com en la Figura 128; montanas de Trinidad,

Cuba C. tninuta, nuevo especie

Careospina hespera Peters and Alayo, new species

FICURES 21-23, 80-81, 99, 121, 127, 141, 150, 159, 169, 181-182,

199, 206, 212

MALE IMAGO (in alcohol).—Length: body, 6.0-

6.5 mm.; fore wings, 6.0-6.5 mm. Upper portion of

eyes and stalks brown, lower portion black. Head

brown, carinae blackish brown. Base of ocelli

blackish brown, remainder white. Thorax and

coxae of legs brown, sutures paler, carinae blackish

brown. Trochanters, femora, and tibiae of prothoracic legs brown, apexes of femora and tibiae

darker, remainder of legs paler; mesothoracic and

metathoracic legs pale, femora with median and

apical darker brown maculae. Wings (Figures 21-

23) : longitudinal veins of fore and hind wings

brown, cross veins paler; membrane of fore and

hind wings hyaline. Abdomen: terga 1 and 2 dark

brown, terga 3-9 light brown, posterior margin of

terga 3-7 darker, color markings on terga 3-9 as

in Figure 127, spiracles and tracheae blackish brown

on segments 3-9; sterna light brown. Genitalia

(Figures 80-81): forceps segment 1 dark brown,

forceps segment 2 washed with brown, remainder

of forceps and penes paler; forceps a little longer

than length of penes. Caudal filaments pale.

FEMALE IMAGO.—Unknown.

MATURE NYMPH (in alcohol).—Head brown, venter paler. Thorax brown, sterna paler, lateral margins of terga black. Legs pale, markings as in male

imagos. Abdomen brown, sterna paler, markings as

in male imagos (Figure 212). Gills gray, tracheae

black. Caudal filaments pale.

Holotype male imago, Cuba: Pinar del Rio Prov.,

Rio Cuzco, near Soroa (field no. 24), March 1965;

paratopotypes: 4 male imagos; paratypes: 2 male

imagos, Cuba: Pinar del Rio Prov., Arroyo Pinar de

Vinales (field no. 22) ; 2 nymphs, Cuba: Pinar del

Rio Prov., Rio Cuzco, near Soroa (field no. 3),

March 1965; 2 nymphs, Cuba: Pinar del Rio Prov.,

Vinales (field no. 11), October 1964; 3 nymphs,

Cuba: Pinar del Rio Prov., Soroa (field no. 13),

December 1964. All types were collected by P. Alayo

D. and are in alcohol. Holotype and one nymphal

paratype were deposited in the collections of the

Institute of Biology, Academy of Sciences of the

Republic of Cuba. One paratopotype, one male

imaginal paratype, and two nymphal paratypes are

in the collections of the University of Utah. One

paratopotype and two nymphal paratypes were deposited in the collections of the National Museum

of Natural History. All remaining types are deposited in the collections of Florida A & M

University.

ETYMOLOGY.—Latin hesperus, meaning west.

DISCUSSION.—Careospina hespera can be distinguished from the other two described species of

Careospina by the following characters. In the male

imagos (1) the forceps of the genitalia are a little

longer than the length of the penes (Figure 80),

(2) the basal two-thirds and apex of the prothoracic

femora are darker brown in color, and (3) the color

markings on abdominal segments 5 and 6 are as in

Figure 127. The nymphs and adults of C. hespera

are associated by the developing wing pads and

color markings on the legs and abdomen of the

nymphs collected from the same locality as the

adults.

Imagos of this species were observed by Dr. P.

Alayo D. flying along a shady stream, on a clear

and sunny spring morning at about 11 A.M. All the

specimens were males, and they were flying leisurely

at about four to five feet above the stream. Dr.

Alayo walked along the stream about twenty meters,

and the flight covered all the water. He reports that

this species is not very common, being very local

and living in only a small fraction of the bodies of

water visited.

Careospina miniita Peters, new species

FIGURES 24-26, 82, 128

MALE IMAGO (pinned).—Length: body, 3.2 mm.;

fore wings, 3.8 mm. Upper portion of eyes and

stalks dark brown, lower portion black. Head dark

brown. Base of ocelli dark brown, remainder white.

Thorax dark brown, sutures paler, pleura with

contrasting pale and dark areas. Coxae of legs dark

brown; prothoracic legs brown, basal two-thirds and
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

apex of femora darker, apex of tibiae darker, tarsi

paler; mesothoracic and metathoracic legs light

brown, a wide median, and narrow subapical darker

bands on femora, apex of tibiae and tarsi darker.

Wings (Figures 24-26) : longitudinal and cross

veins of fore and hind wings brown; membrane of

fore and hind wings hyaline, washed lightly with

brown; costal projection of hind wings acute, short

and well developed, base of projection narrow (Figures 25-26). Abdomen: light brown, segments 1, 9,

and 10 darker; posterior margin of segments 2-8

darker; submedian and sublateral dark brown markings projecting anteriorly to near anterior margin

of terga 2-8 as in Figure 128. Genitalia (Figure

82) : light brown; forceps a little longer than length

of penes. Caudal filaments light brown, annulations

at articulations darker.

FEMALE IMAGO.—Unknown.

MATURE NYMPH.—Unknown.

Holotype male imago, Cuba: Trinidad Mountains, Mina Carlota, 22 March 1935; paratopotype:

1 male imago; paratypes: 5 male imagos, Cuba (no

other data), C. Wright. Holotype, paratopotype,

and paratypes pinned. One paratype was deposited

in each of the collections of Florida A & M University, University of Utah, and National Museum

of Natural History. Remaining types are deposited

in the collections of the Museum of Comparative

Zoology.

ETYMOLOGY.—Latin minutus, meaning small.

DISCUSSION.—Careospina minuta can be distinguished from the remaining species of Careospina

by the following characters. In the male imagos (1)

the forceps of the genitalia are a little longer than

the length of the penes (Figure 82), (2) the apex

of the prothoracic femora is darker brown in color,

and (3) the color markings on abdominal segments

5 and 6 are as in Figure 128.

The four male imagos with spotless wings considered as a possible new species by Eaton (1882,

1884) in the discussion on Hagenuhis caligatus are

the specimens collected by C. Wright and described

herein as C. minuta. An additional unreported

specimen of this series is included herein.

I have seen nymphs and one female subimago of

Careospina which appear to represent an undescribed species. The abdominal color markings are

entirely different than those of the described species

of Careospina. The specimens are recorded from

various localities in Oriente Province, Cuba. The

one female subimago is used in the generic description. Nymphal specimens of this undescribed species

were deposited in the collections of the University

of Utah and Florida A & M University; the female

subimago was deposited in the collections of Florida A & M University.

Careospina annulata Peters, new species

FIGURES 27-29, 83

MALE IMAGO (pinned).—Length: body, 4.0 mm.;

fore wings, 4.6 mm. Upper portion of eyes and

stalks dark brown, lower portion black. Head dark

brown. Base of ocelli dark brown, remainder white.

Thorax dark brown, sutures paler. Coxae of legs

dark brown, trochanters paler, remainder of legs

broken off and missing. Wings (Figures 27-29) :

longitudinal veins of fore and hind wings brown,

cross veins paler; membrane of fore and hind wings

hyaline; costal projection narrow (Figures 28-29).

Abdomen: dark brown, a little paler on anterior

margin of segments 2-7; posterior margin of segments 2-7 with a narrow, darker brown band and

a little paler band ahead of posterior band, thus

giving the abdomen an annulated appearance.

Genitalia (Figure 83): brown; forceps 3y2 times

length of penes. Caudal filaments light brown, annulations at articulations darker.

FEMALE IMACO.—Unknown.

MATURE NYMPH.—Unknown.

Holotype male imago, Haiti: Mt. La Hotte,

1,000 feet, Roche Croix, 14 October 1934, P. J. Darlington, Jr. Holotype pinned and deposited in the

collections of the Museum of Comparative Zoology.

ETYMOLOGY.—Latin annulus, meaning annulated.

DISCUSSION.—Careospina annulata can be distinguished from the remaining species of Careospina

by the following characters. In the male imagos (1)

the costal projection of the hind wings is acute,

short, and well developed; the base of the projection

is narrow (Figures 28-29), (2) the genital forceps

are 3i/2 times the length of the penes (Figure 83),

and (3) the color pattern on the abdominal segments is as given in the description.
NUMBER 62 15

Genus Neohagenulus Traver, 1938

FIGURES 33-42, 84-86, 100, 107, 113, 142, 151, 160, 170, 183-

185, 200, 207.

Neohagenulus Traver, 1938, p. 8; 1959, p. 4.

TYPE-SPECIES.—Neohagenulus julio Traver, by

original designation.

SPECIES INCLUDED.—Neohagenulus julio Traver,

1938, p. 9; N. luteolus Traver, 1938, p. 13; N. tinetus Traver, 1938, p. 12.

DISTRIBUTION.—Puerto Rico.

IMAGO.—Length of male: body, 5.0-7.0 mm.; fore

wings, 5.5-7.0 mm. Length of female: body, 5.0-6.0

mm.; fore wings, 5.0-6.3 mm. Eyes of male meet on

meson of head, lower portion of eyes two-thirds

length of upper portion; eyes of female separated on

meson of head by a length 4 to 7 times maximum

width of an eye. Fore wings of male (Figure 33):

vein Rg of fore wings forked one-fifth to less than

one-fourth of distance from base to margin; vein

MA forked half of distance from base to margin,

fork asymmetrical, distal portion of vein MA sagged

posteriorly; vein MP forked less than half of distance from base to margin, fork symmetrical; cubital area as in Figure 33; cross veins few. Fore wings

of female (Figure 36): vein Rg of fore wings forked

one-sixth to one-fifth of distance from base to margin; vein MA forked less than half to two-thirds of

distance from base to margin, fork a little asymmetrical, distal portion of vein MA a little sagged

posteriorly; vein MP forked one-third of distance

from base to margin, fork symmetrical; fork of veins

MA and MP narrower than those forks in male

wings; cubital area as in Figure 36; cross veins few.

Costal projection of hind wings of male and female

acute, long, and well developed (Figures 34-35, 37-

42), apex of projection located past apex of wings;

apex of wings narrow and rounded (Figures 34—35,

37-42) ; cross veins few. Legs: ratios of segments in

male fore legs, 0.52: 1.00 (2.1 mm.) : 0.05: 0.29:

0.21: 0.14: 0.10. Claws of a pair dissimilar, one

apically hooked (Figure 100), other obtuse, padlike. Male genitalia (Figures 84-86): segments 2

and 3 of forceps short; basal segment of forceps

greatly bowed, base of forceps broad, its inner margin forming an angular bend; length of styliger

plate along median line less than one-fourth to less

than one-third length of maximum width; penes

divided, tubular, apex of each penis lobe bulbous,

basal half of lobes connected by a sclerotized membrane, a subapical, ventral spine on each lobe.

Ninth sternum of female deeply cleft (Figure 107).

Female with a short ovipositor or egg guide extending to anterior margin of segment 8 (Figure

113). Terminal filament longer than cerci.

MATURE NYMPH.—Head prognathous. Antennae

3 times maximum length of head. Mouthparts (Figures 142, 151, 160, 170, 183-185): dorsal hair on

labrum as in Figure 183; submedian and anterolateral areas of hair ventrally; 4 to 5 small

equal-sized denticles on anteromedian emargination

(Figures 184-185). Clypeus as in Figure 183. Left

mandible as in Figure 170. Lingua of hypopharynx

with well-developed lateral processes (Figure 160),

anterior margin deeply cleft; superlingua of hypopharynx as in Figure 160, with a row of hair along

anterior margin. Segment 2 of maxillary palpi equal

to length of segment 1; segment 3 one-half to a

little shorter than length of segment 2, triangular;

a V-shaped ridge near the ventral inner anterolateral

margin of maxillae; hair on maxillae as in Figure

151. Labium as in Figure 142; segment 2 of palpi a

little longer to one and one-third times length of

segment 1; segment 3 one-fifth to one-fourth times

length of segment 2, triangular; paraglossae ventral

to glossae. Fine hair on anterolateral margins of

prothorax only. Legs (Figure 200): claws apically

hooked, denticles on claws progressively larger

apically. Gills (Figure 207) : gills on segments 1-7

alike; each gill deeply forked (Figure 207) . Posterolateral spines on abdominal segments 3-9, spines

progressively larger posteriorly. Terminal filament

longer than cerci.

