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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 62

A Revision of the Leptophlebiidae of the West Indies (Ephemeroptera)

William L. Peters

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1971

A B S T R A C T

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.

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A Revision of the Leptophlebiidae of the West Indies (Ephemeroptera)

William L. Peters

____________

William L. Peters, Entomology, Florida Agricultural and Mechanical University, Tallahassee, Florida 32307.

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 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.

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

Keys to the Genera of the Leptophlebiidae from the West Indies

La clave los generos de Leptophlebiidae de las Antillas

Farrodes Peters, new genus

FIGURES 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 length 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 Rs 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 2½ 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 well-developed 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 one-third 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 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 ICu₁ 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 ICu₁ 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

Clave para las Especies de Farrodes Peters, nuevo género

Farrodes hyalinus Peters, new species

FIGURES 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 sunrise 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

FIGURES 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 77–78): 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., Güines (field no. 43), March 1966; allotype female imago, Cuba: La Habana Prov., Güines (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

FIGURES 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 one-third of distance from base to margin, fork symmetrical; cubital area as in Figure 15; cross veins few. Fore wings of female (Figure 18): vein Rs 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 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 ICu₁ 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

FIGURES 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 R₁, 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 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 Río Prov., Río Cuzco, near Soroa (field no. 57), March 1964; allotype female imago, Cuba: Pinar del Río Prov., Río Cuzco, near Soroa (field no. 25), March 1965; paratypes: 2 nymphs, Cuba: Pinar del Río Prov., Soroa (field no. 51), June 1965; 2 nymphs, Cuba: Pinar del Río Prov., Río 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: Río 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 two-thirds length of upper portion; eyes of female separated on meson of head by a length 3½ 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 Rs 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 2½ 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 well-developed lateral processes (Figure 159), anterior margin deeply cleft; superlingua of hypopharynx as in Figure 159, with a row of hair along anterior margin. Segment 2 of maxillary palpi 1¼ 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 one-fourth 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 ICu₁ 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 equal-sized 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

Clave para las Especies de Careospina Peters, nuevo género

Careospina hespera Peters and Alayo, new species

FIGURES 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 Río Prov., Río Cuzco, near Soroa (field no. 24), March 1965; paratopotypes: 4 male imagos; paratypes: 2 male imagos, Cuba: Pinar del Río Prov., Arroyo Pinar de Vinales (field no. 22); 2 nymphs, Cuba: Pinar del Río Prov., Río Cuzco, near Soroa (field no. 3), March 1965; 2 nymphs, Cuba: Pinar del Río Prov., Viñnales (field no. 11), October 1964; 3 nymphs, Cuba: Pinar del Río 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 minuta 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 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 Hagenulus 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 subimngo 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 3½ times length of penes. Caudal filaments light brown, annulations at articulations darker.

FEMALE IMAGO.—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 3½ times the length of the penes (Figure 83), and (3) the color pattern on the abdominal segments is as given in the description.

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. tinctus 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 Rs 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 Rs 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.

Key to the Species of Neohagenulus Traver

Clave para las Especies de Neohagenulus Traver

Genus Hagenulus Eaton, 1882

FIGURES 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 wings (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 2½ 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 equal-sized 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 1½ to 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 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, 1 ⅛ miles southwest crossing Dick's Pond Trail, Hard-war 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 intermedi ate 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 following 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 equal-sized, blunt denticles are present on the anteromedian emargination of the labrum (Figures 187, 189, 191).

