Re ‘§ iy SS tet eta te! REE Ahh ic ssn agagere aman Raia ce yeah HCI te nha . i i inguin it ieee Heh Mt} nea q bia oy ii i i bee a }) 7 O i a fi \ Wf] - SMITHSONIAN INSTITUTION UNITED STATES NATIONAL MUSEUM PROCEEDINGS OF THE UNITED STATES NATIONAL MUSEUM VOLUME 120 NUMBERS 3554-3567 SMITHSONIAN PRESS WASHINGTON : 1967 Publications of the United States. National Museum The scientific publications of the United States National Museum include two series, Proceedings of the United States National Museum and United States Na- tional Museum Bulletins. In these series are published original articles and monographs dealing with the collections and work of the Museum and setting forth newly acquired facts in the fields of anthropology, biology, geology, history, and technology. Copies of each publication are distributed to libraries and scientific organizations and to specialists and others interested in the various subjects. The Proceedings, begun in 1878, are intended for the publication, in separate form, of shorter papers. These are gathered in volumes, octavo in size, with the publication date of each paper recorded in the table of contents of the volume. In the Bulletin series, the first of which was issued in 1875, appear longer, separate publications consisting of monographs (occasionally in several parts) and volumes in which are collected works on related subjects. Bulletins are either octavo or quarto in size, depending on the needs of the presentation. Since 1902, papers relating to the botanical collections of the Museum have been published in the Bulletin series under the heading Contributions from the United States National Herbarium. FRANK A. TAYLOR Director, United States National Museum II CONTENTS Bram, Ratpeu A. Classification of Culex subgenus Culex in the New World (Diptera: Culicidae). 122 pages, 33 figures. Published March 6, 1967. Capps, Haun W. Review of some species ai eee Stings and descriptions of new species (Lepidoptera, Pyraustidae: Pyraustinae). 75 pages, 104 figures, 7 plates. Published March 15, 1967 Be Mk Be EE on sinc c, ve ee New species: Loxostege pseudohelvialis, L. arizonensis, L. neohelvialis, L. nayaritensis, L. polingi, L. pelotasalis, L. boliviensis, L. saltensis, L. fordi, L. alpinensis, L. neomarculenta, L. pseudobliteralis, L. neobliteralis, L. potosiensis, L. cayugalis, L. ecuadoralis, L. neotro- picalis, L. cochisensis, L. jacalensis, L. yucatanalis, L. corozalis, L. huachucalis, L. marialis, L. purulhalis, L. conisphoralis, L. jaliscalis, L. volcanensis, L. vendalis, L. federalis, L. sinaloensis, L. chiapasalis, L. mojavealis, L. peruensis, L. brasiliensis, L. piuralis. New combinations: Lozostege cynoalis, L. jaralis, L. ramsdenalis, L. conisphora, L. mellinialis, L. intinctalis, L. entephrialis, L. cupreicostalis, L. subcostalis, L. nigripes, L. subcuprea, L. clarissalis, L. aemulalis, L. naranjalis, L. fuscivenalis, L. protealis, L. eneanalis, Ostrinia obliteralis, O. penitalis, Phlyctaenodes parvipicta. Eset, Ropert KE. Amblyceran Mallophaga (biting lice) found on the Bucerotidae (hornbills). 76 pages, 72 figures. Pub- lished February 16, 1967 . New genus: Bucerocolpocephalum. New species: Chapinia fasciati, C. camuri, C. waniti, C. malayensis, C. clayae, C. hoplai, C. boonsongi, C. traylori, C. wenzeli, C. blakei, C. lydae, C. muesebecki, Bucerocolpocephalum emersoni, B. deignant. FLEMINGER, ABRAHAM. Taxonomy, distribution, and _poly- morphism in the Labidocera jollae group with remarks on evolution within the group (Copepoda: Calanoida). 61 pages, 17 figures. Published April 18, 1967. . New species: Labidocera kolpos, L. diandra. Furnt, Ourver S., Jr. Studies of Neotropical caddis flies, II: Types of some species described by Ulmer and Brauer. 20 pages, 5 figures, 2 plates. Published December 30, 1966. New combinations: Mezitrichia albolineata, Oecetis punctipennis. Hanp, Capxt, and BusHneiu, Raven. A new species of burrow- ing acontiate anemone from California (Isophelliidae: Flosmaris). 8 pages, 2 figures. Published March 8, 1967.. New species: Flosmaris grandis. Number 3557 3561 3558 3567 3554 IV CONTENTS Kornicker, Louis S. Euphilomedes arostrata, a new myodo- copid ostracod from Maldive Islands, Indian Ocean. 21 pages, 10 figures. Published February 15, 1967__-______- New species: Huphilomedes arostrata. Kornicker, Louis S. Supplementary description of the myodocopid ostracod Huphilomedes multichelata from the Great Bahama Bank. 16 pages, 6 figures. Published Feb- ruary 16, 1967 . : eye Semone Reisu, Donaup J., and Bee eer i Tcsiing. The benthic Polychaeta and ean eda of Morro Bay, California. 26 pages, 1 figure. Published March 8, 1967 . ScuuttTz, Leonarp P. Review of South American peace tee angelfishes—genus Pterophyllum. 10 pages, 4 plates. Pub- lished February 16, 1967 STEPHENSON, WitiiaAm. A dotmnarison Gi ist ates and American specimens of Hemisquilla ensigera (Owen, 1832) (Crustacea: Stomatopoda). 18 pages, 3 en Published March 8, 1967 d New subspecies: Hemisquilla ensigera ensigera, ‘HL é. Lalbvalietered H. e. australiensis. SterHrenson, WixuiAM, and Regs, May. Some portunid crabs from the Pacific and Indian Oceans in the collections of the Smithsonian Institution. 114 pages, 38 pean! 9 Pee Published March 8, 1967. . New species: Charybdis (Charybdis) pees pian! acerbiter- minalis, P. spiniferus, Thalamita multispinosa, T. philippinensis, T. pseudopoissoni, T. spinimera. StryskaL, Grorce C. Notes on flies captured in treetops in Malaya (Diptera: Empididae, Neriidae, Platystomatidae, Sepsidae, Muscidae). 16 pages, 4 Tee Published De- cember 30, 1966 New species: Hybos es H. meracrus, Pie neme (Meringomeria) leytensis, Dicranosepsis coryphea, Fraserella al- tivolans. New combination: Dichaetomyia prolixa. New name: Genus Fraserella. Topp, E.L. Notes and descriptions of some Neotropical agar- istine moths (Lepidoptera: Nostuidae). 15 pages, 4 plates. Published December 30, 1966. ember ute New genus: Cisaucula. New species: Caularis jamaicensis. New subspecies: J'uerta sabulosa collectiora. New combination: Cisaucula peruviana. 3563 3566 3565 3555 3564 3556 3562 3560 Erocee dines of wae, nited States National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. Volume 120 1967 Number 3554 A NEW SPECIES OF BURROWING ACONTIATE ANEMONE FROM CALIFORNIA (ISOPHELLITDAE: FLOSMARIS) By CapEt Hanp and Raven BusHNeELv! In a series of papers, Hand (1955, 1955a, 1956, 1958) described certain of the sea anemones of central California. The new species herein considered adds to that list. The first specimens were collected in 1949 from intertidal mud flats of San Francisco Bay adjacent to Bay Farm Island, Alameda County, California. The flat is composed of fairly firm sand in which is mixed considerable mud and accumulated mollusk shells in various stages of disintegration. The anemones occur with the base attached to fair sized remains of clam shell, small stones or even pieces of wood some 30-46 cm. beneath the surface. The oral end extends to the surface. Thus, when located in their original habitat, most of the 70 animals so far found, were about 30-46 cm. in length; however, they are only about 6-7 mm. in diameter except at the disc and base. The base may be up to 2.5 cm. in diameter and well covered with bits of clam shell and much sand, which remain adhering when the animal is dug from its burrow. Much care is needed in digging this species from its natural position as it is very easy to misjudge the depth at which the base is located, 1 Both authors: Department of Zoology, University of California, Berkeley, Calif. al 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 and, as frequently happens, the specimen may be cut in two before the base is freed from its attachment. The area inhabited by this species is between the midintertidal and the mean lower-low water level, and they are most easily obtained by searching for them when there is a good minus tide. At these times they are commonly observed with their tentacles extended at the entrance to their burrow, but any movement near them will cause them to contract so that the body is drawn down into the burrow. As one starts to dig for them, they continue to contract until they reach one-third or less of their normal length. When the collector is successful in dislodging them he retrieves an anemone appearing to be about 10-15 cm. long and about 2 cm. in diameter at its widest part (exclusive of the foot). Occasionally, when too much pressure is applied against them in their burrows they will be seen to emit tiny jets of water from cinclis-lke openings in the upper column. We do not believe that this is a natural reaction or one that would occur frequently in the absence of abnormal pressure. Weak areas evidently occur in the upper part of the column, but histologically these do not appear to be typical cinclides. So far in the laboratory, we have not been able to relax specimens to anywhere near their normal length, i.e., 30-46 cm. On preserva- tion they tend to shorten to 15 cm. or less. We wish to thank both Mr. Marshall and Mr. Holleman for making the observations reported herein available to us. This work was supported by a grant from the National Science Foundation, GB 1237, and by a contract with the Atomic Energy Commission, AT (11—1)-34, Project #96. Family Isophelliidae IsOPHELLIIDAE Stephenson, 1935, p. 183. Thenaria (Acontiaria). Sphincter mesogloeal. Mesenteries divisi- ble into macro- and microcnemes; the older microcnemes may, however, be provided with filaments and acontia. Acontia with two categories of nematocysts, basitrichs, and microbasic amastigophores. Genus Flosmaris Flosmaris Stephenson, 1920, p. 441. Isophelliidae with the elongate column divisible into scapus and scapulus, the former with tenaculi. Cinclides present or possibly absent. Margin tenaculate. Tentacles simple, their longitudinal muscles ectodermal as are the radial muscles of oral disc. Sphincter mesogloeal; situated in upper part of scapulus. Twelve pairs of macrocnemes bearing gonads, though number may vary as result of NO. 3554 ANEMONE—HAND AND BUSHNELL 3 asexual reproduction. Macrocnemes all perfect and with strongly restricted to circumscribed retractors. Microcnemes weak, consisting chiefly of parietal muscle. Acontia present on macrocnemes and sometimes on some microcnemes. Cnidom: spirocysts, basitrichs, microbasic amastigophores; also atrichs and holotrichs if catch- tentacles are present. The generic diagnosis above is the one given by Hand (1961). It differs slightly from that of Stephenson (1920) and incorporates cer- tain modifications resulting from the inclusion of Flosmaris bathamae Hand (1961) in this genus. Flosmaris grandis, new species FicuRss 1, 2 Base: Generally circular, adherent and usually larger than scapus. Basilar muscles present and well developed. Column: Elongate, divided into scapus and scapulus, cylindrical and up to 20 cm. long and 2 cm. diameter in the preserved condition. In living condition within their burrows, column is up to 46 cm. long. Diameter of column tapers from less than 0.5 cm. just above base to about 1.5 cm. in region of scapulus in a large individual. Removed from natural surroundings, column usually expanded in upper two-thirds, tapering to meet flaring base, and resembling long, slender goblet (fig. 1). Column translucent or pale white, depending on degree of expansion or contraction of individual. Mesenterial insertions visible as pale white lines. Scapulus short, usually less than one-fifth of total column, some- what more transparent than scapus and thereby distinguished from the latter. Histological preparations show a thinning of all layers in the scapulus. No cinclides or tenaculi present in scapulus and there appear to be fewer gland cells in scapus. Scattered tenaculi present in upper third of scapus, but sand grains rarely found attached to them. Weakened areas somewhat compa- rable to poorly developed cinclides present in upper part of scapus. Mesogloea of column about of the same thickness as endoderm but only about 0.8 as thick as ectoderm. Mesogloeal sphincter (fig. 2a) fairly well developed at top of the scapulus and immediately adjacent to tentacle bases. Circular, endodermal muscles present throughout length of column but not extensively developed. Tentacles and Oral Disc: All the specimens so far examined have only regular tentacles, catch tentacles being unobserved. From 90 to 144 tentacles have been counted on various individuals. Inner tentacles usually hexamerously arranged. Individual tentacles extend to 1.6 cm. in length, and total crown may attain a diameter of as much as 3.5 cm. Tentacles thin, transparent, and gently tapered. 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Only a small amount of coloration is found on animal and is in bars or incomplete rings of white granules against a dark background at bases of tentacles. Lips seldom seen raised above level of disc and are smooth and unpigmented. Tentacles fully retractile and have ectodermal longitudinal muscles. Radial musculature of disc is ectodermal also. Mesenteries: In upper part of body of perfect specimens there are 12 pairs of complete and fertile macrocnemes, including 2 pairs of directives that correspond to the 2 siphonoglyphs, and 36 pairs of Ficure 1.