DISCUSSION.—Traver (1938) established Neohagenulus for three species she described from

Puerto Rico. Excellent descriptions are given in

Traver (1938) for these three species. Included

herein are keys written by Traver (1938) for the

three species and nymphs of two undescribed species

of Neohagenulus and Spanish translations of the

keys.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Species of Neohagenulus Traver

IMAGOS

1. Wings of both sexes distinctly brown-tinged throughout; cross vein prominent.

N. tinctus Traver

Wings not distinctly brown-tinged throughout; cross veins less prominent, except along

costal margin 2

2. Longitudinal veins in wing of male yellowish along costal margin, elsewhere paler; costal

band in wing of female rather dark red brown N. luteolus Traver

Longitudinal veins, also cross veins of costal area and apical third of wing brown; costal

band in wing of female pale brown, usually paler toward apex N. julio Traver

MATURE NYMPHS

1. Pale areas on head between eyes and ocelli distinctly divided by a grayish black line; spines

on fore femur and on hind tibia relatively short N. luteolus Traver

Pale areas on head between eyes and ocelli not divided by a dark line; spines on fore femur

and hind tibia longer 2

2. Second joint of antenna pale; ventral markings, when present, usually confined to lateral

part of posterior margins of sternites S

Second joint of antenna blackish, at least in apical half; ventral markings more extensive.

4

3. Ventral markings usually absent; abdominal tergites of female largely yellowish.

N. species 2

Ventral markings usually present; abdominal tergites of female largely reddish brown.

N. julio Traver

4. Body (female) 7 to 8 mm. in length N. spedes 1

Body (female) 6 to 6i/z mm. in length iV. tinctus Traver

Clave para las Especies de Neohagenulus Traver

IMAGOS

1. Alas en ambos sexos distintamente tenidas de pardo en toda su extension; nervuras transversas prominentes N. tinctus Traver

Alas no distintamente tenidas de pardo en toda su extensidn; nervuras transversas menos

prominentes, excepto a lo largo del margen costal 2

2. Nervuras longitudinales de las alas del macho amarillentas a lo largo del margen costal,

el resto mas palido; banda costal en las alas de la hembra de un color pardo rojizo oscuro.

N. luteolus Traver

Nervuras longitudinales, y tambien las transversales del area costal y tercio apical del ala,

pardas; banda costal en las alas de la hembra de color pardo palido, usualmente mas

palida hacia el apice N. julio Traver

NlNFAS MAOURAS

1. Areas palidas en la cabeza, entre los ojos y ocelos, distintamente divididas por una Hnea

grisaceonegra; espinas del femur anterior y tibia posterior relativamente cortas.

N. luteolus Traver

Areas palidas en la cabeza, entre los ojos y ocelos, no divididas por una linea oscura; espinas

en el femur anterior y la tibia posterior mas largas 2

2. Segundo segmento antenal palido; marcas ventrales, si existen, usualmente connnades a ia

parte lateral de los margenes posteriores de los esternitos 3

Segundo segmento antenal negruzco, por lo menos en su mitad apical; marcas ventrales

mas extensas 4

3. Marcas ventrales usualmente ausentes; tergitos abdominales de la hembra mayormente

amarillentos 2V. especie 2
NUMBER 62 17

Marcas vcntrales usualmente prescntes; tergitos abdominales de la hembra mayormente

pardorojizos N. julio Traver

Longitud cuerpo (hembra) 7-8 mm 2V. especie 1

Longitud cuerpo (hembra) 6-61/2 mm N. tinctus Traver

Genus Hagenulus Eaton, 1882

FICURES 1-3, 43-66, 87-91, 101, 108, 114-118, 129-138, 143-

145, 152-154, 161-163, 171-173, 186-191, 201, 208

Hagenulus Eaton, 1882, p. 207; 1884 (1883-1888), p. 113.—

Morrison, 1919, p. 144.—Ulmer, 1920, p. 117.—Needham

and Murphy, 1924, pp. 10, 11—Ulmer, 1932, p. 203 —

Traver, 1938, p. 7; 1959, p. 2.—Peters, Gillies, and Edmunds, 1964, p. 117.

TYPE-SPECIES.—Hagenulus caligatus Eaton, by

original designation.

SPECIES INCLUDED.—Hagenulus caligatus Eaton,

1882, p. 207; H. eatoni Banks, 1924, p. 423; H.

jamaicensis, new species; H. morrisonae, new species; H. rangelae, new species.

DISTRIBUTION.—Cuba, Haiti, Jamaica, Puerto

Rico.

IMAGO.—Length of male: body, 4.2-6.5 mm.; fore

wings, 4.8-7.2 mm. Length of female: body, 4.5-

6.2 mm.; fore wings, 5.6-7.0 mm. Eyes of male separated on meson of head by a length equal to width

of median ocellus, lower portion of eyes two-thirds

length of upper portion; eyes of female separated on

meson of head by a length 4 times maximum width

of an eye. Wings (Figures 43-66) : vein Rs of fore

wings forked less than one-fourth to one-third of

distance from base to margin, fork wider in male;

vein MA forked less than one-third to half of distance from base to margin, fork wider in male, fork

symmetrical to a little asymmetrical in female, fork

a little asymmetrical to asymmetrical in male, distal portion of vein MA sagged posteriorly in male

and female; vein MP forked less than one-fourth to

half of distance from base to margin, fork wider in

male, fork symmetrical in male and female; cubital

area as in Figures 43, 46, 49, 52, 55, 58, 61, 64; cross

veins few; several to many cross veins in fore wings

of female surrounded with dark clouds (Figures

46, 52, 55, 61) ; several cross veins in fore wing of

male surrounded with dark clouds (Figures 43, 49,

58, 64) . Costal projection of hind wings acute, long,

and well developed, base of projection wide, apex

located near middle to apex of uings (Figures 44-

45, 47-48, 50-51, 53-54, 56-57, 59-60, 62-63, 65-

66); apex of wings narrow to broad and rounded

(Figures 44-45, 47-48, 50-51, 53-54, 56-57, 59-60,

62-63, 65-66); cross veins few. Legs: ratios of segments in male fore legs, 0.61: 1.00 (1.8 mm.) : 0.06:

0.36: 0.28: 0.17: 0.08. Claws of a pair dissimilar, one

apically hooked (Figure 101), other obtuse, padlike. Male genitalia (Figures 87-91) : segments 2

and 3 of forceps short; basal segment of forceps

long, base of forceps broad, its inner margin forming an angular bend (Figures 87-89) ; length of

styliger plate along median line less than half to

more than half length of maximum width; penes

divided, tubular, a subapical, ventral spine present

(Figures 87, 89-91) or absent (Figure 88) on each

lobe. Ninth sternum of female deeply cleft (Figure

108). Female with a well-developed ovipositor or

egg guide extending to just past posterior margin

of segment 10 (Figures 114-118). Terminal filament longer than cerci.

MATURE NYMPH.—Head prognathous. Antennae

2 to 2i/£ times maximum length of head. Mouthparts (Figures 143-145, 152-154, 161-163, 171-173,

186-191): dorsal hair on labrum as in Figure 190,

or Figures 186, 188; submedian and anteromarginal

areas of hair ventrally; 3 to 5 small blunt equalsized denticles on anteromedian emargination (Figures 187, 189, 191); labrum a little (Figure 190)

to greatly (Figures 186, 188) expanded laterally.

Clypeus as in Figures 186, 188, 190. Left mandible

as in Figure 171, or 172, or 173. Lingua of hypopharynx with well-developed lateral processes (Figures 161-163), anterior margin shallowly to deeply

cleft; superlingua of hypopharynx as in Figures

161-163, with a row of hair along anterior margin.

Segment 2 of maxillary palpi equal to length of

segment 1; segment 3 less than one-half times length

of segment 2, triangular; maxillary palpi a little expanded and extending anteriorly past labrum (Figure 154) ; a V-shaped ridge on the ventral, inner

anterolateral margin of maxillae; hair on maxillae

as in Figure 154; to segment 2 of maxillary palpi

li/2

 t o

 less than 2 times length of segment 1;

segment 3 one-half times length of segment 2, triangular; maxillary palpi greatly elongated (Figures

152-153); a large toothlike projection on the inner
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

anterolateral margin of maxillae; hair on maxillae

as in Figures 152-153. Labium as in Figure 145;

segment 2 a little longer than length of segment 1;

segment 3 one-fourth times length of segment 2,

triangular; paraglossae ventral to glossae; to labium as in Figures 143-144; segment 2 less than 2

times length of segment 1; segment 3 one-sixth to

one-eighth times length of segment 2, triangular;

labial palpi greatly elongated (Figures 143-144) ;

paraglossae ventral to glossae. Fine hair on anterolateral margins of prothorax; long hair on inner

and dorsal surfaces of prothoracic tibiae present

(Figures 1-2), or absent (Figure 3). Legs (Figures

1-3, 201): claws apically hooked, denticles on claws

progressively larger apically. Gills (Figures 1, 208) :

gills on segments 1-7 alike, each gill deeply forked.

Posterolateral spines on abdominal segments 5-9,

spines progressively larger posteriorly. Terminal

filament longer than cerci.

DISCUSSION.—Eaton (1882) established Hagenulus for the single species H. caligatus, which was

described only from the adults. Morrison (1919)

described what she thought were the nymphs of

Hagenulus. Later Banks (1924) described a second

species, H. eatoni, which is known only from female

imagos. Traver (1938) discussed the relationships

of Hagenulus with other West Indian genera of

the Leptophlebiidae. Various authors between 1892

and 1956 assigned six new species of the Eastern

Hemisphere Leptophlebiidae to Hagenulus. Peters,

Gillies, and Edmunds (1964) placed four of these

species, H. turbinatus Ulmer (1909), H. scotti

Eaton (1913), H. (?) duliti Demoulin (1954), and

H. fasciatus Kimmins (1956), in other genera occurring only in the Eastern Hemisphere, but tentatively retained H. monstratus Eaton (1892) and

H. karnyi Ulmer (1939) in Hagenulus. I have examined the type series of both H. monstratus and

H. karnyi and neither species represents Hagenulus.

Both species will be transferred to other genera in

a future paper. The true genus Hagenulus is actually restricted to the West Indies.

In 1963, I reared Hagenulus in Jamaica. The

nymphs of this reared material are morphologically

similar to nymphs discussed and keyed by Traver

(1938) as "Jamaica species." The labrum is only

moderately expanded (Figures 3, 190) and the

palpi of the labium (Figure 145) and maxillae

(Figures 3, 154) are only moderately elongated.

The outer edge of the mandibles is smoothly curved

(Figures 3, 173). These nymphs appear to be

adapted to feed on attached periphyton. As this

species is known only from the nymphs and reared

female subimagos, the naming of this species is delayed until male and female imagos are known.

Specimens were collected at a small stream, \\/%

miles southwest crossing Dick's Pond Trail, Hardwar Gap, St. Andrew Parish, Jamaica, and specimens were deposited in the collections of the

University of Utah and Florida A & M University.

The Jamaican nymphs do not resemble the

nymphs described by Morrison (1919) as Hagenulus. The labrum of the nymphs described by Morrison (1919) is greatly expanded laterally (Figures

1, 186), and the palpi of the labium (Figure 143)

and maxillae (Figures 1, 152) are greatly elongated.

The outer edge of the mandibles is strongly angular (Figures 1, 171). Also long hair is present on

the prothoracic tibiae (Figure 1). The remaining

morphological structures are similar to those of the

Jamaican nymphs. These nymphs appear to strain

their food from the current. The Morrison-type

nymph is herein described as H. caligatus.

Nymphs of another Cuban species are intermediate between the Jamaican and Morrison types. The

labrum is a little expanded laterally (Figures 2,

188), but the palpi of the labium (Figure 144) and

maxillae (Figures 2, 153) are greatly elongated.

The outer edge of the mandibles is not greatly

angular (Figure 2, 172). Long hair is present on

the prothoracic tibiae (Figure 2). This intermediate

is described herein as the nymph of H. morrisonae.

A probable evolutionary shift from feeding on

attached periphyton to straining food from the

current has occurred among species of Hagenulus.

As a shift in type of feeding and the associated

changes in mouthpart morphology of the nymphs

are the only characters separating the two groups

in the nymphs or adults, all species are included

within the genus Hagenulus.

Traver (1938) questioned whether the nymphs

described by Morrison were Hagenulus. Eaton's

(1884) figure of the penes of H. caligatus showed

no subapical ventral spines; however, the developing penes dissected by Traver from the nymphs

described by Morrison possessed such spines. As indicated herein the penes of H. caligatus possess

spines.