Key to the Species of Hagenulus Eaton

Clave para las Especies de Hagenulus Eaton

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. 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 of 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., Güines (field no. 33), March 1966, P. Alayo D.; 1 male imago, Cuba: Pinar del Río Prov., Arroyo Pinar de Viñales (field no. 23), P. Alayo D.; I female subimago, Cuba: La Habana Prov., Güines (field no. 33), March 1966, P. Alayo D.; 9 nymphs, Cuba: Pinar del Río Prov., Río Cuzco, near Soroa (field no. 1), March 1965, P. Alayo D.; 1 nymph, Cuba: Pinar del Río Prov., Río Cuzco, near Soroa (field no. 2), March 1965, P. Alayo D.; 14 nymphs, Cuba: La Habana Prov., Güines (field no. 5), March 1966, P. Alayo D.; 3 nymphs, Cuba: Pinar del Río Prov., stream, Viñales (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., Güines (field no. 56), March 1966, P. Alayo D.; 2 nymphs, Cuba: Pinar del Río 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 Río Prov., Santa Cruz de los Pinos, 24–28 March 1939, J. G. Needham. AH 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 R₁ 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 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., Río Boniatico, San Luis (field no. 27), November 1964; allotype, Cuba: Las Villas Prov., Río 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., Río Anaya, Ciego Montero, Rodas (field no. 32), December 1965; 1 male imago, 1 male subimago, 1 female subimago, Cuba: Oriente Prov., Río Guantanamo (field no. 30); 2 nymphs, Cuba: Oriente Prov., Río Nagua, near Turquino (field no. 16), May 1965; 2 nymphs, Cuba: Oriente Prov., Río Yara, Central Estrada Palma, Manzanillo (field no. 17), November 1964; 4 nymphs, Cuba: Oriente Prov., Río Boniatico, San Luis (field no. 26), November 1964; 10 nymphs, same data as for allotype; 6 nymphs, Cuba: Oriente Prov., Río 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 R₁ 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 nymph's 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 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 1½ 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, 133–154

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 R₁ 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 R₁ 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 surrounded 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, Río 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 R₁, 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 R₁ 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 R₁ 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

FIGURES 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 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 Rs 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 well-developed 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.

Borinquena can be distinguished from other West Indian genera of the Leptophlebiidae by the following combination of characters. In the imago (1) the penes of the male genitalia are divided, tubular and a subapical ventral spine occurs on each penis lobe (Figures 92–96), (2) the female possesses a well-developed ovipositor or egg guide and it extends to at least past the anterior margin of abdominal segment 10 (Figures 119–120), and (3) the cross veins in the fore wings of the male and female are not surrounded with dark clouds (Figures 67, 72). In the nymphs (1) the denticles on the claws are progressively larger apically except the apical denticle is much larger (Figures 202–203), (2) 2 large equal-sized denticles and sometimes 1 to 3 small unequal-sized denticles occur on the anteromedian emargination of the labrum (Figures 193–194, 196), and (3) posterolateral spines occur on abdominal segments 6 or 7–9. Borinquena can be distinguished from all genera of the Leptophlebiidae by the following combination of characters. In the imagos (1) the base of vein ICu₁ of the fore wings attaches to vein CuA (Figures 67, 72), (2) the penes of the male genitalia are divided, tubular and a subapical ventral spine occurs on each penis lobe (Figures 92–96), (3) the female possesses a well-developed ovipositor or egg guide and it extends to at least past the anterior margin of abdominal segment 10 (Figures 119–120), and (4) the length of the styliger plate of the male along the median line is one-third to half the length of the maximum width (Figures 92, 94, 96). In the nymphs (1) a V-shaped ridge occurs near the ventral, inner anterolateral margin of the maxillae (Figures 155–156), (2) 2 large equal-sized denticles and sometimes 1 to 3 small unequal-sized denticles occur on the anteromedian emargination of the labrum (Figures 193–194, 196), (3) the denticles on the claws are progressively larger apically except the apical denticle is much larger (Figures 202–203), and (4) posterolateral spines occur on abdominal segments 6 or 7–9.

Key to the Species of Borinquena Traver

Clave para las Especies de Borinquena Traver

Subgenus Borinquena sensu stricto Traver, 1938

FIGURES 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

FIGURES 72, 96, 103, 110, 120, 147, 156, 165, 175, 195–196, 203, 210

IMAGO.—Hind wings absent. Female with a well-developed 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

FIGURES 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 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; paratypes, 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 labium (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 111–113); and a long and well-developed ovipositor or egg guide is present in the remaining genera (Figures 114–120). The females of all genera except Farrodes possess a ninth 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 (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 equal-sized denticles and sometimes 1 to 3 small unequal-sized 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, except 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 revisión de la familia Leptophlebiidae (Ephemeroptera) de las Antillas. La revisión se ha basando en gran cantidad de ninfas y adultos criados o asociados, colectados en varias localidades de las Antillas.