—Flosmaris grandis: a, d-f, live specimens removed from their burrows; ), enlargement of the oral disc and tentacles; c, relaxed and extended preserved specimen. NO. 3554 ANEMONE—HAND AND BUSHNELL 5 microcnemes (fig. 2b). In scapus, especially lower parts, number of microcnemes decreases, and there are approximately half as many mesenteries at base as in oral end (fig. 2c). Dissections of 12 speci- mens and sections of 6 showed from 8 to 12 pairs of mesenteries may be perfect, fertile macrocnemes (fig. 2d). No form of asexual repro- duction has been observed in this species. Ficure 2.—Flosmaris grandis: a, sphincter muscles within the mesogloea; b, cross section of mesenteries in region of actinopharynx; ¢, cross section of column just above pedal disc; d, mesenteries in midregion of the column, 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 The 6 pairs of macrocnemes of first cycle have large restricted retractors (fig. 2b) running throughout their length, while second cycle has only slightly smaller restricted retractors. Directive re- tractors tend to be circumscript. There is a moderately well-developed parietal musculature on macrocnemes, while microcnemes (fig. 26) consist only of parietal muscles, there being no mesenterial sheet. A trifid mesenterial filament and acontium are present on each mac- rocneme, but were not observed on any microcneme. Cnidom (spirocysts, atrichs, basitrichs, and microbasic amastigo- phores) : tentacles microns spirocysts 17-22 3.0—4.0 basitrichs 19-27 X 3.0-4.0 basitrichs 14-16 X 2.5-3.0 scapus basitrichs 18-26 X 3.0-3.5 basitrichs 9-14 2.0-2.5 atrichs (rare) 8-10 3.0 actinopharynz basitrichs 26-34 X 5.0-6.0 basitrichs (rare) 18-20 4.0-5.0 microbasic amastigophores 16-20 X 5.0-6.0 filaments basitrichs 24-29 X 3.0-4.0 basitrichs (rare) 12-16 X 2.0-3.0 microbasic amastigophores 14-18 X 4.5-6.0 acontia basitrichs 20-29 X 2.5-3.0 basitrichs (rare) 16-18 X 2.0—-2.5 microbasic amastigophores 33-42 X 5.0-7.0 Type locality: Bay Farm Island, Alameda Co., Calif. Holotype: Deposited in the United States National Museum as no. 52592. Paratypes: Five additional specimens are deposited in the United States National Museum as nos. 52593 and 52594. Discussion The new species described herein is abundant in the intertidal zone of many parts of San Francisco Bay and also occurs in the estuary separating Alameda from Oakland, Calif. Flosmaris grandis is only the third species assigned to this genus, the others being the type species, I’. phellioides Stephenson, 1920, and F. bathamae Hand, 1961. F’. phellioides was described from a single specimen collected at Hulule Island, N. Male Atoll, Maldive Archipelago, and F. bathamae is known only from Otago Harbor in southern New Zealand. F. bathamae is a small species, the maximum length being about 2.5 cm., NO. 3554 ANEMONE—HAND AND BUSHNELL 7 while F’. grandis is some 18 times longer than this. The size of F. phellroides is not known. ‘There is some question, certainly, whether F. bathamae and F. grandis are cogeneric with F’. phellioides. Stephen- son’s original description of F’. phellioides is very brief and no diagnosis of nematocysts was given. ‘The new species agrees with the type species in having more mesenteries distally than proximally, but differs from F’. bathamae in this respect. Also, no catch tentacles have been observed in /’. grandis or F. phellioides, but they are present in F. bathamae. F. grandis resembles F. bathamae in its restricted re- tractors whereas [’. phellioides is reported to have circumscript retrac- tors. For the time being it seems most reasonable to treat all these species as members of the genus Flosmaris, and what is most needed is fresh material of /’. phellioides. When F. phellioides is rediscovered it should be possible to make a more detailed comparison of these species and, hopefully, to understand better the nature of this genus. Some interesting observations on F’. grandis in captivity have been made by Mr. Edward Arthur Marshall, a student at Oakland City College. We received these observations through Mr. John Holle- man, a biology instructor at the College. Mr. Marshall has kept individuals of this species in unaerated aquaria for several months and has observed that when detached from the substrate and placed on sand in water that the animals actively burrow and reattach. Burrowing is apparently accomplished by extensions of the pedal disc and undulations of the body. After attaching at a given point on the bottom of an aquarium with a 6.5 cm. deep layer of sand, movement from place to place occurs, and individuals were recorded as having moved a distance of 13 cm. overnight. The captive animals accepted a wide variety of foods including almost any small, soft-bodied organism. They readily accept small flies, the larvae of several insects, gnats, small spiders, small gastropods, and land slugs, brine shrimps, copepods, and small marine organisms in general, but refuse earwigs and small beetles. When food objects are dropped near an anemone, they extend the upper part of the body from their burrow toward the food. Successful capture of the food usually follows. When an injured, but living, house fly is placed on the surface of the aquarium, the anemone actively extends toward the surface of the aquarium, and captures it with its tentacles. This occurs only when the fly is still active and moving but trapped in the surface film. One might wonder if this last described behavior occurs in nature, and while no absolute answer is possible, it is readily demonstrable that many, many insects are indeed blown into the Bay. The behavior described may then be one which is adaptively valuable to the species and a perfectly normal bit of behavior as well. § PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Summary A new species of Flosmaris, F. grandis is described. This is the third species known in this genus. The genus is not well understood, and the type species, Ff’. phellioides from the Maldive Islands needs to be restudied. Some observations on the living animals are reported, and the capture of food on the surface of an aquarium is described. Literature Cited Hanp, CapDEtT 1955. The sea anemones of central California, I: The corallimorpharian and athenarian anemones. Wasmann Journ. Biol., vol. 12, no. 3, pp. 345-375. 1955a. The sea anemones of central California, Il: The endomyarian and mesomyarian anemones. Wasmann Journ. Biol., vol. 13, no. 1, pp. 37-99. 1956. The sea anemones of central California, III: The acontiarian ane- mones. Wasmann Journ. Biol., vol. 13, no. 2, pp. 189-251. 1958. Another sea anemone from California and the types of certain Cali- formian anemones. Washington Acad. Sci., vol. 47, no. 12, pp. 411-414. 1961. Two new acontiate New Zealand sea anemones. Trans. Soc. New Zealand. Zool., vol. 1, no. 4, p. 75-89. STEPHENSON, THomas A. 1920. On the classification of Actiniaria, I: Forms with acontia and forms with a mesogloeal sphincter. Quart. Micro. Sci. vol. 64, no. 4, pp. 425-574. 1935. The British sea anemones, vol. 11 (Ray Soe. vol. 121), ix + 426 pp. U.S. GOVERNMENT PRINTING OFFICE:1966 Proceedings of mae United, States National Museum SMITHSONIAN INSTITUTION ¢« WASHINGTON, D.C. ————— eee eee Volume 120 1967 Number 3555 REVIEW OF SOUTH AMERICAN FRESHWATER ANGELFISHES—GENUS PTEROPHYLLUM By Lronarp P. ScHuttz Senior Scientist, Department of Vertebrate Zoology The illustration of the new Plataroides leopoldi Gosse (1963, Inst. Roy. Sci. Nat. Belgique Bull., vol. 39, no. 35, pp. 4-6, pl. 1, fig. 2) so closely resembled the illustration of Plataxoides dumerilii Castelnau (1855, Animaux nouveaux ou rares . . . d’ Amérique du Sud, p. 21, pl. 11, fig. 3) that I decided to review the taxonomic status of all the nominal species of Pterophyllum. I shall attempt (1) to clarify the nomenclatural validity of the generic names Pterophyllum and Pla- taxoides, (2) to review the species complexes of Pterophyllum scalare (=P. evmeker) and Plataxoides dumerilui (=P. leopoldi,), and (3) to discuss the relationship between Pterophyllum altum Pellegrin and P. scalare (Lichtenstein). I have studied all nonaquarium specimens of Pterophyllum available in the following museums: British Museum of Natural History (BMNH); California Academy of Sciences (CAS); Chicago Natural History Museum (CNHM); Museum of Comparative Zoology, Harvard (MCZ); Museum National Histoire Naturelle, Paris (MNHN); Institut Royal des Sciences Naturelles de Belgique, Bruxelles (IRSNB); Stanford University (SU); U.S. National Museum (USNM). To the authorities of the above institutions, I wish to express my gratitude for their help and other courtesies. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 In the ‘‘Tropical Fish Hobbyist” (1953, vol. 1, no. 5, pp. 5-7, 20) I attempted to distinguish three species of aquarium-bred Pterophyllum. Since then I have studied specimens collected from their natural habitat, the syntypes of Pterophyllum altum Pellegrin, a syntype of P. eimekei Ahl, 27 paratypes of Plataxoides leopoldi Gosse, and the holotype of P. dumerilit Castelnau. In my 1953 study I recognized Pterophyllum scalare, P. eimeket, and P. altum as valid species; however, with additional material my conclusions have changed and in this review I now recognize as valid P. scalare, P. altum, and P. dumerilu. P. eimekei is a synonym of P. scalare. In the synonomy for each species I have recorded only the most pertinent references, omitting those related to aquaria. I have attempted to record each locality from which Pterophyllum has been reported by latitude and longitude as found in ‘“The Times Atlas of the World,” 1957, vol. 5 (“The Americas’’); ‘U.S. Board of Geographic Names Gazetteer,’’ no. 56 (‘‘Venezuela’”’) and no. 71 (“Brazil”); Eigenmann’s “The American Characidae,” 1917, pt. 1, pl. 1; and Eigenmann and Allen’s ‘‘Fishes of Western South America,” 1942, pp. 67-81. Since I was unable to find a few localities, specimens from such places are not included in the distributional map for the species (pl. 1). Some of the other records are so unspecific in locality, e.g., ‘Rio Orinoco,” that these cannot be plotted on the map. I have studied the routes followed by the collectors; the latitude and longitude used represents the best estimate for the information available. The specific localities from which specimens of Pterophyllum have been collected are as follows: locality longitude latitude Ambyiacu, Rio, Peru Sel is) Heriey Wy Araguaia, Rio, near Aruana, Brazil 14°54’ S 51°05’ W Atabapo, Rio, Colombia 4°03’ N 67°45’ W Belém [= Par4], Brazil 1°30’ S 48°55’ W Cacaul or Cacauoal [= Cacoal], Brazil 1°59’ S 46°13’ W Cashiboya, Lago above Contamana, Peru 7°19’ S 75°04’ W Casiquiare, Rio, Venezuela 2°01 N- 67°07" Wi Coary, Brazil 4°08’ S_ 63°07’ W Crab Falls, British Guiana 5°17’ N 58°59’ W Cupai [probably Rio Cupari], Brazil 3°04" S. ba 2a Wy Demerara or Georgetown, British Guiana 6°46’ N 58°10’ W Iquitos, Peru 3°oliS* Takano Jutahy, Brazil 3°50 /66rou aN Manaeapuri, Brazil 3°18’ 60°37 Ww Manacapuri, Lago, Brazil 3°10’ S 61°30’ W Manaus, Brazil 3°06’ S 60°00’ W Marajé I, in Amazon, Brazil 145i S 49° W Maximo, Lago, Brazil (?) (?) Monte Alegre, Brazil 2°01’ S 54°04’ W NO. 3555 locality Nazareth, Brazil Obidos, Brazil Pacaya, Rio [see Yarinacocha] Pévas [= Pébas] Cafio, Peru Porto Alegre, Brazil Porto do Méz, Brazil Porto Negro [probably near Manaus], Brazil Purus, Rio, Brazil Rockstone, British Guiana Rupununi River, British Guiana Santarém, Brazil Solimées, Rio, at Manacapurt, Brazil Tabatinga, near Leticia, Brazil Tefé [= Teffé], Rio, Brazil Tonantins, Brazil Ueranduba, Brazil Urubu Rio, Brazil Villa Bella [Rio Ramos], Brazil Xingu, Rio [Pérto Alegre], Brazil Yarinacocha [Rfo Pacaya], Peru ANGELFISHES—SCHULTZ longitude ems. aN UALS 3°10’ S 4°23’ S 1°45’ S 3°42’ 8 6°00’ N 3°55’ N 2°26’ S 3°18’ S 4°14’ 8 3°24’ § 2°46’ S (?) 2°55’ S 2°36’ S 4°23’ 8 5°10’ § Pterophyllum Heckel latitude 3 67°51’ W [probably] 55°30’ W 71°46’ W 52°45’ W 52°10’ W 61°28’ W 58°30’ W 59°06’ W 54°41’ W 60°37’ W 69°44’ W 64°45’ W 67°45’ W (2) 58°25’ W 56°39’ W 52°45’ W 74° W Pierophyllum Heckel, 1840, Ann. Wien. Mus., vol. 2, no. 12, p. 334 [type species: P. scalare Heckel=FPlatax scalaris Cuvier and Valenciennes].—Ahl, 1928, Zool. Anz., vol. 76, p. 252 [revision of 3 species]. Plataxoides Castelnau, 1855, Animaux nouveaux ou rares . Sud . . del’ Amerique du TasBLe 1—Number of vertebrae in Pterophyllum Species and localities dumerilii British Guiana Rio Negro-Amazon Rio Negro, types of leopoldi Belém and Marajo Island altwm Upper Orinoco scalare British Guiana Middle and lower Amazon Belém and Marajo Island Peruvian Amazon 14 15 16 17 18 Abdominal Caudal 12ers 14 1 1 Zi oi Le, fj Pal (el Gees 1 2 26 Ibe 3s a8} 13 6 7 7 ham ge} , p- 21, pl. 11, fig. 8 [type species: P. dumerilit Castelnau]. Total 27 28 29 30 KIO 31 The validity of the generic name Pterophyllum Heckel, long in use by ichthyologists and aquarists, needs to be clarified because Gosse (1963, p. 4) used the generic name Plataxoides Castelnau instead of Pterophyllum. Gosse (loc. cit.) and Whitley (1951, Proc. Roy. Zool. 