Hagenulus can be distinguished from other West

Indian genera of the Leptophlebiidae by the fol-
NUMBER 62 19

lowing combination of characters. In the imagos

(1) the costal projection of the hind wings is long,

acute and well developed (Figures 44-45, 47-48,

50-51, 53-54, 56-57, 59-60, 62-63, 65-66), (2) the

penes of the male genitalia usually possess subapical

ventral spines (Figures 87-91), and (3) few to many

of the cross veins of the fore wings of the males and

females are surrounded with dark clouds (Figures

43, 46, 49, 52, 55, 58, 61, 64). In the nymphs (1)

hair on the dorsal surface of the labrum is not in

two even rows (Figures 1-3, 186, 188, 190), (2) the

denticles on the claws are progressively larger

apically (Figure 201), and (3) posterolateral spines

are present on abdominal segments 5-9; the spines

are progressively larger posteriorly. Hagenulus can

be distinguished from all genera of the Leptophlebiidae by the following combination of characters. In the imagos (1) well-developed hind wings

are present (Figures 44-45, 47-48, 50-51, 53-54,

56-57, 59-60, 62-63, 65-66), (2) the females have

a well-developed ovipositor or egg guide extending

just past the posterior margin of segment 10 (Figures 114-118), (3) the penes of the male genitalia

usually possess subapical spines (Figures 87-91),

and (4) few to many of the cross veins of the fore

wings of the males and females are surrounded with

dark clouds (Figures 43, 46, 49, 52, 55, 58, 61, 64).

In the nymphs (1) the abdominal gills on segments

1-7 are alike; each gill is deeply forked (Figures 1,

208), (2) the denticles on the claws are progressively larger apically (Figure 201), (3) the labrum

is a little (Figures 3, 190) to greatly (Figures 1-2,

186, 188) expanded laterally, and (4) 3 to 5 equalsized, blunt denticles are present on the anteromedian emargination of the labrum (Figures 187,

1«9, 191).

Key to the Species of Hagenulus Eaton

IMAGOS

Cross veins in apical third of cells C, Sc, and R^ of male fringed with brown color (Figure

58); male genitalia as in Figures 89-90; shape of hind wings as in Figures 59-60, 62-63;

costal projection extremely long; Jamaica H. jatnaicensis, new species

Few to many cross veins of fore wings of male with dark clouds but not as above (Figures

43, 49, 64); male genitalia as in Figures 87-88, 91; shape of hind wings as in Figures 44-

45, 47-48, 50-51, 53-54, 56-57, 65-66; costal projection not extremely long 2

Cross veins in basal half of cells C and Sc of fore wings of male surrounded with dark

brown clouds (Figure 43); ovipositor or egg guide of female extending just past abdominal

segment 10 (Figure 114); Cuba H. caligatus Eaton

Cross veins in fore wings of male not as above (Figures 49, 64); ovipositor or egg guide of

female extending to posterior margin of segment 10 (Figures 115-116) 3

Membrane of fore and hind wings of male hyaline, gray; pleura of thorax of male pale

with dark brown markings, especially in diagonal streaks posteromedially from leg bases;

Puerto Rico H. rangelae, new species

Membrane of fore and hind wings hyaline, light brown; pleura of thorax of male not as

above 4

Sterna 2-9 of female with a median, longitudinal and submedian, longitudinal, black lines;

Haiti H. eatoni Banks

Sterna 2-9 of female uniformly light brown; Cuba H. morrisonae, new species

Clave para las Especies de Hagenulus Eaton

IMAGOS

Macho con las nervuras transversas en el tercio apical de las celdillas C, Sc, y R, de las

alas anteriores, rodeadas de color pardo (Figura 58); genitalia del macho segun las Figuras

89-90; forma de las alas posteriores segiin las Figuras 59-60, 62-63; proyecci6n costal de

misma extremadamente larga; Jamaica H. jatnaicensis, new species

De pocas a muchas nervuras transversas rodeadas de color pardo en las alas anteriores del

macho, pero no como se describe en el inciso anterior (Figuras 43, 49, 64); genitalia del
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

macho segiin Figuras 87-88, 91; forma de las alas posteriores segiin las Figuras 44-45, 47-

48, 50-51, 53-54, 56-57, 65-66; proyecci6n costal de la misma no extremadamente larga

2

2. Macho con las nervuras transversas en la mitad basal de las celdillas C y Sc de las alas anteriores, rodeadas por manchitas pardo-oscuras (Figura 43); ovipositor de la hembra sobrepasando el segmento abdominal 10 (Figura 114); Cuba H. caligatus Eaton

Nervuras transversas de las alas anteriores del macho no como en el inciso anterior (Figuras

49, 64); ovipositor de la hembra llegando solamente al margen posterior del segmento

abdominal 10 (Figuras 115-116) 3

3. Machos con la membrana de ambas alas, hialina, gris; pleuras del t6rax pdlidas con marcas

pardo-oscuras, especialmente en forma de trazos diagonales desde las bases de las patas;

Puerto Rico H. rangelae, nuevo especie

Membrana de ambos pares de alas, hialina, pardo-clara; pleuras toracicas del macho no como

se describen en el inciso anterior 4

4. Esternitos 2-9 de la hembra con una Iinea media negra longitudinal y otra submedia; Haiti

H. eatoni Banks

Esternitos 2-9 de la hembra uniformemente pardo-claros; Cuba.

H. marrisonae, nuevo especie

Hagenulus caligatus Eaton

FIGURES 1, 43-48, 87, 101, 108, 114, 129-130, 143, 152, 161, 171,

186-187, 201.

Hagenulus caligatus Eaton, 1882, p. 207; 1884 (1883-1888), p.

113.

MALE IMAGO (in alcohol).—Length: body, 6.5

mm.; fore wings, 7.2 mm. Upper portion of eyes

light brown, lower portion black. Head light brown.

Base of ocelli black, remainder white. Thorax light

brown, sutures paler, sterna and carinae darker,

diagonal streaks posteromedially from bases of

prothoracic legs. Coxae, trochanters, femora and

basal half of tibiae of prothoracic legs light brown,

remainder of legs paler, except apexes of tibiae

dark brown, large dark brown median and apical

maculae on femora; mesothoracic and metathoracic

legs pale, except large dark brown median and

apical maculae on femora, apex of tibiae dark

brown. Wings (Figures 43-45): longitudinal and

cross veins of fore and hind wings hyaline, except

cross veins in basal half of cells C and Sc of the

fore wings surrounded with dark brown clouds;

membrane of fore and hind wings hyaline except

bases of wings brown. Abdomen: terga light brown,

terga 1 and 9 uniformly washed with dark brown,

terga 2-8 with dark brown markings as in Figures

129-130; sterna light brown. Genitalia (Figure 87):

apical half of forceps segment 1 dark brown, segment 2 brown, remainder of forceps and penes

paler; penes with subapical, ventral spines. Caudal

filaments pale, basal half of filaments with wide,

dark brown annulations at alternate articulations,

apical half of filaments with annulations at every

third articulation.

FEMALE IMAGO (pinned).—Length: body, 5.0-

5.8 mm.; fore wings, 7.0 mm. Eyes black. Head and

thorax light brown, carinae darker. Color and

markings of legs as in male imago. Wings (Figures

46-48): longitudinal and cross veins of fore and

hind wings light brown, membrane hyaline, washed

lightly with brown in cells C and Sc in fore wings;

cross veins of fore wings surrounded with black

clouds. Abdomen: light brown; tergum 1 uniformly

washed with dark brown; posterior margin and lateral sides of terga 2-9 with a dark brown band, at

posterolateral angles bands rounded off where they

meet. Ovipositor or egg guide extending just past

posterior margin of segment 10. Color and markings

of caudal filaments as in male imago.

MATURE NYMPH (in alcohol).—Head light brown.

Mouthparts (Figures 143, 152, 161, 171, 186-187):

dorsal hair on labrum as in Figure 186; labrum

greatly expanded laterally (Figures 1, 186). Left

mandible as in Figure 171. Maxillary palpi greatly

elongated (Figures 1, 152); a large toothlike projection on the inner anterolateral margin of maxillae. Labial palpi greatly elongated (Figure 143).

Thorax light brown, lateral margins of terga black.

Legs light brown, markings as in male and female

imagos. Abdomen light brown, markings as in Figure 1. Gills gray, tracheae black. Caudal filaments

light brown, markings as in male and female

imagos.

DISCUSSION.—Eaton (1882) described H. caligatus

from a male subimago and a series of female imagos.
NUMBER 62 21

The above redescription of H. caligatus is based

on the type series examined by me in the Museum

of Comparative Zoology and additional specimens

as listed below. Although the figure of the genitalia

of the male subimaginal type made by Eaton

(1884) shows no subapical, ventral spines on the

penes, examination of this type indicates the presence of spines as in Figure 87.

The labrum of the immature nymphs of H.

caligatus is not as greatly expanded laterally as the

labrum of the mature nymphs. The remaining

mouthparts of the immature nymphs are similar to

those of mature nymphs. Also the labrum of mature male nymphs is not as greatly expanded laterally as the labrum of mature female nymphs.

Hagenulus caligatus can be distinguished from

the remaining species of Hagenulus by the following characters. In the imagos (1) the cross veins in

basal half of cells C and Sc of the fore wings of the

male are surrounded with dark brown clouds (Figure 43), (2) the ovipositor or egg guide of the

female extends just past the posterior margin of abdominal segment 10 (Figure 114), and (3) the

color pattern on the abdomen ot the male is as in

Figures 129-130. In the nymphs (1) long hair is

present on the inner and dorsal surfaces of the

prothoracic tibiae (Figure 1), (2) the labrum is

greatly expanded laterally (Figures 1, 186), and

(3) the left mandible is as in Figure 171. The

nymphs and adults of H. caligatus are associated

by developing wing pads and color markings on the

legs and abdomen of the nymphs collected from the

same locality as the adults.

Additional localities for H. caligatus are: 1 male

imago, Cuba: La Habana Prov., Giiines (field no.

33), March 1966, P. Alayo D.; 1 male imago, Cuba:

Pinar del Rio Prov., Arroyo Pinar de Vinales (field

no. 23), P. Alayo D.; 1 female subimago, Cuba: La

Habana Prov., Giiines (field no. 33), March 1966,

P. Alayo D.; 9 nymphs, Cuba: Pinar del Rio Prov.,

Rio Cuzco, near Soroa (field no. 1), March 1965,

P. Alayo D.; 1 nymph, Cuba: Pinar del Rio Prov.,

Rio Cuzco, near Soroa (field no. 2), March 1965,

P. Alayo D.; 14 nymphs, Cuba: La Habana Prov.,

Giiines (field no. 5), March 1966, P. Alayo D.; 3

nymphs, Cuba: Pinar del Rio Prov., stream, Vinales

(field no. 11), October 1964, P. Alayo D.; 48

nymphs, Cuba: La Habana Prov., Melena del Sur

(field no. 44), February 1965, P. Alayo D.; 62

nymphs, Cuba: La Habana Prov., Giiines (field no.

56), March 1966, P. Alayo D.; 2 nymphs, Cuba:

Pinar del Rio Prov., brook, 29 October 1965, V.

Landa; 29 nymphs, Cuba: La Habana Prov., Vento,

20 April 1930, J. G. Needham; 10 nymphs, Cuba:

Pinar del Rio Prov., Santa Cruz de los Pinos,

24-28 March 1939, J. G. Needham. All specimens

are in alcohol. One male imago and 45 nymphs are

deposited in the collections of the Institute of

Biology, Academy of Sciences of the Republic of

Cuba. One male imago, 1 female subimago, and

33 nymphs were deposited in the collections of

Florida A & M University. Thirty-four nymphs

were deposited in each of the collections of the

University of Utah, Museum of Comparative Zoology, and National Museum of Natural History.

The nymphs of H. caligatus live in the stony

areas of free flowing rivers and streams, clinging

from the underside of the stones, sometimes in considerable numbers. Dr. Alayo has found imagos

and subimagos resting in branches overhanging

the water, 6-8 feet high. He has collected in early

morning a regular number of specimens by sweeping the vegetation near the water and found that

subimagos are more numerous than imagos. Subimagos moult in about 24 hours.

Hagenulus morrisonae Peters and Alayo, new

species

FIGURES 2, 49-54, 88, 115, 131-132, 144, 153, 162, 172, 188-189,

208

MALE IMAGO (in alcohol).—Length: body, 5.2-

5.8 mm.; fore wings, 6.2-7.2 mm. Upper portion of

eyes brown, lower portion black. Head brown.