Tres nuevos géneros, Farrodes, Careospina y Traverina, son estableciodos; también los generos Neohagenulus, Hagenulus, y Borinquena son considerados validos. Dos subgéneros, Borinquena s.s. y Australphlebia, son establecidos dentro de Borinquena. E1 género 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 monotípico, e incluye a T. cubensis de Cuba.

Neohagenulus fué creado por Traver (1938) para tres especies de Puerto Rico, N. julio, N. luteolus, y N. tinctus. Eaton (1882) estableció el género Hagenulus para la especie cubana H. caligatus. Posteriormente Banks (1924) describió 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 hábitos alimenticios de las ninfas de Hagenulus, desde alimentarse del periphyton fijo hasta llegar al proceso de colar materias alimenticias de la corriente fluvial. Borinquena fué establecido por Traver (1938) para dos especies de Puerto Rico, B. carmencita y B. contradicens. En el presente trabajo se describe una especie más, B. traverae de Dominica en el subgénera Australphlebia.

Se discute la filogenia y relaciones de estos géneros antillanos. Basándose en similaridades de morfologia ninfal y venación de las alas anteriores de los adultos, estos seis géneros pertenecen al mismo complejo, pero basándose en la total morfología de ninfas y adultos, aparentemente representan por lo menos dos líneas evolutivas.

Literature Cited

Banks, Nathan

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 Studies, Biological Science Series, 4(4):xii+267 pages.

Demoulin, Georg

1954. Les Ephéméroptères Leptophlebiidae de Bornéo. Bulletin et Annales de la Société Entomologique de Belgiqueé, 90:123–131.

Eaton, Rev. A. E.

1882. An Announcement of New Genera of the Ephemeridae. Entomologist’s Monthly Magazine, 18: 207–208.

1884 (1883–1888). A Revisional Monograph of Recent Ephemeridae or Mayflies. Transactions of the Linnean Society of London (2nd series, Zoology), 3:1–352.

1892. New Species of Ephemeridae from the Tenasserim Valley. Transactions of the Entomological Society of London, 1892(2):185–190.

1913. The Percy Sladen Trust Expedition to the Indian Ocean in 1905, under the Leadership of Mr. Stanley Gardiner, M.A. No. 26, Ephemerida. Transactions of the Linnean Society of London (Zoology), 15: 433–434.

Kimmins, D. E.

1956. New Species of Ephemeroptera from Uganda. Bulletin of the British Museum (Natural History), Entomology, 4:69–87.

Morrison, Emily R.

1919. The Mayfly Ovipositor, with Notes on Leptophlebia and Hagenulus. Canadian Entomologist, 51:139–146.

Needham, James G., and Helen E. Murphy

1924. Neotropical Mayflies. Bulletin Lloyd Library 24, Entomological Series, 4:1–79.

Peters, William L., M. T. Gillies, and George F. Edmunds, Jr.

1964. Two New Genera of Mayflies from the Ethiopian and Oriental Regions (Ephemeroptera: Leptophlebiidae). Proceedings of the Royal Entomological Society of London (B), 33:117–124.

Ross, Herbert H.

1967. The Evolution and Past Dispersal of the Trichoptera. Annual Review of Entomology, 12:169–206.

Traver, Jay R.

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 Série, 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 für Naturgeschichte, 85:1–80.

1920. Übersicht über die Gattungen der Ephemeropteren, nebst bemerkungen über 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 Sunda-Inseln. Teil I. Archiv für Hydrobiologie, Supplement 16:443–580.

Figures

FIGURE 1.—Hagenulus caligatus Eaton, dorsal view of mature female nymph.

FIGURE 2.—Hagenulus morrisonae, new species, dorsal view of mature female nymph.

FIGURE 3.—Hagenulus species from Jamaica, dorsal view of mature female nymph.