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Soc. New South Wales, p. 68) were in error in considering that Pterophyllum Heckel is not available as a generic name with the type species Pterophyllum scalare Heckel (=Zeus scalaris Lichtenstein). Myers (1940, Stanford Ichthy. Bull., vol. 2, no. 1, p. 36) pointed out that Neave (1940, Nomenclator zoologicus, vol. 3, p. 1028) was in error in the following statement: ‘‘Pteropyhllum (pro-lla, Kirby 1825), Harris 1833, in Hitchcock, Rept. Geol. Min. Bot. Zool. Massach., 582-Orth.” I have checked Harris (1833, 1835, Rep. Geol. Mineral. Bot. Zool. Massachusetts) and can verify Myers’ observation that Harris consistently used Pterophylla, leaving Pterophyllum available. Pterophyllum dumerilii (Castelnau) PLATE 2 Plataxoides dumerilit Castelnau, 1855, Animaux nouveaux ou rares... de Amérique du Sud. . . , p. 21, pl. 11, fig. 3 [Pard=Belém]. Plataxoides leopoldi Gosse, 1963, Inst. Roy. Sci. Nat. Belgique Bull., vol. 39, no. 35, p. 4, pl. 1, fig. 2 [mouth of Rio Solimdées, about 90 km above Mana- capurt]. Specimens studied (see page 2 for more precise locality): BMNH 1902-11-4-10-12, Marajé, 3 (27 to 40 mm); BMNH 1902-2-11-4-9, Tonantins, 1 (41 mm); MCZ (out of 14992), Tefé, 2 (33 and 37mm); CNHM (out of 15254); Santarém, 1 (40 mm); CNHM 53846 Rupu- nuni River, 2 (33 and 54 mm); CNHM (out of 54370), Santarém, 1 (37 mm.); USNM 167772, Rupununi River, 1 (440 mm.); USNM 198177, Rio Urubu, 1 (45 mm.); USNM 198178, Rio Purtis, 2 (29 and 31 mm.); IRSNB 460, Rio Solimées. 90 km above Manacapurt, 27 paratypes of P. leopoldi, (34 to 61 mm); MNHN A 254, Para, Brazil, holotype of P. dumerilii (49 mm); MNHN 221-61-3-7, Cacaual, 1 (37 mm); MNHN 221-61-3-6, Caldron, 2 (42 and 51 mm). I examined the holotype of Platazoides dumerilii Castelnau at the Museum National Histoire Naturelle, Paris, and found that Castelnau’s illustration (1855, pl. 11) is in error. In his illustration, a black spot is shown behind the prominent black vertical bar that extends from the middle of the dorsal fin across body to the middle of the anal fin, whereas in the holotype this spot now shows as a trace in front of that dark bar, not behind it. Also, the dark vertical bars anteriorly on the illustration are incorrectly portrayed. On the holotype a short bar extends from the spiny dorsal origin ventrally to the lateral line, another short bar occurs halfway between the eye and the spiny dorsal origin; there is also a dorsoventral bar through the eye and across the cheek. These bars are prominent in well- preserved specimens and their positions differ notably from those of Pterophyllum scalare and P. altum. On the latter two species, the black bar through the eye extends dorso-posteriorly to the origin No. 3555 ANGELFISHES—SCHULTZ 5 of the spiny dorsal fin, replacing the two middle predorsal vertical bars on P. dumerilii. Thus, on the basis of coloration P. dumerilir can always be distinguished from P. scalare and P. altum. The number of vertebrae averages slightly fewer in P. dumeriliz than in the other two species (table 1) and there are also slightly fewer soft dorsal and soft anal rays and fewer oblique scale rows (from rear of head to midbase of caudal fin) (table 2) than in P. scalare and TABLE 2.—Counts recorded for Pterophyllum dumerilii Localities Dorsal fin rays Pectoral fin rays XE XII XIII 18 19 20 21 22 23 24 10 11 British Guiana 3 Wat) val 2 Rio Negro-Amazon 1 8 1 Sieve erly leben 27710 Rio Negro (types of leopoldi) | 4 20 2 Ce cin a5 2 2 Mouth of Amazon 3 1 4 i 383 Holotype of dwmerilii 1 1 1 Localities Anal fin rays Oblique scale rows VI 19 20 21 22 23 24 25 26 27 28] 26 27 28 29 30 31 32.533 British Guiana 3 2) LS al Rio Negro-Amazon 10 2 Deno Tee 83 Th. PP Se mall 1 Rio Negro (types of leopoldi) Zag) 1 3 Ty a ile aa) eantsh nt Srenttl Mouth of Amazon 4 1 2 1 iy <3} Holotype of dwmerilii 1 1 1 P. altum (table 3); however, these differences are not great enough to enable me to identify all specimens on the basis of fin ray and scale counts, even if the dorsal and anal rays are added to the oblique scale rows to form a character index (table 4). An examination of table 2, shows that the specimens of P. dumerilia collected near Belém, at the mouth of the Amazon have a slightly greater number of fin rays and scales than do specimens further up the Amazon—Rio Negro systems. The counts made on the holotype of P. dumerilw from Parad (now Belém) with XII,23 dorsal, VI,24 anal, 11 pectoral fin rays, and 30 scale rows along the side, fit into the higher end of the frequency distributions (table 2) for P. dumerilii; however, since the frequency distributions partly overlap for the various localities and the number of specimens are not sufficient to establish the extent of variability among populations, if such exist, I am recognizing only a single species, P. dumerilii. P. dumerilit has been collected in the Amazon basin and in the Rupununi River of British Guiana (see open circles in map, pl. 1). 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Pterophyllum altum Pellegrin PLATE 3 Pierophyllum altum Pellegrin, 1903a, Bull. Mus. Hist. Nat. Paris, vol. 9, p. 125 [Rio Atabapo, Orinoco]; 1903b, Mem. Soe. Zool. France, vol. 16, p. 252, pl. 4, fig. 4 [Atabapo].—Regan, 1905, Ann. Mag. Nat. Hist., ser. 7, vol. 16, p. 442 [Rio Orinoco].—Higenmann, 1910, Rep. Princeton Univ. Exped. Patagonia 1896-1899, vol. 3 (Zool.), pt. 4, p. 479 [Orinoco].—Ahl, 1928, Zool. Anz., vol. 76, p. 255 [Orinoco].—Schultz, 1949, Proc. U.S. Nat. Mus., vol. 99, p. 167 [Rio Atabapo]; 1953, Tropical Fish Hobbyist, vol. 1, no. 5, pp. 5-7, 20 [key to aquarium-bred angelfish]. Specimens studied (see page 2 for more precise locality): BMNH 1904—6—28-2-3, Rio Orinoco, 2 (59 and 60 mm); MNHN 221-61-1-3, Chaffanjon (Rio Atabapo), 5 syntypes of P. altum (42 to 62 mm); MNHN 221-61-1-1, Chaffanjon, 4 syntypes of P. altum (63 to 75 mm); MNHN 221-61-1-2, Chaffanjon, 5 syntypes of P. altum (59 to 65 mm); USNM 163204, Venezuela, 1 (79 mm); USNM 196007, Colombia, 1 (67 mm); CAS [no number], Rio Casiquiare, 1 (53 mm). Pterophyllum altum Pellegrin has a color pattern identical with that of P. scalare; however, no black spot occurs on the upper midside as in P. dumerilu. The black bar through the eye curves dorso- posteriorly to the origin of the spimy dorsal and is not broken into three separate bars as in P. dumerilia. This species is best recognized from P. scalare by the greater average number of median fin rays, oblique scale rows, and vertebrae (tables 1-4). Undoubtedly P. altum represents the P. scalare type of angelfish in the upper Orinoco, and in having a higher average number of dorsal, anal, oblique scale rows and vertebrae than P. scalare, it might be considered to represent only a subspecies of P. scalare; however, since P. altum has been taken so far only in the upper Orinoco basin, I prefer tentatively to recognize it as a distinct species. Pterophyllum scalare (Lichtenstein) PLATE 4 Zeus scalaris Lichtenstein, 1823, Verz. Doubl. Zool. Univ. Berlin, p. 114 [Brazil]. Platax scalaris Cuvier and Valenciennes, 1831, Histoire naturelle des poissons, vol. 7, p. 237 [Brazil, in “Collection de Bloch, au Musée de Berlin’’]. Pterophyllum scalaris Heckel, 1840, Ann. Wien. Mus., vol. 2, p. 335, pl. 30, figs. 5, 6, 7-7a, 8-8b [Reo Negro]. Pierophyllum scalare Giinther, 1862, Catalogue of the fishes in the British Museum, vol. 4, p. 316 [Rio Cupai; Brazil]—Kner, 1862, Sitz. Akad. Wiss. Wien., vol. 46, p. 295, pl. 1, figs. 1-la—Cope, 1872, Proc. Acad. Nat. Sci. Phila- delphia, p. 250 [Rio Ambyiacu].—Steindachner, 1875, Sitz. Akad. Wiss. Wien., vol. 71, p. 76 [Amazon at Santarém, Monte Alegre, Villa Bella, Obidos, Coary, Ueranduba, Tonantins, Tabatinga, Rio Jutahy, Xingu, Lago Manacapurti, Lago Maximo, Pard, Rio Ambyiacu; ?Barra do Rio Negro].— Higenmann and Eigenmann, 1891, Proc. U.S. Nat. Mus., vol. 14, p. 71 SCHULTZ—PLATE 1 120 NAT. MUS. VOL. PROC. U.S. ae SAL NAPE RST St eS ung kydotajg jo saisads 10} dew [euonnqiisiq wu 4 ’ \\ assay co hia LAE ’ 4 “., wor .o 2. } if =p Jr : C j r me ile fr eect Se we N is Dh Tsctoed| hese a ” < aejoas dd ® unio dd -£ Teun sO PROC. U.S. NAT. MUS. VOL. 120 SCHULTZ—PEATE 2 Pterophyllum dumerilii (Castelnau). SCHUETZ—PEATE 3 VOL. 120 MUS. PROC. U.S. NAT. in. Pterophyllum altum Pellegr SGCHUETZ—PEATE 4 PROC. U.S. NAT. MUS. VOL. 120 NO. 3555 ANGELFISHES—SCHULTZ 7 [Amazon].—Higenmann and Bray, 1894, Ann. New York Acad. Sci., vol. 7 p. 624.—Pellegrin, 1903, Mem. Soc. Zool. France, vol. 16, p. 251 [high Peru; upper Amazon, Tefé; Pard].—Regan, 1905a, Ann. Mag. Nat. Hist., ser. 7, vol. 16, p. 441 [Rio Cupai; Tabatinga; Manaus; Tonantins; Marajo Island in Rio Amazon]; 1905b, Proc. Zool. Soc. London, pt. 1, p. 190 [Rio Negro, Brazil]—lIhering, 1907, Rev. Mus. Paulista, vol. 7, p. 335 [Amazon at Tabatinga and tributaries]—Higenmann, 1910, Rep. Princeton Univ. Exped. Patagonia, 1896-1899, vol. 3 (Zool.), pt. 4, p. 479 [Amazon].— Haseman, 1911, Ann. Carnegie Mus., vol. 7, p. 372 [Santarém, Manaus].— Higenmann, 1912, Mem. Carnegie Mus., vol. 5, p. 521 [Rupununi River; Rockstone; Crab Falls]—Ribeiro, 1915, Arch. Mus. Nac. Rio de Janeiro, vol. 17, p. 56 [Amazon and tributaries]——Ahl, 1928, Zool. Anz., vol. 76, p. 254 [Amazon].—Higenmann and Allen, 1942, Fishes of western South America, p. 406 [Iquitos, Yarinacocha].—Fowler, 1945, Los peces del Peru, Mus. Hist. Nat. Javier Prado, Univ. Nac. Mayor de San Marcos, p. 253, fig. 88 [Amazon, Peru].—Schultz, 1949, Proc. U.S. Nat. Mus., vol. 99, p. 167 [Venezuela]; 1953, Tropical Fish Hobbyist, vol. 1, pp. 5-7, 20 [key to aquarium-bred angelfish]. Pterophyllum eimekei Ahl, 1928, Zool. Anz., vol. 76, p. 252, fig. 1 [Rio Negro in the Amazon]. Specimens studied (see page 2 for more precise locality).—The following have USNM catalog numbers: 163210, probably British Guiana, 20 (28 to 63 mm); 198180, Belém, 5 (72 to 78 mm); 198179, Rio Purts, 4 (46 to 67 mm); 179565 and 179566, Rio Urubu, 54 (33 to 80 mm) ; 163210, probably lower Amazon, 6 (72 to 81 mm); 191591, upper Rio Araguaia, 1 (17.5 mm); 167754, Yarinacocha, 1 (54 mm) ; 167755, Iquitos, 2 (41 and 63 mm); 26664, Tabatinga, 1 (42 mm); 174942, British Guiana, 3 (71 to 81 mm); 175933, Pévas, 3 (27 to 31 mm). The following have BMNH numbers: 1926-3-3-3, Rio Negro, 1 (60 mm); 1925-10-28-452-457, Manacapurt, 6 (44 to 99 mm); 1926-10-27-488, Monte Alegre, 1 (50 mm); 53-3-19-42, Rio Cupai [probably Rio Cupari], 1 [not measured]; 81—5—-13-129, Lago Cashi- boya, 1 (43 mm); 85-1-14-71, Tabatinga, 1 (55mm). The following have MNHN numbers: 221-61-2-1, Rio Negro, 1 syntype of P. emmekei (55 mm): 221-61-3-10, French Guiana, 3 (63 to 71 mm); 221-61-3-2, Tefé, 2 (90 and 101 mm); 221-61—3-3, Peru (Amazon), 5 (41 to 51 mm); 221-61-3-4, Peru (Amazon), 3 (50 to 82 mm); 221-61-3-5, Tefé, 5 (30 to 44 mm); 221-61-3-6, Caldron, 2 (65 and 79 mm); 221-61-3-9, Brazil, 3 (49 to 52 mm). The following have CAS numbers: [no number], Rio Negro, 4 (72 to 84 mm); [no num- ber], Santarém, 11 (34 to 50 mm); IU 17780, Yarinacocha, 1 (47 mm) ; IU 2096, Tabatinga, 1 (43 mm); IU 5106, Lower Amazon, 1 (51 mm); IU 12502, Rupununi River, 1 (34 mm); IU 12504, Essequibo River, 2 (48 and 50 mm); IU 13368, Manaus, 1 (84 mm); [no number] “Porto Negro,” 4 (46 to 61 mm); 5100, Maraj6, 1 (57 mm); TU 12508, Essequibo River, 2 (43 and 44 mm); IU 15988, Iquitos, 4 (42 to 69 mm). The following have MCZ numbers: 14989, Coary, 4 (84 to 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 94 mm); 14993, Manacapurd, 2 (59 and 60 mm); 14984, Pérto do Moz, 2 (47 and 51 mm); 14988, Tonantins, 3 (59 to 94 mm); 14997, Tabatinga, 2 (92 and 99 mm); 14985, Porto do Moz, 20 (37 to 48 mm); 14994, Santarém, 10 (46 to 54 mm); 14998, Obidos, 4 (47 to 71 mm); 14991, Jutahy, 5 (43 to 73 mm); 14992, Jutahy, 3 (39 to 41 mm); 14987, Pérto do Moz, 5 (40 to 44 mm); 14977 to 14980, Tefé, 10 (51 to 101 mm) ; 14977 to 14980, 14982 and 14983, Tefé, 6 (74 to 100 mm); 14996, Santarém, 3 (385 to 46 mm); 14990, Villa Bella, 6 (36 to 41 mm); 14999, Parad, 1 (54 mm); 14995, Santarém, 1 (44 mm); 14976, Monte Alegre, 6 (33 to 89 mm); 14981, Tefé, 10 (63 to98 mm). The following have CNHM numbers: 15254, Santarém, 1 (88 mm); 54371, Manaus, 4 (81 to 90 mm); 54233, Manaus, 1 (49 mm); 53847, Esse- quibo River, 2 (41 and 43 mm); 54370, Santarém, 3 (40 to 56 mm); 53848, Essequibo River, 4 (44 to 47 mm); 15562, Peru, 1 (53 mm). The following have SU numbers: 36799, Pévas, 1 (62 mm); 36660, near Pévas, 1 (28 mm); 2205, Lower Amazon, 1 (59 mm); 36661, near Pévas, 2 (23 and 26 mm); 36797, near Pévas, 1 (37 mm); 54245, Santarém, 3 (45 to 57 mm); 60504, Pévas, 1 (75 mm); 36798, near Pévas, 3 (22 to 36 mm); 36659, near Pévas, 