Basal half of ocelli black, remainder white. Prothorax light brown, a median, and submedian, longitudinal black bands on notum, lateral margins of

notum black; remaining portion of thorax brown,

except pleura lighter, carinae at bases of wings and

legs black. Coxae dark brown, femora and tibiae

brown, remainder of legs paler; a median and apical

dark brown maculae on femora, apexes of tibiae

dark brown. Wings (Figures 49-51): longitudinal

and cross veins of fore and hind wings brown, cross

veins in cells C, Sc, and Rx of the fore wings faintly

edged in dark brown; membrane of fore and hind

wings hyaline, light brown. Abdomen: light brown;

tergum 1 uniformly washed with dark brown, posterior margin of terga 2-9 dark brown, dark brown

anteromedian markings on terga 2-9 as in Figures
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

131-132, markings lighter on terga 2 and 3. Genitalia (Figure 88) : apical half of forceps segment 1

dark brown, segment 2 brown, remainder of forceps

and penes paler; penes without subapical, ventral

spines. Caudal filaments pale, dark brown annulations at articulations, annulations broader on alternate segments.

FEMALE IMAGO (in alcohol).—Length: body,

4.5-6.2 mm.; fore wings, 5.8 mm. Eyes black. Head

brown. Thorax light brown, color markings on

thorax as in male imago. Legs light brown, dark

brown markings as in male imago except bases and

apex of tibiae dark brown. Wings (Figures 52-54):

longitudinal and cross veins of fore and hind wings

brown, cross veins in fore wings surrounded with

dark brown clouds, clouds lighter along posterior

margin of wings; membrane of fore and hind wings

light brown, membrane in cells C and Sc of the

fore wings translucent, remaining membrane in

fore and hind wings transparent. Abdomen: light

brown, dark brown markings as in male imago.

Ovipositor or egg guide extending to anterior margin of segment 10 (Figure 115). Caudal filaments

as in male imago.

MATURE NYMPH (in alcohol).—Head light

brown. Mouthparts (Figures 144, 153, 162, 172,

188, 189): dorsal hair on labrum as in Figure 188;

labrum a little expanded laterally (Figures 2, 188).

Left mandible as in Figure 172. Maxillary palpi

greatly elongated (Figure 153) ; a large toothlike

projection on the inner anterolateral margin of

maxillae. Labial palpi greatly elongated (Figure

144). Thorax light brown, color markings as in

male and female imagos. Legs light brown, color

markings as in male and female imagos. Abdomen

light brown, color markings as in male and female

imagos. Gills gray, tracheae black. Caudal filaments

light brown, markings as in male and female

imagos.

Holotype male imago, Cuba: Oriente Prov.,

Rio Boniatico, San Luis (field no. 27), November

1964; allotype, Cuba: Las Villas Prov., Rio Anaya,

Ciego Montero, Rodas (field no. 31), December

1965; paratypes: 2 female imagos, same data as for

allotype; 1 male subimago, 1 female subimago,

Cuba: Las Villas Prov., Rio Anaya, Ciego Montero,

Rodas (field no. 32), December 1965; 1 male

imago, 1 male subimago, 1 female subimago, Cuba:

Oriente Prov., Rio Guantanamo (field no. 30); 2

nymphs, Cuba: Oriente Prov., Rio Nagua, near

Turquino (field no. 16), May 1965; 2 nymphs,

Cuba: Oriente Prov., Rio Yara, Central Estrada

Palma, Manzanillo (field no. 17), November 1964;

4 nymphs, Cuba: Oriente Prov., Rio Boniatico, San

Luis (field no. 26), November 1964; 10 nymphs,

same data as for allotype; 6 nymphs, Cuba: Oriente

Prov., Rio San Rafael, San Luis, 16 November

1964, V. Landa. All specimens are in alcohol and

all were collected by P. Alayo D., except as marked.

Holotype, allotype, and eight nymphs are deposited

in the collections of the Institute of Biology, Academy of Sciences of the Republic of Cuba. One male

imago, one male subimago, one female imago, one

female subimago, and eight nymphs were deposited

in the collections of Florida A & M University. One

male subimago, one female imago, one female subimago, and four nymphs were deposited in the

collections of the University of Utah. Four nymphs

were deposited in the collections of the National

Museum of Natural History.

ETYMOLOGY.—Species named for Mrs. Emily R.

Morrison who correctly associated the nymphs and

adults of Hagenulus.

Hagenulus morrisonae can be distinguished from

the remaining species of Hagenulus by the following characters. In the imago (1) the cross veins in

cells C, Sc, and Rx of the fore wings are surrounded

with dark brown clouds (Figures 49, 52), (2) the

penes of the male genitalia possess no subapical

ventral spines (Figure 88), and (3) the color pattern on the abdomen of the male and females is

as in Figures 131-132. In the nymphs, (1) long

hair is present on the inner and dorsal surfaces of

the prothoracic tibiae (Figure 2), (2) the labrum

is a little expanded laterally (Figures 2, 188), and

(3) the left mandible is as in Figure 172. The

nymphs and adults of H. morrisonae are associated

by the developing wing pads and color markings

on the legs and abdomen of the nymphs collected

from the same locality as the adults.

Nymphs of this species are commonly found in

free-flowing rivers, under stones in places where

the water is clear and only about six inches to one

foot deep. Nymphs cling to the underside of the

stones. Subimagos are easily found by turning

stones at the margins of the river where nymphs

are plentiful. Dr. Alayo has found these subimagos

at every hour of the day, resting under the stones

and, when disturbed, running a little or making

a short flight. In the evening, about five or six
NUMBER 62 23

o'clock, but in full daylight, he has seen swarms

of adults flying and dancing in the air very near

the water, about one foot or less from the water's

surface and always near the stony places of the

river. Each swarm includes about 30 or 40 individuals, forming a mass in the air about the size of

a ball measuring 1 to li/2 feet in diameter. As the

darkness grows, these groups are more and more

numerous.

Dr. Alayo observes that subimagos apparently

molt during the day shortly after their emergence,

as he has found the imago several times under

stones beside the discarded subimagal skin.

Hagenulus eatoni Banks

FIGURES 55-57. 116. 133-134

Hagenulus eatoni Banks, 1924, p. 423.

FEMALE IMAGO (pinned).—Length: body, 5.6

mm.; fore wings, 6.7 mm. Eyes black. Head light

brown. Thorax light brown; submedian, longitudinal black bands on pronotum; pleura with

black markings, especially in diagonal streaks posteromedially from leg bases. Legs light brown;

median and apical transverse black bands on femora; apex of tibiae and tarsi black. Wings (Figures 55-57): longitudinal and cross veins of fore

and hind wings brown; membrane of fore and hind

wings hyaline, washed lightly with brown especially

in cells C and Sc of fore wings, cross veins in fore

wings surrounded with black clouds except in anal

area. Abdomen: light brown; tergum 1 uniformly

washed with black, terga 2-9 with black markings

as in Figures 133-134; sterna 2-9 with a median,

longitudinal and submedian, longitudinal, black

lines. Ovipositor or egg guide extends to posterior

margin of segment 10 (Figure 116). Caudal filaments light brown, black annulations at articulations.

DISCUSSION.—The above redescription of H.

eatoni is based on the type series of 11 females

collected from Diquini, Haiti. These types are deposited in the collections of the Museum of Comparative Zoology. The species is known only from

the female imago.

One of these specimens possesses fore wings with

distinct black clouds surrounding the cross veins

only in cells C, Sc, and Rx as in Figure 55. All specimens are more yellowish than the type.

I have seen one female imago from Ennery, Haiti,

which appears to represent an undescribed species

of Hagenulus. The abdomen is light brown, except

the anterior half of terga 2-9 is black. Also the

female ovipositor or egg guide extends past abdominal segment 10 by one-fourth its length. This specimen is deposited in the collections of the Museum

of Comparative Zoology and is pinned.

Hagenulus jamaicensis Peters, new species

FIGURES 58-63, 89-90, 117, 135-136

MALE IMAGO (in alcohol).—Length: body, 4.2-

5.7 mm.; fore wings, 4.8-5.4 mm. Upper portion of

eyes light brown, lower portion black. Head light

brown. Base of ocelli black, remainder light brown.

Thorax light brown, sutures paler, pleura with contrasting pale and dark areas. Coxae, trochanters,

and femora of prothoracic legs brown; large, darker,

apical, median, and distal maculae on dorsal and

ventral surfaces of femora, remainder of legs pale,

apexes of tibiae darker; mesothoracic and metathoracic legs pale, large, darker median and distal

maculae on dorsal and ventral surfaces of femora,

apexes of tibiae darker. Wings (Figures 58-60):

longitudinal and cross veins of fore and hind wings

hyaline, washed lightly with brown; cross veins in

apical third of cells C, Sc, and Rt of fore wings surrounded with brown clouds; costal projection of

hind wings acute, long, and well developed (Figures

59-60). Abdomen: terga hyaline, washed lightly

with brown, terga 1, 8-10 darker, posterior margin

of terga 1-9 each with a darker transverse band,

lateral margins of terga 1-8 with a darker longitudinal band, spiracles and tracheae darker, terga

2-8 with darker, sublateral, diagonal bands (Figures 135-136); sterna hyaline, washed lightly with

brown, sterna 8-9 darker. Genitalia (Figures 89-

90): light brown, apex of basal segment of genital

forceps darker; penes with subapical, ventral spines.

Caudal filaments light brown, wide annulations at

articulations darker.

FEMALE IMAGO (in alcohol).—Length: body, 4.6

mm.; fore wings, 5.6 mm. Eyes black. Head light

brown. Color of thorax and legs as in male imago,

except apex of tibiae light brown. Wings (Figures

61-63): longitudinal and cross veins of fore and

hind wings light brown, membrane hyaline, washed

lightly with brown; cross veins of fore wings sur-
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

rounded with dark brown clouds, clouds lighter in

posterior portion of wings. Abdomen: light brown,

darker color markings as in male imago. Ovipositor

or egg guide extending to just past segment 10

(Figure 117). Caudal filaments as in male imago.

MATURE NYMPH.—Unknown.

Holotype male imago, Jamaica: Clarendon Parish, Rio Minho, Frankfield, 7 January 1955, T. H.

Farr; allotype, same data as holotype except collected 10 March 1954; paratopotypes, 18 male

imagos, 4 female imagos. All types are in alcohol.

Four male and one female paratopotypes were deposited in each of the collections of Florida A & M

University, University of Utah, and National Museum of Natural History. Remaining types deposited in the collections of the Science Museum,

Institute of Jamaica.

ETYMOLOGY.—Species named for Jamaica.

DISCUSSION.—Hagenulus jamaicensis can be distinguished from the remaining species of Hagenulus by the following characters. In the imagos (1)

cross veins in apical third of cells C, Sc, and Rx of

fore wings of males are surrounded with brown

clouds (Figures 58, 61), (2) the costal projection

of the hind wings is acute, long, and well developed

(Figures 59-60, 62-63), and (3) the color pattern

on the male and female abdominal segments is as

in Figures 135-136.

Hagenulus rangelae Peters, new species

FIGURES 64-66, 91, 137-138

MALE IMAGO (pinned).—Length: body, 4.3 mm.;

fore wings, 5.0 mm. Eyes gray; eyes separated on

meson of head by a length equal to maximum width

of a lateral ocellus. Head light brown. Base of lateral ocelli light brown, remainder white; base of

median ocellus dark brown, remainder white.

Thoracic nota and sterna light brown, pleura paler

with dark brown to fuscus markings, especially in

diagonal streaks posteromedially from leg bases.

Legs light yellowish brown, becoming paler apically;

femora with broad, transverse, dark brown bands in

middle and apical portions; apical one-fifth of prothoracic and mesothoracic tibiae dark brown; apex

of each segment of prothoracic tarsi dark brown.

Wings (Figures 64-66): longitudinal veins and

cross veins in fore wings dark brown in anterior

portion of wings, fading to light brown in posterior

portion, light brown cross veins may be washed

lightly with black, but occur in no particular pattern on wings; vein Sc of hind wings dark brown,

basal half of all other longitudinal veins dark

brown, apical half light brown, cross veins light

brown; membrane of fore and hind wings hyaline,

gray; cells C, Sc, and Rx of fore wings surrounded

with black clouds. Abdomen: tergum 1 dark brown,

terga 2-7 pale yellowish brown with dark brown

color markings as in Figures 137-138, terga 8 and

9 similar to tergum 7, but obscured with extensive

median brown markings, tergum 10 light brown;

sterna light yellowish brown, the posterolateral

angles a little darker, especially on sterna 8 and 9.