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; 13, hind wing; 14, hind wing enlarged. Traverina cubensis, new species, imago: 15, fore wing, art; 16, hind wing, art; 17, hind wing enlarged, art; 18, fore wing, art; 19, hind wing, art; 20, hind wing enlarged, art. Careospina hespera, new species, art imago: 21, fore wing; 22, hind wing; 23, hind wing enlarged. Careospina minuta, new species, art imago; 24, fore wing; 25, hind wing; 26, hind wing enlarged.

FIGURES 27–48.—Adult fore and hind wings. Careospina annulata, new species, art imago: 27, fore wing; 28, hind wing; 29, hind wing enlarged. Careospina species, art subimago: 30, fore wing; 31, hind wing; 32, hind wing enlarged. Neohagenulus julio Traver, imago: 33, fore wing, art; 34, hind wing, art; 35, hind wing enlarged, art; 36, fore wing, art; 37, hind wing, art; 38, hind wing enlarged, art. Neohagenulus linctus Traver, art imago: 39, hind wing; 40, hind wing enlarged. Neohagenulus luteolus Traver, art imago: 41, hind wing; 42, hind wing enlarged. Hagenulus caligatus Eaton, imago: 43, fore wing, art; 44, hind wing, art; 45, hind wing enlarged, art; 46, fore wing, art; 47, hind wing, art; 48, hind wing enlarged, art.

FIGURES 49–72.—Adult fore and hind wings. Hagenulus morrisonae, new species, imago: 49, fore wing, art; 50, hind wing, art; 51, hind wing enlarged, art; 52, fore wing, art; 53, hind wing, art; 54, hind wing enlarged, art. Hagenulus eatoni Banks, 9 imago: 55, fore wing; 56, hind wing; 57, hind wing enlarged. Hagenulus jamaicensis, new species, imago: 58, fore wing, art; 59, hind wing, art; 60, hind wing enlarged, art; 61, fore wing, art; 62, hind wing, art; 63, hind wing enlarged, art. Hagenulus rangelae, new species, art 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: 7S, ventral view; 74, lateral view. Farrodes grenadae, new species: 75, ventral view; 76, lateral view. Farrodes bimaculalus, new species: 77, ventral view; 78, lateral view. Traverina eubensis, new species: 79, ventral view. Careospina hespera, new species: 80, ventral view; 81, lateral view of penes. Careospina 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.

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 (Australphlebia) 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 (Australphlebia) traverae, new species.

FIGURES 104–120.—Ninth sternum of female: 104, Farrodes hyalinus, new species, imago; 105, Traverina cubensis, 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.

FIGURES 121–138.—Careospina hespera, new species: 121, terminal filaments of art imago. Color patterns of abdominal segments of imago. Farrodes grenadae, new species, art imago: 122, dorsal view; 123, lateral view. Farrodes bimaculatus, new species: 124, dorsal view of abdominal segments 6–10, art imago. Traverina cubensis, new species, art imago: 125, dorsal view; 126, lateral view. Careospina hespera, new species: 127, lateral view of art imago. Careospina minula, new species: 128, lateral view of art imago. Hagenulus caligatus Eaton, art imago: 129, dorsal view; 130, lateral view. Hagenulus morrisonae, new species, art imago: 131, dorsal view; 132, lateral view. Hagenulus eatoni Banks, art imago: 133, dorsal view; 134, lateral view. Hagenulus jamaicensis, new species, art imago: 135, dorsal view; 136, lateral view. Hagenulus rangelae, new species, art imago: 137, dorsal view; 138, lateral view.

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.

FIGURES 148–163.—Ventral view of right maxilla of mature nymph: 148, Farrodes hyalinus, new species; 149, Travtrina species; 150, Careospina hespera, new species; 151, Neohagenulus julio Traver; 152, Hagenulus caligatus Eaton (reduced to 70% of mandible proportions); 155, Hagenulus morrisonae, new species (reduced to 70% of mandible proportions); 154, Hagenulus species; 155, Borinquena (B.) carmencita Traver; 156, Borinquena (Australphlebia) Iraverae, 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.

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 species.

FGURES 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. Hagenutus 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.

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; 20S, 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

Baetis, 30

Baetisca, 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

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

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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