2 (37 and 46 mm); 36796, near Pevas, 3 (47 to 74 mm). P. scalare (Lichtenstein) is the common angelfish of aquarists. Since 1928, when Ahl described P. eimeker from the mouth of the Rio Negro in the Amazon basin considerable confusion has occurred as to the validity of that species. P. ewmekei was thought to average fewer dorsal and anal rays and fewer oblique rows of scales (Schultz, 1953) and as indicated by Ahl (1928) and Ladiges (1949, Deutsche Aquar. Terr., year 2, no. 3, pp. 50-52); however, most of their counts were based on aquarium specimens, undoubtedly inbred strains in aquaria, which most probably did not represent the species com- plex of natural populations found in the Amazon basin. The counts recorded in tables 3 and 4 are based on specimens from definite natural localities and not on aquarium-reared material. The counts for the syntype of P. eimeker with XIII, 24 dorsal, VI, 26 anal, 10 pectoral fin rays, and 36 scale rows on the side fall nearly in the middle of the frequency distributions for Amazon speci- mens of P. scalare (tables 3 and 4). Thus, I conclude that P. eimeker is a synonym of P. scalare. The differences between the color patterns of P. scalare and P. dumerilii, discussed under the latter species and shown in plates 1 and 3, should make identification easy; however, P. scalare and P. alitum cannot be distinguished on the basis of coloration. The chief differences are the greater number of soft dorsal and soft anal fin rays and the number of scales in P. altum (tables 3 and 4). P. altum has 27 to 31 soft dorsal rays whereas P. scalare rarely has more than 26. ANGELFISHES—SCHULTZ 3555 NO. lb OF Sb Fh Sb Ch Ib OF 6E BSE LE OE GE FE SE CE TE OF 66 8 SMO OBIS ONDIIGO mn Au AN 1 a £& re ov. 9T i! cE (i KE € GL ao AY Oo Oriwmo onl co re oD Seon 19 ic.) st re nN aA NON & TE Of 66 8% 16 9% GS HF 8 CG ITA IA A shui Uy [eUy a Nn nN x Onn s oOo tron oO oD OD ws MRMINOO mn AQ oO rl OD ra rs 00 mozeury uUeIANIOg esuneqey ‘soouljos “y ‘Ayeine “yY -surZUeUOT, 9}9L AIB09 ‘nindvovury, ‘sning “y ‘nqnig “YW -sneue OIZ0TV OJUO {0199 N OO wedny “Y ‘eT VIEA ‘SOPIAO Wyre} URS ZO OP OWT ‘T olereyy “TA9TP ‘erensery “Wy reddy euemny Ys 9LD]DI8 oo00ulIO oY Joddq wnqyD cL IL OF sAvl ye10490 SoT}ITBOOT pu setoedg re MN N WAN Nr &T g G IT ¥G TE O€ 62 8% LG 9S GS FE EZ rN 0€ 6 9 9& 6G, 1G) ATX TEX IX De shel uy [es1oq uozeury UeIANIOg esureqey ‘sooufog “y ‘AyeIne “y :sujueUOL, 93°.L Arv09 ‘naindeovuryy ‘snmg “WY ‘nqnig “y ‘sneuey OIZ0[V eJUOJ {0139 N Oyo wedng “Y {ed VITA :Soplag “UI9IeyUBS ZOW OP 01100 "J ofereyy Ug Te erensery “yy seddq eueMpy Ye aLD]D98 oo00uLIO OT reddy wnyD SoTiITBO0T pu sooedg wunj[e ‘dq puv oivypeos wan{pAydo19jg of papsooas squnog—g§ AIsVL 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Although proportional measurements were made on all three species of Pterophyllum, the great variability even at nearly equal sizes suggests that little reliance can be placed on measurements for iden- tification purposes. Tapie 4.—Character index: Total number of rays in dorsal and anal fins added to number of scale rows for Pterophyllum Species and localities 83 85 87 89 91 93 95 97 99 101 103 105 107 109 111 113 115 117 119 121 123 125 84 86 88 90 92 94 96 98 100 102 104 106 108 110 112 114 116 118 120 122 124 126 altwm Upper Rio Orinoco oy iy, Eh scalare British Guiana Leh Bot ee 5 Upper R. Araguaia Belém; Marajo I. Pérto do Moz 5 Santarém; Obidos, Villa Bella; R. Cupari; Porto Negro; Monte Alegre Sea Sees lies eal Manaus; R. Urubu; R. Purus; Manacapurt; Coary By hy Als th a) a ube al Tefé 1 Pa 6 Tonantins; R. Jutahy; R. Solimées; Taba- tinga il 983 Peruvian Amazon Bey ey aly 83 dumerilii British Guiana 1 Rio Negro-Amazon 2 3 Py al 1 Rio Negro (type of leopoldi) 1 sky Le Srl at Mouth of Amazon, Cacaual 1 1 to we OO _ bo i = = be | cn — bo for} e =" on e bo bo An inspection of tables 3 and 4 indicates that for P. scalare there may be certain areas where distinct populations exist; the specimens from Pérto do Moz, Santarém, Obidos, for example, average fewer soft dorsal fin rays than specimens from Manaus, Rio Urubu, and Tefé. Specimens from British Guiana have about the same counts as those listed from Pérto do Moz. Before any conclusion can be made for the “population” at the mouth of the Amazon, large series will need to be studied to find out the meaning of the bimodal nature of the frequency distribution for specimens from Belém and Marajé Island. P. scalare ranges in the Amazon Basin and in the Rupununi and Essequibo Rivers of British Guiana with a single record from French Guiana (MNHN 221-61-3-10), the exact locality unknown. The known localities represented by specimens studied by me are recorded in plate 1. U.S. GOVERNMENT PRINTING OFFICE: 1967 Proceedings of the United States National Museum SMITHSONIAN INSTITUTION +¢« WASHINGTON, D.C. Volume 120 1967 Number 3556 eee SOME PORTUNID CRABS FROM THE PACIFIC AND INDIAN OCEANS IN THE COLLECTIONS OF THE SMITHSONIAN INSTITUTION By WILLIAM STEPHENSON and May Regs! Introduction The material on which the following report is based was received for identification as part of a program of systematic research on the marine fauna of the tropical Pacific area, with studies of related faunas as required, sponsored jointly by the Biology Branch of the Office of Naval Research [Contract NONR 1354(09)] and the Division of Biology and Medicine of the Atomic Energy Commission [Contract AT (30-1)2409]. No attempt has been made in this study to incor- porate material previously identified and cataloged in the collections of the Smithsonian Institution. The collections as received included 96 species, but only 82 are reported below. The remaining are western American species, which have been included in a monograph by Dr. John S. Garth and the senior author (Garth and Stephenson, 1966), and species of Ovalipes, which are to be included in a revision of the genus. Initially, 12 of the 82 species were new, but Crosnier (1962) de- scribed 5 of them. No doubt a considerable number of Indo-West- Pacific species still await description, and the time for a comprehensive monograph is still distant. 1 Both authors: Department of Zoology, University of Queensland, St. Lucia, Brisbane, Australia. Manuscript submitted for publication in February 1965. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 As the group becomes better known, identifications become easier only in certain directions. In the present work some clarification is effected by recognizing the validity of species previously synonymized. As more species are discovered, however, the distinctions between them become narrower, and the difficulties of relating them to the older descriptions increase. Until type material is reexamined and redescribed, several groups must remain confused. In the present work these are termed “‘complexes,” and the Portunus gladiator and P. longispinosus complexes have caused particular difficulty. A more fundamental problem concerns the importance attached to differences in male secondary sexual characteristics. Shapes of male abdomens have long been recognized as having diagnostic importance, and since Stephenson (1945), following Brocchi (1875), increasing stress has been laid upon male first pleopods. In both these structures the extent of ‘‘expectable” variation has been known in general terms, and few cases of greater variation have been described (e.g., male pleopods of P. orbitosinus by Gordon, 19388, and male abdomens of P. granulatus by Crosnier, 1962). In the pres- ent work unexpected variability in male pleopod structure has been noted in several cases. In one species (P. orbitosinus) there is approxi- mately continuous gradation between wide extremes, and in three others (Thalamita stimpsoni, T. danae, and Portunus argentatus) there is distinct male dimorphism. If a species is defined in terms of a dis- tinct morphological gap between two forms, 7’. stimpsoni, T. danae, and P. argentatus should each be split into two species. However, apart from male pleopods and abdomens, no structural differences could be found. For the purposes of the present paper, P. argentatus was taken as a “‘test case” of the concept of a portunid species (see p. 19). This concept implies a distinct morphological gap in general facies, as well as in the shape of the male abdomens and structure of the male pleopods. If no distinctions were noted in general facies, groups with different pleopods are here recorded as different “forms.” Pos- sibly they are incipient species. Details of synonymy vary from species to species; where there had been past confusion, reasonable detail is given, but in other cases only critical references are cited. Brief morphological comments are given upon species adequately treated in recent literature, and fuller rede- scriptions are restricted to inadequately described species. Subfamily headings follow Stephenson and Campbell (1960) ; within subfamilies genera are in alphabetical order, and within genera species are also in alphabetical order, except where similar species are grouped into ‘“complexes.”’ NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 3 Geographical distributions adhere closely to the expected pattern, with the better known species extending from East Africa and the Red Sea to India, Japan, Australia, and the mid-Pacific islands. Only one of the present species (Thalamita picta) and one species of Ovalipes (see later work) are known to cross the mid-Pacific boundary. Listed specimens within the 16 main locality groupings are recorded in chronological order. The locality groupings are: East Africa, in- cluding Saudi Arabia and the Red Sea; Malay Peninsula, including Rangoon and Sumatra; China, including Hong Kong and Formosa; Philippines, including the Sulu Sea and North Borneo; Celebes; Australia; Melanesia, including New Guinea, Fiji, and the Solomon Islands; Palau Islands, including the Carolines; the Marianas, includ- ing Yap and Guam; Japan; the Marshalls; the Gilbert Islands; Samoa; Fanning Island; Hawaii; and the Tuamotu and Society Islands. Throughout, two abbreviations are used: ‘‘Alb.”’ for collections of the U.S. Fish Commission Steamer Albatross either from the Philip- pines, Japan, or Samoa, and ‘‘GVF”’ for ‘George Vanderbilt Founda- tion.”” And, throughout, dimensions are those of total breadths, including the last anterolateral teeth. In cases of damage, the approximate estimated breadth of the undamaged specimen is given. Specimens considered too small or too damaged for identification have been excluded from this account. It is obvious that this paper could never have been written but for the kindness of Dr. Fenner A. Chace, Jr., who asked us, in the name of the Smithsonian Institution, to report upon the collections. We are also deeply grateful to the following museums and their directors for allowing us to borrow specimens for comparative study: Australian Museum, Sydney (Dr. J. W. Evans), Western Australian Museum (Dr. W. D. L. Ride), Queensland Museum (the late Mr. G. Mack), Central Marine Fisheries Research Institute, India (Dr. S. Jones), Muséum National d’Histoire Naturelle, Paris, Laboratoire de Zoologie (Arthropodes) (Dr. J. Forest). The following carcinologists have helped materially either in loans or in examination of specimens or by their advice: Dr. John C. Yaldwyn, Dr. Ray W. George, Mme. Daniéle Guinot, Dr. Lipke B. Holthius, Dr. John S. Garth, and Dr. Raymond B. Manning. Thanks are due to Mr. E. Hollywood, University of Queensland photographer, for photographic illustrations, the Research Grants Committee of the University of Queensland, and the Smithsonian Institution for financial assistance. VOL. 120 4 PROCEEDINGS OF THE NATIONAL MUSEUM Checklist Subfamily CaroprrininaE Borradaile Genus Libysies A. Milne Edwards truncatifrons (de Man) Genus Carupa Dana tenutpes Dana Subfamily MacroprpinaE Stephenson and Campbell Genus Parathranites Miers orientalis Miers Subfamily Capuyrinar Alcock Genus Lissocarcinus Adams and White orbicularis Dana Genus Caphyra Guérin rotundifrons (A. Milne Edwards) Subfamily PorruninAE Stephenson and Campbell Genus Charybdis de Haan Genus Portunus Weber—Continued (Charybdis) anisodon de Haan (Charybdis) callianassa (Herbst) (Charybdis) curtilobus, new species (Charybdis) feriatus (L.) (Charybdis) ef. feriatus (L.) (Charybdis) hellerit (A. Milne Edwards) (Charybdis) japonica (A. Milne Edwards) (Charybdis) miles (de Haan) (Charybdis) natator (Herbst) (Charybdis) orientalis Dana (Goniohellenus) truncata (Fabricius) (Goniohellenus) vadorum Alcock (Gonioneptunus) bimaculata (Miers) (Goniosupradens) erythrodactyla (Lamarck) (Goniosupradens) obtusifrons Leene Genus Portunus Weber acerbiterminalis, new species argentatus (A. Milne Edwards) brockizi (de Man) dubius (Laurie) euglyphus (Laurie) P. gladiator complex gladiator (Alcock) pseudoargentatus Stephenson granulatus (H. Milne Edwards) hastatoides Fabricius P. longispinosus complex emarginatus Stephenson and Campbell tranjae Crosnier P. longispinosus complex—Con. macrophthalmus Rathbun tenuicaudatus Stephenson species (unidentifiable) orbitosinus Rathbun pelagicus (L.) pubescens (Dana) pulchricristatus (Gordon) rubromarginatus (Lanchester) ef. rubromarginatus (Lanchester) rugosus (A. Milne Edwards) sanguinolentus (Herbst) spiniferus, new species spinipes (Miers) tenuipes (de Haan) trituberculatus (Miers) tuberculosus (A. Milne Edwards) tweediet (Shen) Genus Scylla de Haan serrata (ForskAl) Genus Thalamita Latreille admete (Herbst) auauensis Rathbun bouviert Nobili chaptalit (Audouin) coeruleipes Jacquinot corrugata Stephenson and Rees crenata (Latreille) dakini Montgomery danae Stimpson demant Nobili forestt Crosnier gatavakensis Nobili NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 5 Genus Thalamita Latreille—Continued Genus Thalamita Latreille—Continued gloriensis Crosnier pseudopoissoni, new species granosimana Borradaile quadrilobata Miers imparimana Alcock sexlobata Miers integra Dana sima H. Milne Edwards mitsiensis Crosnier spinifera Borradaile multispinosa, new species spinimana Dana oculea Alcock spinimera, new species parvidens (Rathbun) stephensoni Crosnier philippinensts, new species stimpsont A. Milne Edwards picta Stimpson Genus Thalamitoides A. Milne Edwards pilumnoides Borradaile quadridens A. Milne Edwards poissoni (Audouin and Savigny) tridens A. Milne Edwards prymna (Herbst) Subfamily PopoputHaLMINnAk Borradaile Genus Podophthalmus Lamarck vigil (Weber) Subfamily CaTopTRinaE Borradaile, 1907 Genus Carupa Dana, 1851 Carupa tenuipes Dana Carupa tenuipes Dana, 1851, p. 129; 1852a, p. 85; 1852b, pp. 279-280.