Genitalia (Figure 91): penes pale yellowish brown;

penes with subapical, ventral spines; forceps broken

off and missing. Caudal filaments broken off and

missing.

FEMALE IMAGO.—Unknown.

MATURE NYMPH.—Unknown.

Holotype male imago, Puerto Rico: San Rangel,

27 June, Otero. Holotype pinned and deposited in

the collections of the Museum of Comparative

Zoology, Cambridge.

ETYMOLOGY.—Species is named for type locality,

San Rangel.

DISCUSSION.—Hagenulus rangelae can be distinguished from the remaining species of Hagenulus

by the following characters. In the male imagos

(1) the cross veins in cells C, Sc, and Rx of the fore

wings are surrounded with black clouds (Figure

64), (2) the membrane in the fore and hind wings

is hyaline, gray, and (3) the color pattern on the

abdominal terga is as given in the descriptions and

Figures 137, 138.

Genus Borinquena Traver, 1938

FICURES 67-72, 92-96, 102-103, 109-110, 119-120, 146-147, 155-

156, 164-165, 174-175, 192-196, 202-203, 209-210

Borinquena Traver, 1938, p. 16; 1959, p. 2.

TYPE-SPECIES.—Borinquena (B.) carmencita

Traver, by original designation.

SPECIES INCLUDED.—Borinquena (B.) carmencita

Traver, 1938, p. 18; B. (B.) contradicens Traver,

1938, p. 20; B. (A.) traverae, new species.

DISTRIBUTION.—Puerto Rico, Dominica, St. Lucia.

IMAGO.—Length of male: body, 5.5-5.7 mm.; fore

wings, 5.2-5.6 mm. Length of female: body, 3.5-6.0
NUMBER 62 25

mm.; fore wings, 5.0-^6.0 mm. Eyes of male separated

on meson of head by a length 1 to 3 times maximum width of a lateral ocellus or eyes meet on

meson of head, lower portion of eyes two-thirds to

three-fourths length of upper portion; eyes of female

separated on meson of head by a length 4 to 5 times

maximum width of an eye. Wings (Figures 67-72):

vein Rg of fore wings forked less than one-fourth to

one-fourth of distance from base to margin, vein

MA forked half of distance from base to margin,

fork asymmetrical, distal portion of vein MA sagged

posteriorly; vein MP forked a little less than half

of distance from base to margin, fork symmetrical;

cubital area as in Figures 67, 72; cross veins few.

Hind wings present (Figures 68-71), or absent.

When present, hind wings well developed, costal

projection acute, long, and well developed, apex of

projection located half distance from base, apex of

wings narrow and rounded (Figures 70-71), cross

veins few (Figures 70-71), or hind wings reduced,

no costal projections or cross veins (Figures 68-69),

apex of wings pointed (Figures 68-69). Legs: ratios

of segments in male fore legs, 0.52: 1.00 (2.2 mm.):

0.05: 0.29: 0.29: 0.27: 0.23. Claws of a pair dissimilar; one apically hooked (Figures 102-103), other

obtuse, padlike. Male genitalia (Figures 92-96):

segments 2 and 3 of forceps short, basal segment of

forceps long and narrow, slightly bowed at apex, its

inner margin forming an angular bend; length of

styliger plate along median line one-third to half

length of maximum width; penes divided, tubular,

a subapical, ventral spine on each penis lobe. Ninth

sternum of female deeply cleft (Figures 109-110).

Female with a well-developed ovipositor or egg

guide extending to posterior margin of segment 10

(Figure 120) to past posterior margin of segment

10 by half its length (Figure 119). Terminal filament longer than cerci.

MATURE NYMPH.—Head prognathous. Antennae

3 times maximum length of head. Mouthparts (Figures 146-147, 155-156, 164-165, 174-175, 192-196):

dorsal hair on labrum as in Figures 192, 195; submedian and anterolateral areas of hair ventrally; 2

large equal-sized denticles and sometimes 1 to 3

small unequal-sized denticles on anteromedian

emargination (Figures 193-194, 196). Clypeus as

in Figures 192 and 195. Left mandible as in Figures

174 and 175. Lingua of hypopharynx with welldeveloped lateral processes (Figures 164-165), anterior margin shallowly to deeply cleft; superlingua

of hypopharynx as in Figures 164—165, with a row

of hair along anterior margin. Segment 2 of maxillary palpi one and one-fourth to one and one-half

times length of segment 1; segment three one-third

to half length of segment 2, triangular; a V-shaped

ridge near the ventral, inner anterolateral margin

of maxillae; hair on maxillae as in Figures 155-156.

Labium as in Figures 146-147; segment 2 of palpi

equal to a little longer than length of segment 1;

segment three one-fifth to one-third length of segment 2, triangular; paraglossae ventral to glossae.

Fine hair on anterolateral margins of prothorax

only. Legs (Figures 202-203): claws apically hooked,

denticles on claws progressively larger apically

except apical denticle much larger. Gills (Figures

209-210) : gills on segments 1-7 alike; each gill consists of one long, slender portion (Figure 209), or

each gill deeply forked (Figure 210). Posterolateral

spines on abdominal segments 6 or 7-9, spines

progressively larger posteriorly. Terminal filament

slightly longer than cerci.

DISCUSSION.—Traver (1938) established Borinquena for two species described by her from Puerto

Rico. Excellent descriptions of these two species,

B. carmencita and B. contradicens, are given in

Traver (1938). Herein one new species of Borinquena is described, B. traverae, from Dominica.

Collections recently made available to me contain

another new species from St. Lucia; however, this

species is not described herein, as it is known only

from the male and female subimagos and nymphs.

The adults of B. traverae and the undescribed

species from St. Lucia possess no hind wings, the

adults of B. carmencita possess reduced hind wings

(Figures 68-69), and the adults of B. contradicens

possess well-developed hind wings (Figures 70-71).

I have found no other adult morphological character that successfully separates B. traverae and the

species from St. Lucia from the remaining described

species of Borinquena. The abdominal gills of the

nymphs of B. traverae and the undescribed species

from St. Lucia are deeply forked (Figure 210),

while the abdominal gills of the nymphs of B. carmencita and B. contradicens consist of one long,

slender portion (Figure 209). I have found no

other nymphal morphological character that successfully separates B. traverae and the undescribed

species from the remaining described species of

Borinquena. Herein a new subgenus, Australphlebia, is established for the species traverae.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Borinquena can be distinguished from other West of characters. In the imagos (1) the base of vein

Indian genera of the Leptophlebiidae by the fol- ICux of the fore wings attaches to vein CuA (Figlowing combination of characters. In the imago (1) ures 67, 72), (2) the penes of the male genitalia

the penes of the male genitalia are divided, tubular are divided, tubular and a subapical ventral spine

and a subapical ventral spine occurs on each penis occurs on each penis lobe (Figures 92-96), (3) the

lobe (Figures 92-96), (2) the female possesses a female possesses a well-developed ovipositor or egg

well-developed ovipositor or egg guide and it ex- guide and it extends to at least past the anterior

tends to at least past the anterior margin of ab- margin of abdominal segment 10 (Figures 119—

dominal segment 10 (Figures 119-120), and (3) 120), and (4) the length of the styliger plate of the

the cross veins in the fore wings of the male and male along the median line is one-third to half the

female are not surrounded with dark clouds (Fig- length of the maximum width (Figures 92, 94, 96).

ures 67, 72). In the nymphs (1) the denticles on In the nymphs (1) a V-shaped ridge occurs near the

the claws are progressively larger apically except ventral, inner anterolateral margin of the maxillae

the apical denticle is much larger (Figures 202- (Figures 155-156), (2) 2 large equal-sized denticles

203), (2) 2 large equal-sized denticles and some- and sometimes 1 to 3 small unequal-sized denticles

times 1 to 3 small unequal-sized denticles occur on occur on the anteromedian emargination of the

the anteromedian emargination of the labrum labrum (Figures 193-194, 196), (3) the denticles

(Figures 193-194, 196), and (3) posterolateral on the claws are progressively larger apically exspines occur on abdominal segments 6 or 7-9. cept the apical denticle is much larger (Figures

Borinquena can be distinguished from all genera of 202-203), and (4) posterolateral spines occur on

the Leptophlebiidae by the following combination abdominal segments 6 or 7-9.

Key to the Species of Borinquena Traver

IMAGOS

1. Hind wings absent; apex of penes of male imago acute (Figure 96); Dominica.

B. (A.) traver at, new species

Hind wings present, although sometimes greatly reduced (figures 68-71); apex of penes of

male imago blunt (Figures 92-95) Borinquena sensu stricto 2

2. Hind wings reduced, with 2 longitudinal veins, costal projection absent (Figures 68-69);

paired, small submedian protuberances on posterior margin of styliger plate of male

genitalia (Figure 92); Puerto Rico B. (B.) carmencita Traver

Hind wings well developed, with 4 longitudinal veins, well developed costal projection

(Figures 70-71); 1 small median protuberance on posterior margin of styliger plate of

male genitalia (Figure 94); Puerto Rico B. (B.) contradicens Traver

MATURE NYMPHS

1. Abdominal gills 1-7 deeply forked (Figure 210); abdominal terga 2-7 with a pale, narrow,

median, longitudinal band; Dominica B. (A.) traverae, new species

Abdominal gills 1-7 each consists of 1 long, slender portion (Figure 209); abdominal terga

2-7 not as above Borinquena sensu stricto 2

2. Two large equal-sized denticles and 3 small unequal-sized denticles on anteromedian emargination of labrum (Figure 194); a wide, blackish, band across head between lateral ocelli,

usually extending lateral to eyes; Puerto Rico B. (B.) carmencita Traver

Two large equal-sized denticles only on anteromedian emargination of labrum (Figure 193);

color markings on head not as above; Puerto Rico B. (B.) contradicens Traver

Clave para las Especies de Borinquena Traver

IMAGOS

1. Alas posteriores ausentes; penes del imago macho con el apice agudo (Figura 96); Dominica.

B. (A.) traverae, nuevo especie
NUMBER 62 27

Alas posteriores presentes, aunque a veces muy reducidas (Figuras 68-71); penes del imago

macho con el apice romo (Figuras 92-95) Borinquena sensu stricto 2

Alas posteriores reducidas, con solo dos nervuras longitudinales y la proyecci6n costal ausente

(Figuras 68-69); genitalia del macho con un par de pequenas protuberancias submedias en

el margen posterior del styliger (Figura 92); Puerto Rico B. (B.) carmencita Traver

Alas posteriores bien desarrolladas, con 4 nervuras longitudinales y la proyeccion costal bien

desarrollada (Figuras 70-71); genitalia del macho con una pequefia protuberancia media

en el margen posterior del styliger (Figura 94); Puerto Rico . . B. (B.) contradicens Traver

NlNFAS MADURAS

Branquias abdominales 1-7 profundamente bifurcadas (Figura 210); tergitos abdominales

2-7 con una estrecha banda media longitudinal de color daro; Dominica.

B. (A.) traverae, nuevo especie

Branquias abdominales 1-7 no bifurcadas, consistentes en unas largas y delgadas dntas

(Figura 209); tergitos abdominales 2-7 no como se describen en el inciso anterior.

Borinquena sensu stricto 2

Labrum con das denticulos grandes de tamafio igual y tres dentfculos pequenos y desiguales

en su emarginadcm antero-media (Figura 194); una ancha banda negruzca a travel de la

cabeza, entre los ocelos laterales, usualmente extendida en los lados hasta los ojos; Puerto

Rico B. (B.) carmencita Traver

Labrum con solo dos dentfculos grandes de tamafio igual en su emarginacion antero-media

(Figura 193); coloraci6n de la cabeza distinta a la descrita en el inciso anterior; Puerto

Rico B. (B.) contradicens Traver

Subgenus Borinquena sensu stricto Traver, 1938

FICURES 67-71, 92-95, 102, 109, 119, 146, 155, 164, 174, 192-

194, 202, 209

IMAGO.—Hind wings well developed, costal projection acute, long, and well developed, apex of

projection located half distance from base, apex of

wings narrow and rounded (Figures 70-71), cross

veins few (Figures 70-71) or hind wings reduced,

no costal projection or cross veins (Figures 68-69),

apex of wings pointed (Figures 68-69). Female with

a well-developed ovipositor or egg guide extending

to past posterior margin of segment 10 by half its

length (Figure 119).

MATURE NYMPH.—Abdominal gills on segments

1-7 alike, each gill consists of one long, slender portion (Figure 209).