—Leene, 1940, pp. 165-168, figs. 1, 2.—Stephenson and Campbell, 1960, p. 88, pl. 2 (fig. 1).—Crosnier, 1962, pp. 19-20, figs. 16-23, pl. 1 (fig. 1).—Miyake, Sakai, and Nishikawa, 1962, p. 128 (record only). Carupa laeviscula Heller, 1862, p. 520; 1865, p. 27, pl. 2 (fig. 2).—Alcock, 1899, p. 26.—Leene, 1938, pp. 9-10.—Sakai, 1939, p. 373, pl. 44 (fig. 3).—Edmond- son, 1954, pp. 226-227, figs. 3b, 4e, 4f, 4g. Marertau.—Melanesia: Mariemberg, New Guinea, May 10, 1929, Albert W. Herre, 1 female. Marianas: Saipan, coral heads, 1945, A. H. Banner, 1 male. Marshalls: Lagoon reef, Latoback Is., Rongarik, Aug. 18, 1947, F. M. Bayer, F. C. Zimmerman, 1 male; Aug. 21, 1947, 1 male. Hawai: Waikiki Marine Lab., Honolulu, under stones near shore, Feb. 9, 1942, G. S. Mansfield, 1 female. Tuamotus and Societies: Sta. 29a-57, outer reef, Tickahau Atoll, Maiai Is.; Apr. 14, 1957, Bredin Exped., 1 female; Huahine Is., head of Baie de Maroe, from branching coral, Apr. 30, 1957, Bredin Exped., 1 male; Huahine, off Point Teffaao, Sta. 90a-57, from dead coral, May 2, 1957, Bredin Exped., 1 female; Moorea, Society Is., outside barrier reef, 200 yds. E. of pass to Papetoai Bay, 70 ft., May 15, 1957, J. Randall, 1 female. M®ASUREMENTS.—Males, 8-11 mm.; females, 9-22 mm. REMARKS.—Specimens resemble Crosnier’s (1962) figure 16, rather than his figure 17. 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Disrripution.—Madagascar to Hawaii, including Australia and Japan. Genus Libystes A. Milne Edwards, 1867 See Stephenson and Campbell, 1960, p. 85, for synonymy. Libystes truncatifrons (de Man) Catoptrus nitidus A. Milne Edwards, 1870, p. 82.—Ortmann, 1894, p. 687.— Alcock, 1900, p. 307.—Laurie, 1906, p. 422 (in part). Rathbun, 1911, pp. 238-239.—Tesch, 1918, pp. 179-180, pl. 9 (figs. 4, 4a-d).—Sakai, 1936, p. 135, pl. 35 (fig. 2).—Balss, 1938, p. 29.—-Sakai, 1939, p. 372, pl. 44 (fig. 2). Goniocaphyra truncatifrons de Man, 1887b, p. 339, pl. 14 (fig. 1); 1890, p. 67.— Zehntner, 1894, p. 163, pl. 8 (fig. 12)—Borradaile, 1900, p. 577. Libystes truncatifrons (de Man).—Stephenson and Campbell, 1960, pp. 85-86 (in key).—Crosnier, 1962, pp. 16-17, figs. 11-15. ?Catoptrus nitidus A. Milne Edwards.—Edmondson, 1946, p. 277; 1954, p. 224, figs. 2e-g. MareErtaAL.—Tuamotus and Societies: Both coll. Bredin Exped., Huahine Is., Sta. 84a-57, head of Baie de Maroe, from branching coral, Apr. 5, 1957, 1 male (6 mm.). Tickahau Lagoon, Sta. 10a-57, Apr. 11, 1957, 1 ovig. female (9 mm.). Remarks.—lIf Tesch (1918, pp. 177-178) is correct in stating that there are no reasons for maintaining the genus Catoptrus A. Milne Edwards, 1870, then the above synonymy follows. Serene (pers. comm.) has drawn our attention to Barnard’s sugges- tion (1954, p. 100) that C. inequalis Rathbun (1906), which is very close to the present species, may be a synonym of L. edwardsi Alcock (1900). The male abdomen resembles that figured by Crosnier (1962, fig. 14); unfortunately no male pleopods are present so that Edmondson’s (1954) figures 2f and g cannot be checked. DistTRIBUTION.—Madagascar to Hawaii, including Mauritius, Amirante, Coetivy, Ceylon, Amboina, Batavia, Japan, Samoa, and Fiji. Subfamily MAcropipinaE Stephenson and Campbell, 1960 Genus Parathranites Miers, 1886 Parathranites orientalis (Miers) Lupocyclus (Parathranites) orientalis Miers, 1886, pp. 186-187, pl. 17 (figs. 1, la, b, c). Parathranites orientalis (Miers).—Alcock, 1899, pp. 17-18.—Sakai, 1939, pp. 376-877, pl. 43 (fig. 2).—Barnard, 1950, pp. 148, 149, figs. 29 i-1.—Stephenson, 196la, pp. 97-98, figs. 1B, 2H, pls. 1 (fig. 2), 4B.—Crosnier, 1962, p. 22, fig. 24. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES Yi; MarertAL.—Philippines: Sta. 5154, Bakun Point, 5°14’50’N., 119°58’45”E., 12 fm., coarse sand, Feb. 19, 1908, Alb., 1 fragmented female (ca. 18 mm.). Distripution.—Madagascar, Seychelles, Ki Is., Admiralty Is., India, Andamans, Solomon Bank, Japan, and eastern Australia. Subfamily CapuyrinaE Alcock, 1899 Genus Caphyra Guérin, 1832 Caphyra rotundifrons (A. Milne Edwards) Camptonyx rotundifrons A. Milne Edwards, 1869, p. 156, pl. 7 (figs. 11, 12). Caphyra rotundifrons (A. Milne Edwards).—Stephenson and Campbell, 1960, pp. 101-102, figs. 1H, 2J, 3A—C, 3K, pls. 3 (fig. 4), 5J.—Crosnier, 1962, p. 30, fig. 39, pl. 1 (fig. 2). Marrriau.—Tuamotus and Societies: Matu Uta Is., Papeete Harbor, Tahiti, Sta. 42-57, outer reef, Apr. 20, 1957, Bredin Exped., 1 male (7 mm.) ; Sta. 62-57, Bora Bora, inner edge of outer reef, Apr. 25, 1957, Bredin Exped., 1 female (7 mm.). Distripution.—Mauritius to Fiji and Marianas, including Australia. Genus Lissocarcinus Adams and White, 1849 Lissocarcinus orbicularis Dana Lissocarcinus orbicularis Dana, 1852a, p. 288, pl. 18 (figs. a-e); 1852b, p. 86.— Alcock, 1899, pp. 20-21—Leene, 1938, p. 7.—Sakai, 1939, pp. 379-380, pl. 45 (fig. 1).—Barnard, 1950, pp. 145-146, fig. 28g.—Edmondson, 1954, p. 230, fig. 6b.—Stephenson and Campbell, 1960, pp. 95-96, pl. 3 (fig. 2.)— Forest and Guinot, 1961, p. 27, figs. 15a, b, 16.—Crosnier, 1962, pp. 25-27, figs. 26, 27, 31. Marertau.—Hawaii: All from or near the Waikiki Marine Lab., Honolulu, in 1942; March 22, G. S. Mansfield, 1 male, 1 female; April 30, G. S. Mansfield, 1 female; May 22, G. S. Mansfield, 1 male (damaged); May 23, Mansfield and Bonne, 2 females (1 ovig.); May 30, 1 male. Tuamotus and Societies: All coll. Bredin Exped. 1957; Sta. 13-57 Tikahau Atoll, from Holothuria atra, ocean reef, April 12, 1 female (damaged); Sta. 77-57 Uturoa, Raiatea, 1-3 ft., April 28, 1 female; Sta. 79-57, Taoru Is., Raiatea, reef near shore, shallow water, April 29, 1 male, 1 female; Sta. 84-57, head of Baie de Maroe, Huahine, from holothurian, sandy shoal, 2-3 ft., April 30, 1 male; Sta. 86-57, Baie de Bourayne, Huahine, sandy reef, May 1, 1 male. MerasuREMENTS.—Males, 7-10 mm.; females, 8-12 mm.; ovig. female, 9 mm. 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Remarxs.—After prolonged preservation, no traces of pigmentation remained. Distrrpution.—Africa to Hawaii and Tuamotus, including Australia and Japan. Subfamily PoRTUNINAE Stephenson and Campbell, 1960 Genus Charybdis de Haan, 1833 Subgenus Charybdis de Haan, 1833 Charybdis (Charybdis) anisodon (de Haan) Portunus anisodon de Haan, 1835, p. 42. Charybdis (Charybdis) anisodon (de Haan).—Leene, 1938, pp. 64-67, figs. 29, 30.— Stephenson, Hudson, and Campbell, 1957, p. 493, pl. 1 (fig. 1).—Crosnier, 1962, pp. 81-82, figs. 141-142, pl 6 (fig. 1). Charybdis anisodon (de Haan).—Sakai, 1939, pp. 405-406. MareriaL.—Malay Peninsula: Cham Han Bight, Siam, Jan. 9, 1924, H. M. Smith, 1 male, Chantabun River at Lem Sing, Siam, May 7, 1927, H. M. Smith, 2 females; Thailand, Sta. 26, entrance to Trat Bay, Gulf of Thailand, flat mud bottom, 11°58’30’N., 102°44’05”E., 5 m., Oct. 29, 1957, GVF, 1 male; Sta. 27, east coast, Gulf of Thailand, S. of Trat Bay off Lam Son Village near Cambodian border, 11°57’00"N., 102°44’45”E., 10 m., Oct. 29, 1957, GVF, 2 males; Thailand, anchorage in Mae Nam Chantaburi River at Tha Chalaep Harbor, Chantaburi Province, Sta. 134, Dec. 24, 1957, GVF, 1 female. Philippines: Manila Bay, Dec. 7, 1907, Alb., 1 male; Manila Bay, outside of breakwater, mud, small rocks, Dec. 12, 1907, Alb., 1 male; Tacloban Anchorage, about ship, hand dredge, 3 fm., Apr. 12, 1908, Alb., 2 males (1 fragmented), 1 female. M f&ASUREMENT.—Males, 12-45 mm.; females, 13-28 mm. ReMARKsS.—In larger males the borders of ultimate and penultimate abdominal segments form an uninterrupted curve instead of the in- dented separation shown by Crosnier (1962, fig. 142). DistTRIBUTION.—Madagascar and Red Sea to New Caledonia, Japan, and Australia. Charybdis (Charybdis) callianassa (Herbst) ?Cancer callianassa Herbst 1789, pl. 54 (fig. 7) (fide Leene, 1938). Charybdis (Goniosoma) callianassa (Herbst).—Chopra, 1935, pp. 489-491, fig. 11, pl. 9 (fig. 1). Charybdis (Charybdis) callianassa (Herbst).—Leene, 1938, pp. 81-84, figs. 41-43.— Stephenson, Hudson, and Campbell, 1957, pp. 493-495, figs. 1B-D, 2C, 3D, pls. 1 (fig. 2), 4A. Marertau.—Malay Peninsula: East coast Gulf of Thailand, 8S. of NO. 3556 PORTUNID CRABS—-STEPHENSON AND REES 9 Trat Bay off Lam Son Village, near Cambodian border, 11°57’00’N.., 102°44’45”E. GVF Sta. 27, Oct. 29, 1957, 1 male (without chelipeds) (32 mm.). Distripution.—Karachi to eastern Australia. Charybdis (Charybdis) curtilobus, new species PLATE 1A Mareriau.—Philippines: Sta. 5594, near Mount Putri, 4°14’20’N., 117°53’12”E., 11 fm., Sept. 30, 1909, Alb., 1 female (21 mm., holo- type, cat. no. 111729). Description—Front: 6-toothed. Medians rounded, protruding slightly beyond submedians, separated by narrow incision. Subme- dians broader, lobelike, with inner border curved and outer border running straight backward. Laterals roundedly triangular. Front distinctly separated from and prominent beyond inner supraorbital angles. These strongly arched and short (about as broad as lateral frontal lobes). Anterolateral teeth: Six. First four stout and square cut, but with distinct anterior points becoming sharper in more posterior teeth. Fifth tooth sharp and the smallest. Sixth distinctly the largest. Carapace: Relatively broad, B./L.=1.7. Posterolateral borders strongly convergent forming curve with nearly straight posterior border. Bearing sparse pile of fine hairs on frontal and lateral por- tions, microscopically granular throughout. Following ridges present: Frontals (short), protogastrics (convex anteriorly), mesogastrics (con- tinuous sinuous line), epibranchials (interrupted at cervical groove but not in midline), cardiac (obscure), anterior mesobranchial (short, oblique), posterior mesobranchial (forward surface only ridgelike, remainder an extensive granulated area). Chelipeds: Right larger. Granular and pilose on upper and outer surfaces. Anterior border arm with two spines, posterior border smooth. Wrist normal. Upper surface palm with two granular carinae terminating in spines about four-fifths distance along seg- ment. Outer surface with three carinae, uppermost the most granu- lar. Inner surface with ill-developed, smooth, central carina. Under surface microscopically granular. Fifth lee: Merus short (L./B.=1.7), with posterior spine. RemARKS.—This species resembles C. rathbuni Leene (1938, pp. 97-99, fig. 52) but differs as follows: a. Broader carapace (B./L. ca. 1.7 instead of ca. 1.4) due mainly to the longer last anterolateral tooth. b. Narrower inner orbital lobe which is not much wider than the lateral frontal lobe. The specific name refers to this feature. 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 c. Center of mesobranchial areas of carapace is covered with a large diffusely granular area which has no counterpart in C. rathbuni. This effectively obliterates the two most posterior mesobranchial ridges. d. Only two spines on upper surface of palm (excluding that at wrist articulation) against four in C. rathbuni. Charybdis (Charybdis) feriatus (Linnaeus) Cancer feriatus L. 1758, p. 627 (fide Holthius, 1962, pp. 234-235). Cancer cruciatus Herbst, 1790, pl. 8 (fig. 58) (fide Alcock, 1899). Charybdis (Charybdis) cruciata (Herbst).—Leene, 1938, pp. 24-27, figs. 1, 2.— Stephenson, Hudson, and Campbell, 1957, pp. 495, 497, figs. 2E, 3F, pls. 1 (fig. 3) 4B.—Crosnier, 1962, pp. 75-77, figs. 130-132. Charybdis cruciata (Herbst).—Sakai, 1939, pp. 403-404, pl. 82 (fig. 3)—Miyake, Sakai, and Nishikawa, 1962, p. 128 (record only). Marerriau.— Malay Peninsula: Sta. 10001, from Thailand, pur- chased, received Aug. 24, 1955, R. E. Elbel, 1 male (120 mm.); Thailand, NE. of Goh Krah I., middle of Gulf of Thailand, 08°38.2’N., 101°14.6’E., GVF Sta. 40, Oct. 31, 1957, 1 male (55 mm.). ReMARKS.— Justifications for changing the long-accepted name of this species are given by Holthius (1962). DistriButTion.— Madagascar, South Africa, India, to Japan and Australia. Charybdis cf. feriatus (Linnaeus) Mareriau.—China: Namru-2, Taipei, 12 mi. S. of Tau Hsui, Sta. AT-51, beach, 1957, R. E. Kuntz, 1 male (27 mm.). Remarks.— Probably a juvenile, as evidenced by the flexibility of the male pleopods. With many features of C. fervatus, except the basal antennal joint does not touch the front. Charybdis (Charybdis) hellerii (A. Milne Edwards) Goniosoma hellerii A. Milne Edwards, 1867, p. 282. Charybdis (Goniosoma) merguiensis Chopra, 1935, pp. 484-486, fig. 8. Charybdis (Charybdis) hellerii (A. Milne Edwards).—Leene, 1938, pp. 44-49, figs. 15-17. Charybdis merguiensis Chopra.—Barnard, 1950, p. 168, figs. 27d, 32b. Charybdis hellerit (A. Milne Edwards)—Edmondson, 1954, pp. 247-248, figs. 32e, f. Charybdis (Charybdis) helleri (A. Milne Edwards).—Stephenson, Hudson, and Campbell, 1957, pp. 497-498, figs. 1A, 2I, 3J, pls. 1, (fig. 4), 4C, 5B.— Crosnier, 1962, pp. 77-78, figs. 183-135, pl. 5 (fig. 1). Martertau.— Malay Peninsula: Rangoon, Burma, Gordon E. Gates, 1 male (85 mm.). Melanesia: Sta. 4, Bougainville I., received Jan. 10, 1945, W. A. Bartos, 1 female (72 mm.). Distripution.— Mediterranean and East African coast to Hawaii, including Australia. NO. 3656 PORTUNID CRABS—STEPHENSON AND REES 11 Charybdis (Charybdis) japonica (A. Milne Edwards) Goniosoma japonicum A. Milne Edwards, 1861, p. 373. Charybdis (Charybdis) japonica (A. Milne Edwards).—Leene, 1938, pp. 30-35, figs. 5-7. Charybdis japonica (A. Milne Edwards).—Sakai, 1939, pp. 400-401, pl. 45 (fig. 5).— Miyake, Sakai, and Nishikawa, 1962, p. 128 (record only). MareriAu.—China: Tan Shui Beach, northwest coast of Taiwan, Aug. 15, 1965, R. E. Kuntz, 1 male. Japan: Tangku, Fan Memorial Institute of Biology, May 8, 1929, C. J. Shen (coll. and ident.), 1 male, 1 female. MEASUREMENTS.— Males, 34, 63 mm.; female, 73 mm. DistRIBUTION.—Red Sea, China, and Japan. As Leene notes, the absence of specimens from intermediate localities is surprising. Charybdis (Charybdis) miles (de Haan) Portunus (Charybdis) miles de Haan, 1835, p. 41, pl. 11 (fig. 1). Charybdis (Charybdis) miles (de Haan).—Leene, 1938, pp. 38-42, figs. 10-13.— Stephenson, Hudson, and Campbell, 1957, pp. 500-501, figs. 2H, 3I, pls. 2 (fig. 3), 4F. Charybdis miles (de Haan).—Sakai, 1939, p. 405, pl. 46 (fig. 2).—Miyake, Sakai, and Nishikawa, 1962, p. 128 (record only). Marterrau.—Philippines: Sta. 5442, west coast of Luzon, South Fernando Point Light, 16°30’36”N., 120°11’06”E., 45 fm., coarse sand, May 11, 1909, Alb., 1 female (22 mm.). DisTRIBUTION.—India to Japan and Australia, including South China Sea and Hong Kong. Charybdis (Charybdis) natator (Herbst) Cancer natator Herbst, 1789, pl. 40 (fig. 1) (fide Leene, 1938). Charybdis (Charybdis) natator (Herbst).—Leene, 1938, pp. 93-97, figs. 50, 51.— Stephenson, Hudson, and Campbell, 1957, pp. 501-502, figs. 2G, 3H, pls. 2 (fig. 4), 4J.—Crosnier, 1962, pp. 82-83, figs. 143-144, pl. 13 (fig. 2). Charybdis natator (Herbst).—Sakai, 1939, p. 407.—Barnard, 1950, pp. 169-170, Mareriau.—Malay Peninsula: Singapore, Mar. 20, 1934, Herre collection, 1 male. Australia: Low tide, Little Lagoon, Groote Eylandt, on sand bar, about one-fourth mile from south shore, speared by native, May 11, 1948, 1 male. MrEASuREMENTS.—Males, 55, 98 mm. DistriBuTION.—Southeast Africa to Japan and Australia. Charybdis (Charybdis) orientalis Dana Charybdis orientalis Dana, 1852a, p. 285, pl. 17 (fig. 10); 1852b, p. 85.—Sakai, 1939, pp. 407-408, pl. 83 (fig. 2). 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Charybdis (Charybdis) orientalis Dana.—Leene, 1938, pp. 68-72, figs. 32-34 (excluding some synonymy).—Stephenson, Hudson, and Campbell, 1957, pp. 502-503, figs. 2B, 3B, pls. 3 (fig. 1), 4G.—Crosnier, 1962, pp. 80-81. Not Charybdis orientalis Edmondson, 1946, p. 281, fig. 173e (=C. hawaiensis). Marertau.—China: Tan Shui Beach, northwest coast of Taiwan, Aug. 15, 1956, R. E. Kuntz, 1 female (44 mm.). Distripution.—Madagascar and East Africa to Japan and Australia. Subgenus Goniohellenus Alcock, 1899 Charybdis (Goniohellenus) truncata (Fabricius) Portunus truncatus Fabricius, 1798, p. 365 (fide Leene, 1938). Charybdis (Goniohellenus) truncata (Fabricius)—Leene, 1938, pp. 118-121, figs. 66, 67—Stephenson, Hudson, and Campbell, 1957, pp. 503-504, figs. 2D, 3E, pls. 3 (fig. 3), 41—Stephenson, 196la, p. 117.—Crosnier, 1962, pp. 87, 89, figs. 149-150, pl. 8 (fig. 1) —Miyake, Sakai, and Nishikawa, 1962, p. 128 (record only). Martertau.—Philippines: Sta. 5442, west coast of Luzon, South Fernando Point Light, 16°30’36”N., 120°11’06”E., 45 fm., coarse sand, May 11, 1909, Alb., 1 male (23 mm.). Distripution.—Madagascar, Ceylon, India, Japan, Philippines, and Australia. Charybdis (Goniohellenus) vadorum Alcock Charybdis (Goniohellenus) hoplites var. vadorum Alcock, 1899, p. 67. Charybdis (Goniohellenus) vadorum Alcock.—Leene, 1938, pp. 114-117, figs. 63-65. Marteriau.—Philippines: Sta. 5442, west coast of Luzon, South Fernando Point Light, 16°30’36”N., 120°11’06”E., 45 fm., coarse sand, May 11, 1909, Alb., 1 male (17 mm.), 1 female (21 mm.). Remarks.—The carapace ornamentation is slightly different from that figured by Leene, the mesobranchial granulated areas being distinctly separated from the posterolaterals. The frontal incisions separating the lateral frontal teeth from the remainder are deep and relatively broad, more so than Leene’s figure 63. This causes diffi- culties in her key (pp. 22-23). Distripution.—Red Sea and Persian Gulf to Hong Kong. Subgenus Gonioneptunus Ortmann, 1893 Charybdis (Gonioneptunus) bimaculata (Miers) Goniosoma variegatum var. bimaculatum Miers, 1886, p. 191, pl. 15 (fig. 3). Charybdis (Gonioneptunus) bimaculata (Miers).—Leene, 1938, pp. 126-129, figs. 70, 71.—Stephenson, Hudson, and Campbell, 1957, pp. 504-505, figs. 2J, 3K, pls. 3 (fig. 4), 4H, 5A. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 13 Charybdis bimaculata (Miers).—Sakai, 1939, pp. 410-412, figs. 10, 11—Miyake, Sakai, and Nishikawa, 1962, p. 128 (record only). Charybdis (Gonioneptunus) subornata (Ortmann).—Leene, 1938, pp. 122-125, figs. 68, 69. Charybdis (Gonioneptunus) whitelegget (Ward).—Leene, 1938, pp. 125-126. MareriaLt.—Philippines: Sta. 5391, Tubig Point, Destacado 1., 12°13/05”N., 124°05’03”E., 118 fm., Mar. 13, 1909, Alb., 1 female (15 mm.). Remarks.—As indicated by Sakai (1939, pp. 410-412), C. subornata is probably a synonym of C. bimaculata. Stephenson, Hudson, and Campbell (1957, pp. 504-505) supported this synonymy, adding C. whitelegger. The present specimen has long last anterolateral teeth as in Leene’s figure (fig. 68) of C. subornata. Distrisution.—India and Maldives to Japan and eastern Australia. Subgenus Goniosupradens Leene, 1938 Charybdis (Goniosupradens) erythrodactyla (Lamarck) Portunus erythrodactylus Lamarck, 1818, p. 259. Thalamita erythrodactyla (Lamarck).—H. Milne Edwards, 1834, p. 464. Charybdis (Goniosoma) erythrodactyla (Lamarck).— Rathbun, 1906, p. 872, pl. 4. Charybdis (Goniosupradens) erythrodactyla (Lamarck).—Leene, 1938, pp. 134-137, figs. 77-80.—Crosnier, 1962, p. 86, pl. 7 (fig. 1). Charybdis erythrodactyla (Lamarck).—Sakai, 1939, pp. 408-409, pl. 83 (fig. 1).— Holthuis, 1953, p. 6.—Forest and Guinot, 1961, p. 30. Matertau.—Marshalls: Outer reef, Namu I., Bikini Atoll, col- lected at night, by light, Aug. 7, 1947, F. M. Bayer, 2 males. Tuamotus and Societies: Bora Bora, Farepiti Point Sta. 50-57, Apr. 23, 1957, Bredin Exped., 1 ovig. female; Sta. 126-57, reef, Nuarei Bay, Moorea, May 12, 1957, Bredin Exped., 1 male. MEASUREMENTS.—Males, 55-94 mm.; ovig. female, 81 mm. DistriBuTIon.—Mauritius and Red Sea to Hawaii, Tahiti, and Marquesas, including Japan. Not recorded from Australia. Charybdis (Goniosupradens) obtusifrons Leene Charybdis obtusifrons Leene, 1936, p. 124, figs. 11, 12.—Sakai, 1939, pp. 409-410, pl. 83 (fig. 3). Charybdis (Goniosupradens) obtusifrons Leene, 1938, pp. 140-148, figs. 85-87.— Sankarankutty, 1961, pp. 123-124.—Crosnier, 1962, pp. 84-85, figs. 146, 146 bis a-e, pl. 6 (fig. 2). Martertau.—Melanesia: Bougainville I., received Jan. 10, 1945, W. A. Bartos, 2 males (dried) (8, 13 mm.). Remarks.—Because of their fragile state, detailed examination was impossible. Compared with Leene’s figures and description, the differences in present specimens are: Submedian frontal teeth are broader; protogastric ridges are equidistant between frontals and 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 mesogastrics; and mesobranchial ridges are longer and slightly in- clined. DistTRIBUTION.— Madagascar, Red Sea, India, and Japan. Genus Portunus Weber, 1795 Portunus acerbiterminalis, new species FIGuRE 1; PLATE 1B Materiau.—East Africa: U-48-74 Saudi Arabia, Tarut Bay, Ras Tanura, fish pier at night, May 26, 1948, Erdmann, 1 male (frag- mented, two walking legs, one fifth leg missing) (16 mm., holotype, cat. no. 112-656); U-48-89, Saudi Arabia, Tarut Bay, 1% mi. N. of Ras Tanura, 100 yds. offshore, June 7, 1948, Erdmann, 1 male (19 mm.), 1 female (missing most appendages, with only one left cheliped, one walking leg, one left fifth leg) (ca. 19 mm.). All specimens soft, initially formalin preserved. Descriprion.—Front: 4-lobed, upraised, protruding well beyond inner supraorbital angles. Medians much smaller, more rounded, and less projecting than laterals, which are obtusely rounded. Inner supraorbital angle very obtuse. ‘Two fissures on upper border of orbit, both slightly open. Single suborbital fissure. Suborbital tooth reasonably prominent. Anterolateral teeth: Nine, first blunt, remainder sharp; fourth, sixth, and eighth slightly smaller than intervening odd-numbered teeth. Ninth distinctly elongate. Carapace: Reasonably broad, approximately twice as broad as long, with conspicuously elevated areas (some almost tubercular), separated by regions with dense pubescence. Posterior-posterolateral junction elevated, forming an obtuse or right angle. Following areas recog- nizable: Protogastrics diffuse and broadly separated; mesogastrics each divided into two portions, median with almost ridgelike termination, lateral with dense cluster of granules to form a tubercle; metagastric continuous across midline with ridged border; central longitudinal patch present, running from level of protogastrics but not quite confluent with metagastrics; between metagastrics and cardiacs a small, central, transverse, granular patch; paired cardiacs, conspicu- ously elevated, each bearing a tubercle, broadly separate in midline; median postcardiac an unusually conspicuous, tuberculate elevation; lateral postcardiacs conspicuous, but diffuse; anterolaterals, sizable patch opposite third anterolateral tooth, smaller one opposite fifth, and smaller still opposite seventh; epibranchial feebly developed laterally, well developed toward center; anterior and posterior meso- branchials present, former with short ridgelike border sometimes NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 15 tuberculate, latter variable, either diffuse or tuberculate; postero- laterals well developed, sometimes merging with posterior meso- branchials. Two additional small, paired, granular patches present, one midway between epibranchials and metagastrics, the other midway between anterior mesobranchials and cardiacs. Figure 1.—Portunus acerbiterminalis, new species, male holotype: a, pleopod; b, pleopod tip, under surface; c, abdomen. (Scales=a, 1 mm.; b, 0.1 mm.;¢, 5 mm.) Chelipeds: Short, stout, right stouter than left, with squamiform markings on under surface. Posterior border of arm with two well- developed spines, anterior border with four spines. Wrist with well-developed spines on inner and outer surfaces. Hand with well- developed shelflike carina separating upper from inner surface, and bearing on each left chela a terminal spine (none on right chela). T'wo carinae on outer surface of hand, both regularly granular, lowermost very prominent. Well-developed granular carina on inner surface of hand. Fingers short, stout but thin. 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Fifth leg: Merus, relatively stout (B/L ca. 1.5), posterodistal border coarsely serrated. Third maxilliped: Anteroexternal angle conspicuously expanded in lateral direction. Male abdomen: Elongate, ultimate segment lanceolate with rounded tip, twice as long as broad. Penultimate segment, distally parallel sided, then converging regularly, 1.4 times as long as broad. Male first pleopod: Short, stout, robust, smoothly but conspicuously curved. Subterminal armature on outer surface, of scattered spinules with portions of three or four (only) visible in profile view. Inner side without subterminal armature or with a single bristle in profile view. Under surface with scattered reasonably numerous spinules. Remarks.