DISCUSSION.—The subgenus Borinquena sensu

stricto occurs in Puerto Rico. The characters above

will distinguish it from the subgenus Australphlebia.

Australphlebia, new subgenus

FICURES 72, 96, 103, 110, 120, 147, 156, 165, 175, 195-196, 203,

210

IMAGO.—Hind wings absent. Female with a welldeveloped ovipositor or egg guide extending to posterior margin of segment 10 (Figure 120).

MATURE NYMPH.—Abdominal gills on segments

1-7 alike, each gill deeply forked (Figure 210).

ETYMOLOGY.—Latin australis, meaning southern;

Greek phlebo, meaning vein.

TYPE-SPECIES.—Borinquena (A.) traverae Peters,

new species.

Borinquena (Australphlebia) traverae Peters,

new species

FICURES 72, 96, 103, 110, 120, 147, 156, 165, 175, 195-196, 203,

210

MALE IMAGO (in alcohol).—Length: body, 5.5-

5.7 mm.; fore wings, 5.2-5.6 mm. Upper portion of

eyes light brown, lower portion black; eyes separated on meson of head by a length equal to maximum width of a lateral ocellus. Head light brown,

carinae darker. Base of lateral ocelli light brown,

remainder white; base of median ocellus dark

brown, remainder white. Thorax and coxae of legs

brown, except prothoracic notum and carinae

darker, sutures paler. Legs pale, outer surface of

femora and apex of tibiae of prothoracic legs dark

brown; apex of metathoracic femora dark brown,

remainder of femora washed lightly with dark

brown. Wings (Figure 72): longitudinal veins of

fore wings brown, cross veins paler; membrane of

fore wings hyaline, except distal third of cells C and
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sc translucent; hind wings absent. Abdomen: terga

dark brown, terga 2-7 with a paler, narrow, median,

longitudinal, band; sterna paler. Genitalia (Figure

96): brown, styliger plate darker. Caudal filaments

light brown; basal half of filaments with darker

annulations at articulations.

FEMALE IMAGO.—Unknown.

FEMALE SUBIMAGO (in alcohol).—Length: body,

6.0 mm.; fore wings, 6.0 mm. Eyes black. Head dark

brown. Color of thorax as in male imago. Color and

markings on legs as in male imago, except apex of

prothoracic tibiae paler; outer surfaces of metathoracic femora washed with dark brown, apex of

inner surface dark brown. Wings: longitudinal veins

of fore wings light brown, paler posteriorly, cross

veins pale; membrane of fore wings transparent;

hind wings absent. Abdomen: dark brown, color

markings as in male imago, sterna paler. Ovipositor

extending just past posterior margin of abdominal

segment 10 (Figure 120). Caudal filaments broken

off and missing.

MATURE NYMPH (in alcohol).—Head and thorax

brown, venter paler. Legs pale, brown color markings as in male imago and female subimago, except

prothoracic tibiae entirely pale. Abdomen dark

brown, sterna paler, color markings on terga as in

male imago and female subimago. Gills gray, tracheae black. Caudal filaments pale.

Holotype male imago, Dominica: 0.4-0.5 mile

east of Pont Casse, 8 April-24 July 1964; paratopotypes, 4 male imagos, 8 male subimagos, 1 female

subimago, 13 nymphs; para types, 1 male subimago,

5 nymphs, Dominica: Dleau Gommier, 17 April

1964. All types were collected by O. S. Flint, Jr.,

and are in alcohol. One male imaginal, three male

subimaginal, and four nymphal paratopotypes were

deposited in each of the collections of Florida A & M

University and the University of Utah. Remaining

types deposited in the collections of the National

Museum of Natural History.

ETYMOLOGY.—Species is named for Dr. Jay R.

Traver, Amherst, Massachusetts, who has greatly

contributed to the knowledge of the West Indian

Ephemeroptera.

DISCUSSION.—The subgenus Australphlebia occurs

in Dominica and St. Lucia. The characters above

will distinguish it from the subgenus Borinquena

sensu stricto. The nymphs and adults of B. traverae

are associated by the developing wing pads, the

color markings on the legs, and the color pattern on

the terga of nymphs collected from the same locality

as the adults.

Phylogeny and Relationships of the West Indian

Genera

The phylogeny and relationships of the West Indian genera of Leptophlebiidae will not be fully

understood until these genera are compared in

greater detail with the remaining genera occurring

in the Neotropical region. At present, study of all

Neotropical Leptophlebiidae, the dominant mayfly

family in South and Central America, is not complete enough to make adequate comparisons. Some

phylogenetic conclusions, however, can now be

drawn relative to the West Indian genera.

All six leptophlebiid genera occurring in the West

Indies possess similarities. Venation of the adult fore

wings of all genera is similar (Figures 15, 21, 33, 43,

67), except for the cubital area of the fore wings of

Farrodes (Figures 6, 9, 12). The penes of the male

genitalia are partially to entirely divided (Figures

73-96). In the nymphs, denticles are present on the

anteromedian emargination of the labrum (Figures

176-196), and a V-shaped ridge is present on the

dorsal surface of the maxillae near the inner anterolateral margin (Figures 148-156). Except for those

species of Hagenulus with nymphs specialized for

straining food from the current, the mouthparts of

the nymphs are morphologically similar. The abdominal gills are long and slender (Figures 204,

206-210), except for the platelike gills possessed by

Traverina (Figure 205).

While similarities do occur, there are basic morphological differences, especially in the adults. The

first cubital intercalary attaches to vein CuP in the

fore wings in all genera (Figures 15, 21, 33, 43, 67),

except Farrodes. In Farrodes the first cubital intercalary is independent of both veins CuA and CuP

(Figures 6, 9, 12). The penes of the male genitalia

of Farrodes possess large apical appendages (Figures

73-78), while the penes of the remaining genera

possess either subapical ventral spines (Figures 79,

84-87, 89-96) or no spines (Figures 80-83, 88). No

female ovipositor or egg guide occurs in Farrodes;

a short ovipositor or egg guide occurs in Neohagenulus, Careospina, and Traverina (Figures Hill 3); and a long and well-developed ovipositor or

e

Sg guide is present in the remaining genera (Figures 114-120). The females of all genera except
NUMBER 62 29

Farrodes possess a ninlh abdominal sternum that is

deeply cleft apically (Figures 105-110). The ninth

sternum of the female of Farrodes is entire (Figure

104). The terminal filament in Careospina is greatly

reduced and has only 6 or 7 segments (Figure 121),

but the terminal filament in all other genera is

longer than the cerci. It is apparent that the six

West Indian genera are not all closely related.

Farrodes appears to be the most primitive genus

occurring in the West Indies and may represent a

separate line of evolution. However, it appears to

be a member of the same complex. The first intercalary in the cubital area of the adult fore wings

is not connected either to vein CuA or CuP (Figures 6, 9, 12), but the first intercalary connects to

vein CuP (Figures 15, 21, 33, 43, 67) in all other

West Indian genera. The female imagos possess no

ovipositor or egg guide. A large ventral appendage

arises from the apex of each penis lobe of the male

genitalia (Figures 73-78). The ninth sternum of

the female is entire apically (Figure 104), but the

ninth sternum is greatly cleft in all other West

Indian genera (Figures 105-110). Nymphs of

Farrodes are morphologically similar to the other

West Indian genera. No representative from the

surrounding continental area has been found that

is closely related to Farrodes.

Traverina appears to be more specialized than

Farrodes and may represent a line of evolution separate from Farrodes. The first intercalary in the

cubital area of the adult fore wings connects to vein

CuP (Figures 15, 18). The female imagos possess

an ovipositor or egg guide extending to the anterior

margin of abdominal segment 8 (Figure 111). Each

penis lobe of the male genitalia possesses a subapical, ventral spine (Figure 79). It is reasonable to

assume the ventral appendage in Farrodes has reduced to a ventral spine in Traverina. The ninth

sternum of the female is greatly cleft apically (Figure 105). Nymphs of Traverina are morphologically

similar to nymphs of the other West Indian genera

except for the abdominal gills. The gills are platelike and terminate in 3 processes, with the median

projection longer than the laterals (Figure 205).

No representative from the surrounding continental

area has been found that is closely related to

Traverina.

Careospina appears to be closely related to Traverina but is more apomorphic. The female imagos

possess an ovipositor or egg guide extending to the

anterior third of abdominal segment 8 (Figure

112). The subapical, ventral spines are secondarily

lost on the penes of the male genitalia (Figures 80-

83). The terminal filament is greatly reduced, with

6-7 segments; the basal 3 segments are short, but

the remaining segments are long (Figure 121). All

other genera from the West Indies possess a terminal filament that is at least a little longer than

the cerci. Nymphs of Careospina are morphologically similar to the other West Indian genera. No

representative from the surrounding continental

area has been found that is closely related to

Careospina.

Neohagenulus appears to be related to Traverina

and Careospina but is much more specialized. The

female ovipositor or egg guide extends to the anterior margin of abdominal segment 8 (Figure 113).

The costal projection of the hind wings, however,

is acute, long, and well developed (Figures 34-35,

37-42). While the costal projection in Traverina

and Careospina is well developed, it is much shorter

in length (Figures 16-17, 22-23). Subapical ventral

spines occur on the penes of the male genitalia;

however, the penes are shorter than those of Traverina and Careospina and the basal half of the

penis lobes is connected by a sclerotized membrane

(Figures 84-86). Nymphs of Neohagenulus are

morphologically similar to the other West Indian

genera. No representative from the surrounding

continental area has been found that is closely related to Neohagenulus.

Hagenulus and Borinquena appear to be closely

related. Both seem to be related to Neohagenulus,

but the genera appear to be more apomorphic than

Neohagenulus. It is difficult to determine which of

the two genera, Hagenulus or Borinquena, is more

primitive. The adults of both genera are specialized.

The female ovipositor or egg guide is well developed

and extends to at least the posterior margin of

abdominal segment 10 (Figures 114-120). The

length of the styliger plate in the males is one-third

to more than half its maximum width (Figures 87-

89, 91-92, 94, 96). Subapical, ventral spines occur

on the penes of the male genitalia (Figures 87, 89-

96). All known species of Hagenulus, however,

possess well-developed hind wings (Figures 44-45,

47-48, 50-51, 53-54, 56-57, 59-60, 62-63, 65-66)

with a long and well-developed costal projection

similar to that of Neohagenulus. The hind wings

of Borinquena (B.) contradicens are well developed
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(Figures 70-71) and similar to those of Neohagenulus and Hagenulus, but the hind wings of B. (B.)

carmencita are greatly reduced and without a costal

projection (Figures 68-69), while the hind wings

of B. (A.) traverae are entirely absent. The nymphs

of Hagenulus and Borinquena are morphologically

similar to the other West Indian genera. The

nymphs of Hagenulus, however, appear more

primitive than those of Borinquena. The denticles

on the tarsal claws of Hagenulus, as well as those of

Neohagenulus, Careospina, and Traverina, are

progressively larger apically (Figures 198-201), but

the denticles on the tarsal claws of Borinquena are

progressively larger apically, except the apical denticle is much larger (Figures 202-203). At least 3

small, equal-sized denticles occur on the anteromedian emargination of the labrum of Hagenulus,

as well as the labrum of Neohagenulus, Careospina,

and Traverina (Figures 178-191). Two large equalsized denticles and sometimes 1 to 3 small unequalsized denticles occur on the anteromedian

emargination of the labrum of Borinquena (Figures 192-196). It appears Hagenulus is more primitive than Borinquena.

Nymphs of two species of Hagenulus have mouthparts secondarily specialized for straining food from

the current. It appears that this specialized type of

feeding has evolved in situ since the West Indies

broke up into several isolated islands. Only the

Cuban species have such feeding modifications,

being fully developed in H. caligatus and partially

developed in H. morrisonae. Based on the highly

specialized mouthparts, Traver (1960) suggested

Hagenulus (as described by Morrison) is closely

related to the Central and South American genera

Hermanella, Ulmeritus, and Traverella, and to the

Asian genus, Choroterpides. The foregoing study

has shown that Hagenulus is most closely related

to the other West Indian genera. Preliminary investigation indicates that Hermanella and Traverella are closely related and both may be closely

related to Ulmeritus. Recent research by Peters (in

manuscript) shows Choroterpides is most closely

related to the Asian species of Choroterpes.

Adults of the six West Indian genera of Leptophlebiidae are extremely varied except for the venation of the adult fore wings. The nymphs of these

six genera, however, are all morphologically similar

except for the type of mouthparts in two species of

Hagenulus and the abdominal gills of Traverina.

These six genera are some of the best examples, in

the Leptophlebiidae, of differential rates of evolution between adults and nymphs. Based on similarity of nymphal morphology and the venation of the

adult fore wings, all six genera belong to the same

complex, and based on total morphology of the

nymphs and adults they appear to represent at least

two diverging lines of evolution.

No North American leptophlebiid genus appears

closely related to the West Indian genera, even

though Cuba is in close proximity to Florida. Many

groups of insects indicate a faunal connection between Florida and the West Indies. Berner (1950)

stated that although other species of insects have

become established both in Florida and in the

West Indies, the mayflies have been unable to do

this principally because of the ecological conditions

of southern Florida. He further stated that a great

majority of mayflies described from the West Indies are inhabitants of mountain streams, and even

if these mayflies were accidentally introduced into

Florida, establishment would be impossible in the

sandy bottomed streams. Almost the entire mayfly

fauna of Florida appears to be Nearctic in nature.

Families and genera of mayflies occurring in Florida are Siphlonuridae (Isonychia), Baetidae (Baetis,

Callibaetis, Centroptilum, Cloeon, Pseudocloeon),

Oligoneuriidae (Homoeoneuria), Heptageniidae

(Heptagenia, Pseudiron, Stenonema), Ametropodidae (Siphloplecton), Leptophlebiidae (Choroterpes, Habrophlebia, Habrophlebiodes, Leptophlebia, Paraleptophlebia), Ephemerellidae (Ephemerella), Tricorythidae (Tricorythodes), Behningiidae (Dolania), Ephemeridae (Ephemera, Hexagenia, Pentagenia), Polymitarcidae (Tortopus),

Neoephemeridae (Neoephemera), Caenidae (Brachycercus, Caenis), and Baetiscidae (Baetisca). Of

these only the widespread American genera Baetis,

Callibaetis, Centroptilum, Tricorythodes, and

Caenis occur in the West Indies.

Leptophlebiidae and Baetidae are the only families possessing endemic West Indian genera. Traver

(1938) established Cloeodes for three new species

from Puerto Rico. Based on published reports and

collections available to me, the remaining West

Indian mayfly fauna is composed of species of the

widespread genera Baetis, Callibaetis, Centroptilum,

Paracloeodes, Lachlania, Leptohyphes, Tricorythodes, Euthyplocia, and Caenis. All of these genera

occur in the Nearctic and Neotropical regions, ex-
NUMBER 62 31

cept Euthyplocia, which occurs only in the Neotropical region. Although representatives of the

families Siphlonuridae (Isonychia only), Heptageniidae (as a montane extension from the Nearctic), Ephemeridae, and Polymitarcidae have been

reported in the tropics of the New World, none of

these families are known to occur in the West

Indies.

Based on large collections of Leptophlebiidae

available to me from Central and South America,

no species of Neohagenulus, Borinquena, Hagenulus, Traverina, Farrodes, or Careospina occur in

Central or South America. However, Hagenulopsis

Ulmer (1919) and three undescribed genera appear

to be closely related to Borinquena. Hagenulopsis

occurs from Brazil northward to Honduras, while

the new genera are known from Northern Brazil.

Reared or associated nymphs and adults of these

four genera are available. The mouthparts, tarsal

claws, and abdominal gills of the nymphs of all

these genera are morphologically similar to Borinquena. The fore wing venation and tarsal claws of

the adults of these genera are also morphologically

similar. The hind wings are absent in all four of the

South American genera. Hagenulopsis and the undescribed genera possess long female ovipositors or

egg guides and the associated elongated male genital forceps and styliger plate. The penes of the

males are tubular and divided, with a subapical,

ventral spine on each lobe, much as those of Borinquena. Preliminary investigation indicates Hagenulopsis and the undescribed genera are apomorphic

to Borinquena. In the future, I plan to study the

relationships of these four genera to the West Indian genera and to other South American genera

in detail.

According to Ross (1967) many inter-American

tropical elements of the Trichoptera appear to

have evolved an unusual number of species in the

area around the Caribbean and adjacent parts of

Mexico and South America. The best evidence for

this regional evolution is the increasing number

of endemic caddisfly genera being found in the

West Indies that are closely related to tropical elements in the surrounding continental areas. Further, Ross (1967) stated it is becoming probable

that Central America and the West Indies have

been connected at times and broken up into several

isolated land masses at other times, at least since

mid-Cretaceous, and possibly with the fracturing

reaching its maximum degree during mid-Tertiary.

At the present the relationships of the West Indian

Leptophlebiidae clearly appear to be with those of

Central and South America.

Ross (1967) stated that each element of the Trichoptera of the inter-American and circum-Caribbean area has arisen from a temperate ancestor and

evolved into a primarily tropical lineage. Until all

genera of the Leptophlebiidae occurring in South

and Central America have been studied, the probable ancestor of the West Indian genera and related

continental genera is not known with certainty.

Resumen

Este trabajo es una revisi6n de la familia Leptophlebiidae (Ephemeroptera) de las Antillas. La

revisirin se ha basando en gran cantidad de ninfas

y adultos criados o asociados, colectados en varias

localidades de las Antillas.

Tres nuevos ge*neros, Farrodes, Careospina y

Traverina, son estableciodos; tambten los generos

Neohagenulus, Hagenulus, y Borinquena son considerados validos. Dos subg^neros, Borinquena s.s.

y Australphlebia, son establecidos dentro de Borinquena. El g^nero Farrodes contiene las nuevas

especies F. hyalinus de Jamaica, F. grenadae de

Grenada, y F. bimaculatus de Cuba. Careospina se

establece para las nuevas especies C. hespera y C.

minuta de Cuba, y C. annulata de Haiti. Traverina

es monodpico, e incluye a T. cubensis de Cuba.

Neohagenulus fu6 creado por Traver (1938) para

tres especies de Puerto Rico, N. julio, N. luteolus,

y N. tinctus. Eaton (1882) establecid el g^nero

Hagenulus para la especie cubana H. caligatus.

Posteriormente Banks (1924) describi6 otra especie,

H. eatoni de Haiti. En el presente trabajo se describen tres especies mas, H. jamaicensis de Jamaica,

H. morrisonae de Cuba, y H. rangelae de Puerto

Rico. La ninfa descrita por Morrison como de

Hagenulus se establece como perteneciente a la

especie Hagenulus caligatus. Se discute un probable

cambio evolutivo en los habitos alimenticios de las

ninfas de Hagenulus, desde alimentarse del periphyton fijo hasta llegar al proceso de colar materias

alimenticias de la corriente fluvial. Borinquena hie"

establecido por Traver (1938) para dos especies de

Puerto Rico, B. carmencita y B. contradicens. En el

presente trabajo se describe una especie mas, B.
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

traverae de Dominica en el subg£nero Australphlebia.

Se discute la filogenia y relaciones de estos g£neros antillanos. Basandose en similaridades de morfologia ninfal y venaci6n de las alas anteriores de

los adultos, estos seis g^neros pertenecen al mismo

complejo, pero basandose en la total morfologfa de

ninfas y adultos, aparentemente representan por lo

menos dos lineas evolutivas.

Literature Cited

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1924. Descriptions of New Neuropteroid Insects. Bulletin

of the Museum of Comparative Zoology, 65:421-455.

Berner, Lewis

1950. The Mayflies of Florida. University of Florida

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Demoulin, Georg

1954. Les Ephemeropteres Leptophlebiidae de Borneo.

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Eaton, Rev. A. E.

1882. An Announcement of New Genera of the Ephemeridae. Entomologist's Monthly Magazine, 18:

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1884 (1883-1888). A Revisional Monograph of Recent

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Kimmins, D. E.

1956. New Species of Ephemeroptera from Uganda.

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1919. The Mayfly Ovipositor, with Notes on Leptophlebia

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Needham, James G., and Helen E. Murphy

1924. Neotropical Mayflies. Bulletin Lloyd Library 24,

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1964. Two New Genera of Mayflies from the Ethiopian

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Ross, Herbert H.

1967. The Evolution and Past Dispersal of the Trichoptera. Annual Review of Entomology, 12:169-206.

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1938. Mayflies of Puerto Rico. Journal of Agriculture of

the University of Puerto Rico, 22:5-42.

1944. Notes on Brazilian Mayflies. Boletim do Museu

Nacional, Nova Serie, Zoologia, 22:1-52.

1959. Uruguayan Mayflies. Family Leptophlebiidae: Part

I, Revista de la Sociedad Uruguaya de Entomologia, 3:1-19.

1960. Uruguayan Mayflies. Family Leptophlebiidae: Part

II. Revista de la Sociedad Uruguaya de Entomologia, 4:19-26.

Ulmer, Georg

1909. Ephemerida von Madagaskar und den Comoren. In

A. Voeltzkow, Reise in Ostafrika in Jahren 1903-

1905, 2:365-368.

1919. Neue" Ephemeropteren. Archiv fiir Naturgeschichte,

85:1-80.

1920. Ubersicht uber die Gattungen der Ephemeropteren,

nebst bemerkungen uber einzelne Arten. Stettiner

Entomologische Zeitung, 81:97-144.

1932. Aquatic Insects of China. Article VI. Revised Key

to the Genera of Ephemeroptera. Peking Natural

History Bulletin, 7:195-218.

1939. Eintagsfliegen (Ephemeroptera) von den SundaInseln. Teil I. Archiv fiir Hydrobiologie, Supplement 16:443-580.
NUMBER 62 33

FIGURE 1.—Hagenulus caligatus Eaton, dorsal view of mature female nymph.
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Hagenulus morrisonae, new species, dorsal view of mature female nymph.

FIGURE 3.—Hagenulus species from Jamaica, dorsal view of mature female nymph.
NUMBER 62 35

22 25

FIGURES 4-26.—Adult fore and hind wings. Schematic wings, showing abbreviations of venational

terminology used in this paper: 4, fore wing; 5, hind wing. Farrodes hyalinus, new species: 6,

fore wing; 7, hind wing; 8, hind wing enlarged. Farrodes grenadae, new species: 9, fore wing;

10, hind wing; 11, hind wing enlarged. Farrodes bimaculatus, new species: 12, fore wing; IS,

hind wing; 14, hind wing enlarged. Traverina cubensis, new species, imago: 15, fore wing, $;

16, hind wing, $ ; 17, hind wing enlarged, $; 18, fore wing, $ ; 19, hind wing, 9 : 20, hind wing

enlarged, $. Careospina hespera, new species, $ imago: 21, fore wing; 22, hind wing; 23, hind

wing enlarged. Careospina rninuta, new species, $ imago; 24, fore wing; 25, hind wing; 26, hind

wing enlarged.
36

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

28

*-• '

37

39 41

44

47

FIGURES 27—48.—Adult fore and hind wings. Careospina annulata, new species, $ imago: 27,

fore wing; 28, hind wing; 29, hind wing enlarged. Careospina species, 9 subimago: 30, fore

wing; 31, hind wing; 32, hind wing enlarged. Neohagenulus julio Traver, imago: 33, fore wing,

$; 34, hind wing, $; 35, hind wing enlarged, $; 36, fore wing, $; 37, hind wing, 9 ', 38, hind

wing enlarged, 9. Neohagenulus tinctus Traver, $ imago: 39, hind wing; 40, hind wing enlarged. Neohagenulus luteolus Traver, $ imago: 41, hind wing; 42, hind wing enlarged.

Hagenulus caligatus Eaton, imago: 43, fore wing, $; 44, hind wing, $; 45, hind wing enlarged,

$; 46, fore wing, 9 ; 47, hind wing, 9 ; 48, hind wing enlarged, 9 .
NUMBER 62 37

70

FIGURES 49-72.—Adult fore and hind wings. Hagenulus morrisonae, new species, imago: 49, fore

wing, $; 50, hind wing, $ ; 51, hind wing enlarged, $; 52, fore wing, 9; 53, hind wing, 9 ; 54,

hind wing enlarged, $. Hagenulus eatoni Banks, $ imago: 55, fore wing; 56, hind wing; 57,

hind wing enlarged. Hagenulus jamaicensis, new species, imago: 58, fore wing, $; 59, hind

wing, $; 60, hind wing enlarged, $; 61, fore wing, 9; 62, hind wing, 9; 63, hind wing enlarged, 9 • Hagenulus rangelae, new species, $ imago: 64, fore wing; 65, hind wing; 66, hind

wing enlarged. Borinquena (B.) carmencita Traver: 67, fore wing; 68, hind wing; 69, hind wing

enlarged. Borinquena (B.) contradicens Traver: 70, hind wing; 71, hind wing enlarged. Borinquena (Australphlebia) traverae, new species: 72, fore wing.
FIGURES 73-85.—Genitalia of male imago. Farrodes hyalinus, new species: 73, ventral view; 74,

lateral view. Farrodes grenadae, new species: 75, ventral view; 76, lateral view. Farrodes btmaculatus, new species: 77, ventral view; 78, lateral view. Traverina cubensis, new species: 79, ventral

view. Careospina hespera, new species: 80, ventral view; 81, lateral view of penes. Careospma

minuta, new species: 82, ventral view. Careospina annulata, new species: 83, ventral view.

Neohagenulus julio Traver: 84, ventral view. Neohagenulus tinctus Traver: 85, ventral view.
NUMBER 62 39

102 103

FIGURES 86-103.—Genitalia of male imago. Neohagenulus luteolus Traver: 86, ventral view.

Hagenulus caligatus Eaton: 87, ventral view. Hagenulus morrisonae, new species: 88, ventral

view. Hagenulus jamaicensis, new species: 89, ventral view; 90, ventral view of penes enlarged.

Hagenulus rangelae, new species: 91, ventral view. Borinquena (B.) carmencita Traver: 92,

ventral view; 93, ventral view of penes enlarged. Borinquena (B.) contradicens Traver (after

Traver, 1938): 94, ventral view; 95, ventral view of penes enlarged. Borinquena (AustralphUbia)

traverae, new species: 96, ventral view. Claws of fore leg of male imago. 97, Farrodes hyalinus,

new species; 98, Traverina cubensis, new species; 99, Careospina hespera, new species; 100,

Neohagenulus julio Traver; 101, Hagenulus caligatus Eaton; 102, Borinquena (B.) carmencita

Traver; 103, Borinquena (AustralphUbia) traverae, new species.
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 104-120.—Ninth sternum of female: 104, Farrodes hyalinus, new species, imago; 105,

Traverina cubcnsis, new species, imago; 106, Careospina species, subimago; 107, Neohagenulus

julio Traver, imago; 108, Hagenulus caligatus Eaton, imago; 109, Borinquena (B.) carmencita

Traver, imago; 110, Borinquena (Australphlebia) traverae new species, subimago. Lateral view

of ovipositor or egg guide of female: 111, Traverina cubensis, new species, imago; 112, Careospina species, subimago; 113, Neohagenulus julio Traver, imago; 114, Hagenulus caligatus Eaton,

imago; 115, Hagenulus morrisonae, new species, imago; 116, Hagenulus eatoni Banks, imago;

117, Hagenulus jamaicensis, new species, imago; 118, Hagenulus species, subimago; 119, Borinquena (B.) carmencita Traver, imago; 120, Borinquena (Australphlebia) traverae, new species,

subimago.
NUMBER 62 41

125 126

mm

132

137

FIGURES 121-138.—Careospina hespera, new species: 121, terminal filaments of $ imago. Color

patterns of abdominal segments of imago. Farrodes grenadae, new species, $ imago: 122, dorsal

view; 123, lateral view. Farrodes bimaculatus, new species: 124, dorsal view of abdominal segments 6-10, $ imago. Traverina cubensis, new species, $ imago: 125, dorsal view; 126, lateral

view. Careospina hespera, new species: 127, lateral view of $ imago. Careospina minuta, new

species: 128, lateral view of $ imago. Hagenulus caligatus Eaton, $ imago: 129, dorsal view;

130, lateral view. Hagenulus morrisonae, new species, $ imago: 131, dorsal view; 132, lateral

view. Hagenulus eatoni Banks, $ imago: 133, dorsal view; 134, lateral view. Hagenulus jamaicensis, new species, $ imago: 135, dorsal view; 136, lateral view. Hagenulus rangelae, new

species, $ imago: 137, dorsal view; 138, lateral view.
42

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 139-147.—Labium of mature nymph: 139, Farrodes hyalinus, new species; 140, Traverina

species; 141, Careospina hespera, new species; 142, Neohagenulus julio Traver; 143, Hagenulus

caligatus Eaton (reduced to 70% of mandible proportions); 144, Hagenulus morrisonae, new

species (reduced to 70% of mandible proportions); 145, Hagenulus species; 146, Borinquena (B.)

carmencita Traver; 147, Borinquena (Australphlebia) traverae, new species.
NUMBER 62 43

FIGURES 148-163.—Ventral view of right maxilla of mature nymph: 148, Farrodes hyalinus, new

species; 149, Traverina species; 150, Careospina hespera, new species; 151, Neohagenulus julio

Traver; 152, Hagenulus caligatus Eaton (reduced to 70% of mandible proportions); 153, Hagenulus morrisonae, new species (reduced to 70% of mandible proportions); 154, Hagenulus species;

155, Borinquena (B.) carmtncita Traver; 156, Borinquena (Australphlebia) traverae, new species.

Hypopharynx of mature nymph: 157, Farrodes hyalinus, new species; 158, Traverina species;

159, Careospina hespera, new species; 160, Neohagenulus julio Traver; 161, Hagenulus caligatus

Eaton (reduced to 70% of mandible proportions); 162, Hagenulus morrisonae, new species (reduced to 70% of mandible proportions); 163, Hagenulus species.
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 164-175.—Hypopharynx of mature nymph: 164, Borinquena (B.) carmencita Traver; 165,

Borinquena (Australphlebia) traverae, new species. Left mandible of mature nymph: 166, Farrodes hyalinus, new species; 167, Traverina cubensis, new species; 168, Traverina species; 169,

Careospina hespera, new species; 170, Neohagenulus julio Traver; 171, Hagenulus caligatus

Eaton; 172, Hagenulus morrisonae, new species; 173, Hagenulus species; 174, Borinquena (B.)

carmencita Traver; 175, Borinquena (Australphlebia) traverae, new spedes.
NUMBER 62 45

179

AA/Wv

180

FIGURES 176-196.—Labrum of mature nymph. Farrodes hyalinus, new species: 176, labrum; 177,

enlargement of denticles on labrum. Traverina cubensis, new species: 178, labrum; 179, enlargement of denticles on labrum. Traverina species: 180, enlargement of denticles on labrum. Careospina hespera, new species: 181, labrum; 182, enlargement of denticles on labrum. Neohagenulus

julio Traver: 183, labrum. Neohagenulus luteolus Traver: 184, 185, enlargement of denticles on

labrum. Hagtnulus caligatus Eaton: 186, labrum (reduced to 50% of mandible proportions); 187,

enlargement of denticles on labrum. Hagenulus morrisonae, new species: 188, labrum (reduced

to 70% of mandible proportions); 189, enlargement of denticles on labrum. Hagenulus species:

190, labrum; 191, enlargement of denticles on labrum. Borinquena (B.) contradicens Traver:

192, labrum; 193, enlargement of denticles on labrum. Borinquena (B.) carmencita Traver: 194,

enlargement of denticles on labrum. Borinquena (Australphlebia) traverae, new species: 195,

labrum; 196, enlargement of denticles on labrum.
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

211 212

FIGURES 197-212.—Fore nymphal claw: 197, Farrodes hyalinus, new species; 198, Traverina species;

199, Careospina hespera, new species; 200, Neohagenulus julio Traver; 201, Hagenulus caligatus

Eaton; 202, Borinquena (B.) carmencita Traver; 203, Borinquena (Australphlebia) traverae, new

species. Nymphal abdominal gills: 204, Farrodes hyalinus, new species; 205, Traverina cubensis,

new species; 206, Careospina hespera, new species; 207, Neohagenulus julio Traver; 208, Hagenulus morrisonae, new species; 209, Borinquena (B.) carmencita Traver; 210, Borinquena (Australphlebia) traverae, new species. Dorsal view of nymphal abdominal segments 5-10: 211, Farrodes

hyalinus, new species; 212, Careospina hespera, new species.
Index

(Page numbers of principal accounts in boldface.)

Ametropodidae, 30

annulata, Careospina, 12, 14, 31

(Australphlebia) Borinquena, 2, 3, 4, 5, 25, 26, 27, 31, 32

Baetidae, 30

Raetis, 30

fiaetisca, 30

Baetiscidae, 30

Behningiidae, 30

bimaculatus, Farrodes, 6, 7, 8, 31

Borinquena, 1, 2, 3, 4, 5, 24, 25, 29, 30, 31

Borinquena (Australphlebia), 2, 3, 4, 5, 25, 26, 27, 31, 32

traverae, 24, 25, 26, 27, 28, 30, 32

Borinquena (Borinquena), 2, 3, 4, 5, 25, 26, 27, 31

carmencita, 24, 25, 26, 27, 30, 31

contradicens, 24, 25, 26, 27, 29, 31

(Borinquena) Borinquena, 2, 3, 4, 5, 25, 26, 27, 31

Brachycercus, 30

Caenidae, 30

Caenis, 30

caligatus, Hagenulus, 17, 18, 19, 20, 30, 31

Callibaetis, 30

Careospina, 2, 3, 4, 5, 11, 28, 29, 30, 31

annulata, 12, 14, 31

hespera, 12, 13, 31

minuta, 12, 13, 14, 31

carmencita, Borinquena (B.), 24, 25, 26, 27, 30, 31

Centroptilum, 30

Choroterpes, 30

Choroterpides, 30

Cloeodes, 30

Cloeon, 30

contradicens, Borinquena (B.), 24, 25, 26, 27, 29, 31

cubensis, Traveriva, 10, 31

Dolania, 30

duliti, Hagenulus (?), 18

eatoni, Hagenulus, 17, 18, 19, 20, 23, 31

Ephemera, 30

Ephemerella, 30

Ephemerellidae, 30

Ephemeridae, 30

Euthyplocia, 30

Farrodes, 2, 3, 4, 5, 6, 28, 29, 31

bimaculatus, 6, 7, 8, 31

grenadae, 6, 7, 8, 31

hyalinus, 6, 7, 31

fasciatus, Hagenulus, 18

grenadae, Farrodes, 6, 7, 8, 31

Habrophlebia, 30

Habrophlebiodes, 30

Hagenulus, 1, 2, 3, 4, 17,18, 28, 29, 30, 31

caligatus, 17, 18, 19, 20, 30, 31

duliti, 18

eatoni, 17, 18, 19, 20, 23, 31

fasciatus, 18

"Jamaica species," 18

jamaicensis, 17, 19, 23, 24, 31

karnyi, 18

monstratus, 18

morrisonae, 17, 19, 20, 21, 22, 23, 30, 31

rangelae, 17, 19,20,24,31

scotti, 18

turbinatus, 18

Heptagenia, 30

Heptageniidae, 30

Hermanella, 30

hespera, Careospina, 12, 13, 31

Hexagenia, 30

Homoeoneuria, 30

hyalinus, Farrodes, 6, 7, 31

Isonychia, 30

"Jamaica species," Hagenulus, 18

jamaicensis, Hagenulus, 17, 19, 23, 24, 31

julio, Neohagenulus, 15, 16, 17, 31

karnyi, Hagenulus, 18

Lachlania, 30

Leptohyphes, 30

Leptophlebia, 30

Leptophlebiidae, 1, 3, 4, 28, 30

luteolus, Neohagenulus, 15,16, 31

minuta, Careospina, 12, 13, 14, 31

monstratus, Hagenulus, 18

morrisonae, Hagenulus, 17, 19, 20, 21,22, 23, 30, 31

Neoephemera, 30

Neoephemeridae, 30

Neohagenulus, 1, 2, 3, 4, 5, 15, 28, 29, 30, 31

julio, 15, 16,17, 31

luteolus, 15, 16, 31

species 1, 16

species 2, 16

tinctus, 15, 16, 17, 31

Oligoneuriidae, 30

Paracloeodes, 30

Paraleptophlebia, 30

Pentagenia, 30

Polymitarcidae, 30

Pseudiron, 30

Pseudocloeon, 30

rangelae, Hagenulus, 17, 19, 20, 24, 31

scotti, Hagenulus, 18

Siphlonuridae, 30

47
48

Siphloplecton, 30

species 1, Neohagenulus, 16

species 2, Neohagenulus, 16

Stenonema, 30

tinctus, Neohagenulus, 15, 16, 17, 31

Tortopus, 30

traverae, Borinquena (A.), 24, 25, 26, 27,28, 30

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Traverella, 30

Traverina, 2, 3, 4, 9, 28, 29, 30, 31

cubensis, 10, 31

Tricorythidae, 30

Tricorythodes, 30

turbinatus, Hagenulus, 18

Ulmeritus, 30

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