—This species differs considerably from P. hastatordes, although it keys out similarly in Stephenson and Campbell (1959). It differs obviously in the carapace ornamentation. The strongly embossed carapace is reminiscent of P. petreus, but the present species differs as follows: Spines on inner and outer sides of wrist are approximately equal size; last anterolateral tooth is dis- tinctly longer (see Crosnier, 1962, fig. 86, pl. 4, fig. 1); has angled posterior-posterolateral junctions of the carapace. It resembles P. rugosus in its embossed carapace but differs in the details of granulation and in having a 4-lobed front. It possibly resembles P. tuberculosus more closely than any other species but differs in having: A more conspicuously produced antero- external border of the merus of the third maxilliped; less protruding median frontal teeth; a much larger first anterolateral tooth; a longer ninth anterolateral tooth; a differently shaped epibranchial carapace ridge; and the posterior border of the arm of the cheliped bearing two distinct spines instead of one and a doubtful second. Portunus argentatus (A. Milne Edwards) FIGURE 2 Amphitrite argentata White, 1847, p. 146 (descriptio nulla). Neptunus argentatus A. Milne Edwards, 1861, pp. 332, 339, pl. 31 (figs. 4, 4a, 4b). Portunus (Achelous) argentatus (A. Milne Edwards).—Rathbun, 1906, p. 871.— Edmondson, 1954, pp. 238-239, figs. 14, 15. Portunus argentatus (A. Milne Edwards).—Stephenson, 1961a, pp. 105-106, figs. 1F, 3D, pls. 2 (fig. 2), 4D, 5A.—Crosnier, 1962, pp. 50-51, figs. 71, 75, 77, 80, 81, pl. 3 (fig. 1). Matrriau.—As indicated below, two forms are distinguishable in adult males, designated forms A and B respectively. These are listed separately, as are females and juveniles. NO. 3556 PORTUNID CRABS—-STEPHENSON AND REES 17 MALES (FORM A) China: Sta. 5308, China Sea, vicinity Hong Kong, 21°54’N., 115°42’E., 62 fm.; Nov. 4, 1908, Alb.; 1 Sacculina infected male; Sta. 5309, China Sea, vicinity Hong Kong, 21°53’N., 115°51’E., 62 fm., green mud, Novy. 4, 1908, Alb., 4 males. PTE] Lene UPC IT ver a — Ficure 2.—Portunus argentatus (A. Milne Edwards), male: a, form B, pleopod, Philippines, Sta. 5442, Alb., 30 mm.; b, abdomen, same specimen; c, form A, abdomen, Philippines, Sta. 5442, Alb.,29 mm. (Scales=a, 1 mm.; b, c, 5 mm.) Philippines: Sta. 5152, Pajumajan I., 5°22’55”N., 120°15’45”E., Feb. 18, 1908, Alb., 1 male; Sta. 5442, west coast of Luzon, South Fernando Point Light, 16°30’36”N., 120°11/06”E., 45 fm., coarse sand, May 11, 1909, Alb., 16 males (1 damaged). 221—520—67——_2 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 MALES (FORM B) Philippines: Sta. 5442, west coast of Luzon, South Fernando Point Light, 16°30’36”N., 120°11’06”E., 45 fm., coarse sand, May 11, 1909, Alb., 3 males. FEMALES China: Sta. 5303, China Sea, vicinity Hong Kong, 21°44’N., 114°48’E., 34 fm., blue mud, Aug. 9, 1908, Alb., 1 female; Sta. 5304, China Sea, vicinity Hong Kong, 21°46’N., 114°47’E., 34 fm., black mud, Aug. 9, 1908, Alb., 1 female; Sta. 5309, China Sea, vicinity Hong Kong, 21°53’N., 115°51’E., 62 fm., green mud, Nov. 4, 1908, Alb., 2 females; Takao, Formosa, Dec. 3 and 4, 1914, Fred Baker, 2 females (1 soft and fragmented). Philippines: Port Binanga, ship’s side, 6 fm., Jan. 8, 1908, Alb., 1 female (deformed abdomen); Panabutan Bay, Mindanao, electric light, soft mud and sand, Feb. 5, 1908, Alb., 1 female; Sta. 5152, Pajumajan I[., 5°22'55”N., 120°15/45”E., Feb. 18, 1908, Alb., 1 female; Sta. 5157, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°12’30”N., 119°55’50”E., 18 fm., fine sand, Feb. 21, 1908, Alb., 1 female (damaged); Sta. 5442, west coast of Luzon, South Fernando Point Light, 16°30’36” N., 120°11’06”E., 45 fm., coarse sand, May 11, 1909, Alb., 22 females (4 ovig., 2 Sacculina infected, 1 damaged), 1 damaged unsexable specimen; Sta. 5480, Tacbue Point (Leyte), 10°44’36”N., 125°19’K., 62 fm., fine sand, July 29, 1909, Alb., 1 female. JUVENILES Philippines: Port Binanga, ship’s side, 6 fm., Jan. 8, 1908, Alb., 2 juvs. MEASUREMENTS.—Males (form A), 21-39 mm. (1 damaged male, ca. 40 mm.); males (form B), 30-83 mm.; females, 18-36 mm.; ovig. females, 35-37 mm.; juvs. 14 mm.; damaged unsexable specimen, ca. 27 mm. Remarks.—In most specimens, and after long preservation, there are no detectable spots on the dactylus of the swimming legs and the general nacreous luster is far from obvious. Most males have ple- opods resembling those figured and described by Stephenson (1961a) with short, curved, distally swollen appendages bearing a complex and dense subterminal armature. These possess abdomens with rel- atively broad, sharply converging penultimate segments. In three cases male pleopods are quite different, with long gradually tapering appendages bent at more than aright angle near their centers, and with inconspicuous subterminal armature consisting of small spi- nules on the outer surface (fig. 2a). These possess abdomens with rel- atively narrow, gradually converging penultimate segments. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 19 This second form of pleopod resembles that figured by Edmondson (1954, fig. 14 c,d), differing only in subterminal armature which Edmondson shows as a distinct row of sizable bristles on the outer sur- face. This form of pleopod has also been figured by Crosnier (1962, figs. 77, 80, 81). The corresponding abdomen is apparently inter- mediate between those mentioned above. Apart from male pleopods and abdomens, no structural heterogene- ity could be found in the present material. There are two possibil- ities : a. The material belongs to two distinct species, separable only in adult males and effectively only by pleopods. To accept this would involve several difficulties. First, the two “species”? would be sym- patric in distribution, or at least seriously overlapping, since both occur in one collection. Second, to establish which was the true P. argentatus would necessarily await reexamination of Milne Edwards’ material. Third, it would involve the assumption of reproductive isolation, with females being capable of fertilization with only one or another type of male. Until the detailed role of pleopods in ferti- lization has been determined, this must remain doubtful. Fourth, there are other cases of heterogeneity in male pleopods (e.g., Thalamita stimpsom, T. danae, P. orbitosinus—see text), and these should be treated similarly. This presents particular difficulty in P. orbitosinus where pleopod variation is more continuous than in the present case. b. The second possibility is that a species can be polymorphic as regards male pleopods. As stated in the Introduction, for the purposes of the present paper, P. argentatus was taken as the ‘‘test case” of the concept of a portunid species. This concept implies a distinct morphological gap between other species in general facies as well as pleopod structure. Thus there are two male forms of P. argentatus, form A corre- sponding with that figured and described by Stephenson (1961a) and form B corresponding to that given in figure 2a. Possibly Edmond- son’s (1954) and Crosnier’s (1962) specimens belong to a third form, resembling form B in general shape of the pleopod, but differing in its much better developed bristles. DistripuTion.—Natal to Honolulu, including Japan and Australia. Portunus brockii (de Man) Neptunus brockii de Man, 1887a, pp. 328-331, pl. 13 (fig. 4). Neptunus (Hellenus) brockii de Man.—Alcock, 1899, pp. 438-44.—Shen, 1937, p. 111, figs. 7, 8e, 8f. Portunus brocki (de Man).—Stephenson and Campbell, 1959, pp. 106-107, figs. 2G, 3G, pls. 2 (fig. 3), 4G, 5G. Marerrau.—Philippines: Subig Bay, China Sea, off southern Luzon, shore, seine, sand, Jan. 7, 1908, Alb., 1 male; Sta. 5160, Tinakta 20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 I., Sulu Archipelago, Tawitawi Group, 5°12’40”N., 119°55’10’E.., 12 fm., sand, Feb. 22, 1908, Alb., 1 male; Catabata, Mindanao, below mouth of river, seine, May 20, 1908, Alb., 1 male; Port San Vicente, 130 ft. seine, Nov. 18, 1908, Alb., 1 male, 1 ovig. female; Buena Vista, Guimaras Is., Iloilo Strait, seine in mouth of river, Jan. 14, 1909, Alb., 1 male; near Mariveles, Luzon, 1913, A.M. Reese, 1 male. Palau: Sta. 12-832, Madalai district, west end of Koror I., mangrove shore grading into mud and sand flat, July 9, 1955, GVF, 1 male. MEASUREMENTS.—Males, 12-29 mm.; ovig. female, 15 mm. Remarxs.—A male “16 mm., Port San Vicente, Nov. 18, 1908 Alb.” has a very broad, female-like abdomen, but fully developed male pleopods are present. Distripution.—Amboina, Andamans, Singapore, and Australia. Portunus dubius (Laurie) FIGuRE 3; PLATE 2A Neptunus (Achelous) dubia Laurie, 1906, pp. 416-417, fig. 9. MareriAu.—Philippines: Sta. 5140, Jolo Lt., 6°08’45’N., 121°03’E., 76 fm., fine coral sand, Feb. 15, 1908, Alb., 1 female (13 mm.); Sta. 5432, Corandagos I., 10°37’50”N., 120°12’E., 51 fm., Apr. 8, 1909, Alb., 1 male (12 mm.). Description.—Front: Protruding beyond inner supraorbital angles and 4-toothed. Teeth sharp and directed dorsally, laterals the larger and inclined outward. Inner supraorbital angles rounded, upper border of orbit with single fissure, lower border with acute forward-directed tooth. Anterolateral teeth: Nine, first the stoutest, gradually decreasing in size with eighth the smallest and ninth slightly larger. Carapace: Very long, breadth about 1.3 times length. Surface microscopically granular among which major patches of granules are visible. Protogastrics diffuse and merging with ill-developed meso- gastrics, which in turn connect by a central granular line to trans- versely situated metagastrics. Patch behind orbit in line with third anterolateral tooth; further diffuse patch opposite fourth to ninth teeth, and merging with epibranchial. Paired cardiac merges with median postcardiac. Pair of lateral postcardiacs. Mesobranchials large and diffuse but terminates abruptly on posterolateral margins by granular ridges. Fifth leg: Merus short, length 1.8 times breadth, and with finely denticulated posterior margin. Propodus with fine hairs but no spines. Third maxilliped: Hirsute and produced laterally. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 2h Male abdomen: Penultimate segment with regularly tapering sides, ultimate segment triangular. Male first pleopod: Grossly swollen so that normal distinction be- tween basal and terminal elongate portions cannot be made. Inner side curves regularly and bears forwardly directed bipinnate bristles along half its length. Outer surface bears conspicuous flattened lobe thinly beset with elongate bipinnate bristles or hairs, these extending to upper surface where they are denser. Appendage terminates Ficure 3.—Portunus dubius (Laurie), male, Philippines, Sta. 5432, Alb.: a, pleopod; b, abdomen. (Scales=a, 1 mm.; b, 5 mm.) abruptly and bears subterminal aperture directed upward. Folds of cuticle surrounding this aperture give a rosette appearance and are densely covered with minute inwardly directed spinules. Remarks.—This species resembles P. orbicularis (Richters) which has been redescribed recently by Crosnier (1962). It differs in pos- sessing a spiniform front, more granular carapace, and a male pleopod which is different from all other known species of the genus. The abdomen of the single male in the present collection is nar- rower, and the ultimate segment is longer than figured by Laurie (1906, fig. 9). Distrisution.—Previously from Gulf of Manaar and Negombo, Ceylon. Portunus euglyphus (Laurie) Figure 4; PLATE 28 Neptunus (Amphitrite) euglyphus Laurie, 1906, pp. 413-414, figs. 6, 7. 22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Portunus (Achelous) granulatus Edmondson, 1954, p. 239, figs. 16a, b (not fig. 17a). Not Lupea granulata H. Milne Edwards, 1834, p. 454. Marerrau.—Philippines: Guijulugan, Negros, shore seine, Apr. 2, 1908, Alb., 1 male (19 mm.); Paluan Bay, Mindoro, 130 ft. seine, Dec. 11, 1908, Alb., 1 male (18 mm.). Description.—Front: 4-lobed, outer lobes roundedly triangular and protruding beyond the much smaller, similar shaped medians. Ficure 4.—Portunus euglyphus (Laurie), male, Philippines, Guijulugan, Alb.: a, pleopod; b, pleopod tip, upper surface; c, abdomen. (Scales=a, 1 mm.; b, 0.1 mm.; c, 5 mm.) These merge with epistomial projection so that in some views front appears 3-lobed. Inner orbital angle short. Two incisions on upper border and rounded tooth on lower. Anterolateral teeth: Anterolateral borders practically straight. First tooth relatively stout, fourth and fifth the smallest, then in- creasing in size to ninth, much the largest. Apart from first, all bear sharp, forwardly directed points. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 23 Carapace: With inconspicuous pubescence and distinct granulated areas. Frontals and orbitals poorly developed. Mesogastrics and metagastrics distinct granular transverse lines connected by broad central granular patch. Protogastrics with larger anterior and smaller posterior region, latter with ridgelike anterior boundary. Cardiacs, median postcardiac, and lateral postcardiacs all conspicuously raised, rounded, granular areas. Mesobranchials resolvable into three patches. Epibranchials and posterolaterals well developed. Posterior-pos- terolateral junction rounded. Chelipeds: Pubescent and markedly granular, especially on inner surfaces. Posterior border of arm with two sharp spines, anterior border with four spines. Wrist with inner and outer spine. Hand with two well-marked outer carinae, upper being granular. Two carinae on upper surface of hand, inner being the better developed and terminating in a spine just before end of segment. No recognizable carinae on inner surface of hand, this completely covered with large rounded granules showing through fairly dense pubescence. Fifth leg: Posterior borders of merus, carpus, and propodus orna- mented only with hairs. Third maxilliped: Anteroexternal angle markedly produced laterally. Male abdomen: Penultimate and ultimate segment gradually taper- ing, latter three-fifths length of former. Male abdomen and entire undersurface shining. Male first pleopod: Short and stout with undulating outer side and sharply curved tip pointing outward. Outer surface bearing long, and inner surface bearing shorter bipinnate hairs. Terminal armature ill developed and consisting of three or four minute spinules on upper surface and five or six on lower. Remarks.—In its general facies this species resembles P. gladiator Stephenson and Campbell (1959) but differs from it in having a much shorter last anterolateral spine so that the anterolateral borders appear almost without curvature, in the absence of serrations on the postero- distal border of the fifth merus, and in possessing a male pleopod unique to the genus. The species is probably closest to P. granulatus, particularly in the eranular areas of the carapace and the form of male pleopod, but differs conspicuously in carapace shape, in having a longer last anterolateral tooth, and in the characteristic form of its male pleopod. Edmond- son’s (1954) figure 16b of this pleopod obviously refers to the present species and not to P. granulatus. DistrIBUTION.—Previously known only from the Gulf of Manaar and Negombo, Ceylon, and Hawaii. 24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 The Portunus gladiator Complex Crosnier (1962) concluded that P. pseudoargentatus Stephenson (1961a) is synonymous with his P. gladvator; however, the present collection contains three specimens identical with P. pseudoargentatus and two identical with P. gladiator Crosnier. ‘These differ in a sufficient number of small particulars for P. pseudoargentatus to stand as a valid species. Crosnier suspected that numerous confusions had occurred in the past regarding P. gladiator and particularly that P. gladiator Stephenson and Campbell (1959) differed from the species he recognized under this name. Reexamination of Stephenson and Campbell’s material confirms that this is so. The penultimate segment of the male abdomen in Stephenson and Campbell’s species is not swollen in its distal third, and the ultimate segment is more rounded and elongate. (Note that the plates in Stephenson and Campbell are mislabelled, the male abdomen of P. gladiator is plate 4, figure I, not figure J.) In addition, the carapace granulation is much coarser and more obvious, with posterolaterals almost merging with mesobranchials, and these again almost merging with the epibranchials. In addition, the serrated distal border of the fifth merus bears a posterior spine, not an anterior spinous projection. Until the “true” P. gladiator has finally been decided upon, by reference to Fabricius type, the correct name for P. gladiator Stephenson and Campbell must remain un- decided. Yet other authors have described different species under P. gladiator. Thus the shape of the male abdomen figured by Sakai (1939, fig. 5a) differs from those figured similarly by Shen (1937, fig. 2b) and Crosnier (1962, fig. 72), and described by Alcock (1899) and Barnard (1950). Sakai’s figure is identical with that of P. pseudoargentatus Stephenson (1961a, pl. 4F), and his colored plate (pl. 47, fig. 3) is generally similar to P. pseudoargentatus and specifi- cally bears an identical pigmentation of the propodus and dactylus of the fifth leg. It is concluded that P. gladiator Sakai and P. pseudoargentatus are synonyms. De Haan’s (1835) description and figures of Portunus (Amphitrite) gladiator appear to resemble P. pseudoargentatus more closely than P. gladiator Crosnier in possessing a parallel-sided male abdomen. The position as regards shape of the merus of the fifth leg is more ambiguous. The left side of de Haan’s plate 1, figure 5, shows a relatively broad merus, approximating to P. gladiator Crosnier and the right side a relatively elongate one, approximating to P. pseudo- argentatus (see p. 25). Because de Haan’s plate shows pigmentation of the fifth leg resembling that of P. pseudoargentatus and because the NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 25 last anterolateral tooth points in a similar direction, his species is here regarded (tentatively) as being a synonym of P. pseudoargentatus. Portunus gladiator (Alcock) Neptunus (Amphitrite) gladiator Alcock, 1899, pp. 35-36.—Shen, 1937, pp. 101-108, figs. 2a-c. Monomia gladiator Barnard, 1950, p. 156. Portunus gladiator Crosnier, 1962, pp. 51-54, figs. 72, 76, 78, 82-83, pl. 3 (fig. 2). ?Portunus gladiator Fabricius, 1798, p. 368. Not Portunus gladiator Stephenson and Campbell, 1959, pp. 110-111, figs. 2J, 3J, pls. 3 (fig. 2), 41, 5J. Materiau.—Africa: Natal, from W. F. H. Rosenberg, 1 male (70 mm.), 1 female (66 mm.). Portunus pseudoargentatus Stephenson Portunus (Amphitrite) gladiator De Haan, 1835, p. 39, pl. 1 (fig. 5). Neptunus (Amphitrite) gladiator Sakai, 1939, pp. 390-391, text-fig. 5a, pl. 47 fig. 3). a Sa iadnoke Stephenson, 196la, pp. 109-111, figs. 2A, 3F, pls. 2 (fig. 4), 4F, 5D. ?Portunus gladiator Fabricius, 1798, p. 368. MarterraAL.—Philippines: Lembeh Strait, Celebes, June 14, 1929, Herre collection, 1 male (25 mm.). Japan: Fukura, Awaji, Y. Hirase, 1 female (28 mm.); Shimizu, Suruga, shore, Oct. 14, 1906, Alb., 1 male (20 mm.). Remarxks.—This species differs in the following details from P. gladiator, as described and figured by Crosnier (1962) (the comparison has been facilitated by Crosnier’s kindness in forwarding one of his named specimens) : a. A more strongly embossed carapace with coarser granulation. In particular the cardiac, lateral postcardiacs, posterolaterals, and two mesobranchials are larger and less diffuse than in P. gladiator. b. Form of male abdomen. The ultimate segment relatively longer and without concave lateral borders, penultimate segment not swollen in distal third. c. Relatively narrower fifth merus. In P. pseudoargentatus breadth is about 0.75 times length, in P. gladiator about 0.9 times. d. Differently armed anterodistal border of fifth merus. In P. pseudoargentatus it is finely serrated, in P. gladiator it bears small but distinct teeth. e. The last anterolateral tooth is not directed slightly posteriorly, as in P. gladiator. Portunus granulatus (H. Milne Edwards) Figure 5 Lupea granulata H. Milne Edwards, 1834, p. 454. Neptunus (Achelous) granulatus (H. Milne Edwards).—Alcock, 1899, pp. 45-46.— Sakai, 1939, p. 397, pl. 81 (fig. 3). 26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Portunus (Achelous) granulatus (H. Milne Edwards).—Rathbun, 1906, p. 871, pl. 41 (fig. 6); 1911, p. 205, pl. 15 (fig. 10).—Balss, 1938, p. 31—Edmond- son, 1954, p. 2389 (in part). Neptunus granulatus (H. Milne Edwards).—Balss, 1924, p. 3 (excluding synonymy). Portunus granulatus (H. Milne Edwards).—Stephenson and Campbell, 1959, pp. 108-110, figs. 21, 31, pls. 3 (fig. 1), 4J, 51 (not pl. 41 as stated in text) .— Sankarankutty, 1961la, p. 104; 1961b, p. 124.—Crosnier, 1962, pp. 57-58, figs. 89, 92, 94a, b. Not Portunus (Achelous) granulatus Edmondson, 1954, figs. 16a, b = P. euglyphus (Laurie) 1906. AA A AS Ficure 5.—Portunus granulatus (H. Milne Edwards), male abdomens: a, Hawaii, shallow water Mokuleia, Degener, 27 mm.; b, Hawaii, reef off Mokuleia, Degener, 26 mm.; ¢, Philippines, Capurmypugan Pt., A/b., 26 mm.; d, Marianas, Agfayan Bay, Baker, 23 mm.; e, Philippines, Pandanon Is., Mar. 24, 1909, A/b., 28 mm., right branchial area swollen. (Scale=5 mm.) MateEriAuL.— Philippines: Zamboanga, Mindanao, May 1904, Dr. E. A. Mearns, 1 male; Capurmypugan Point, Mindanao, 150 ft. seine, May 9, 1908, Alb., 3 males, 1 female, 4 ovig. females; Galera Bay, Verde Is. Passage and Batanga Bay, 150 ft, seme, June 9, 1908, Alb., 1 male; Tara Is., Mindora Strait, 130 ft. seme, Dec. 15, 1908, Alb., 1 ovig. female; Sirinas I., Nakoola Bay, near Alphonzo XIII, Dec. 30, 1908, Alb., 1 damaged male; near Sta. 5412, Pandanon Is., between Cebu and Bohol, seine, Mar. 23, 1909, Alb., 1 male; Pandanon Is., between Cebu and Bohol, seine, 130 ft., Mar. 24, 1909, Alb., 4 males. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 27 Melanesia: Pacific Ser. A, received Nov. 13, 1944, Lt. George H. Penn, Malaria Control Team 28, Com. M. J. B. Rons, 7th Fleet, Advance Base 3, 1 male [Amsterdam Island, then Dutch New Guinea, coll. Aug. 15—-Oct. 3, 1944, fide F. A. Chace, Jr.]; Florida I., Solomon Is., Sta. P2, Oct. 28, 1944, R. E. Kuntz, 2 females, 1 ovig. female. Marianas: Sta. 112-65, Ajayan River, Guam, June 19, 1945, D. H. Johnson, 2 females; Sta. 110-63, Bijia Point, Guam, June 22, 1945, McElroy and Markley, 9 males, 7 females; Agfayan Bay, Guam, 145/R.H.B. 150, July 7, 1945, R. H. Baker, 5 males, 1 female; Ackang Bay, Guam, Sta. 151/R.H.B. 158, July 16, 1945, R. H. Baker, 5 males, 4 females, 1 juv. Japan: Okinawa, Oct. 5, 1945, W. B. S. Thomas, 1 male. Marshalls: E1—9, Arno Atoll, received 1952, Hiatt, 1 male. Samoa: U-39-521-591, Tutuila I., reef at Alofau, June 3, 1939, L. P. Schultz, 1 male. Hawaii: Shallow water on reef at Mokuleia, Oahu, 1938, Degener, 1 male; reef off Mokuleia, Waialua, Oahu, received June 23, 1939, Otto Degener, 2 males (1 fragmented), 1 ovig. female. Tuamotus and Societies: East side of Papetoai Bay, Moorea, poison, shore reef, 3-20 ft., Apr. 30, 1957, J. Randall, 1 male. MEASUREMENTS.— Males, 9-27 mm.; females, 6-21 mm.; ovig. females, 14-21 mm.; juv., 5 mm. RemarKs.—One specimen, a large male (27 mm., Pandanon I., Mar. 24, 1909) has the right branchial cavity swollen, presumably due to a parasite, and has a broader than usual abdomen. Crosnier has drawn attention to the remarkable variation in the form of the male abdomen. In the specimens he examined, the penultimate segment was approximately as long as broad and re- sembled those figured by Rathbun and Sakai; however, a penultimate segment much longer than broad is shown by Gordon and again by Stephenson and Campbell. In the present collection most of the specimens resemble Crosnier’s, but these intergrade to a few others with elongate penultimate and ultimate segments (figs. 5a-e). Edmondsons’ P. granulatus is discussed under P. euglyphus (Laurie). DistriputTion.— Madagascar and Red Sea to Hawaii, Samoa, and Fiji, including Japan and Australia. Portunus hastatoides Fabricius Portunus hastatoides Fabricius, 1798, p. 368.—Stephenson and Campbell, 1959, pp. 101-102, figs. 2D, 3D, pl. 1 (fig. 4), pls. 4D, 5D.—Crosnier, 1962, pp. 68-69, figs. 96, 109, 117, 122-123.—Miyake, Sakai, and Nishikawa, 1962, p- 128 (record only). Neptunus (Amphitrite) hastatoides (Fabricius).—De Haan, 1835, pp. 39-40, pl. 1 (fig. 3). Neptunus (Hellenus) hastatoides (Fabricius).—Alcock, 1899, pp. 38-39, Sakai, 1939, pp. 391-392, pl. 47 (fig. 1). Neptunus (Hellenus) hastatoides var. unidens Laurie, 1906, pp. 414-415. 28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Hellenus hastatoides (Fabricius).—Barnard, 1950, pp. 158-159. Marertau.—China: Kowloon, electric light, Sept. 14, 1908, Alb., 1 juv; Takao, Formosa, Dec. 3 and 4, 1914, F. Baker, 6 males, 1 female, 2 ovig. females. Philippines: Sandakan Bay, Borneo, seine, Mar. 2, 1908, Alb., 1 female; Sta. 5182, Antonia I., off eastern Panay, 11°36’40’N., 123°26'35”E., 26 fm., Mar. 27, 1908, Alb., 1 female; Sta. 5208, Taratara I., 11°45’53”N., 124°42’50”E., 26 fm., Apr. 14, 1908, Alb., 1 male; Sta. 5342, Endeavour Point, (S.), 10°56’55”N., 119°17'24”E., 12-25 fm., Dec. 23, 1908, Alb., 1 male; Sta. 5360, Corregidor Lt., 14°21’N., 120°41’E., 12 fm., Feb. 8, 1909, Alb., 2 females, 1 ovig. female; Sta. 5442, west coast of Luzon, South Fernando Point Light, 16°30’36”N., 120°11’06”E., 45 fm., coarse sand, May 11, 1909, Alb., 4 males, 4 females, 7 ovig. females; Sta. 5461, Caringo Is., 13°57’42"N., 123°06’42”E., 11 fm., June 14, 1909, Alb., 1 ovig. female; Sta. 5594, near Mount Putri, 4°14’20’’N., 117°53’12’E., 11 fm., Sept. 30, 1909, Alb., 1 Sacculina infected male, 1 ovig. female; Sandakan Harbor, British North Borneo, June 28, 1929, Herre col- lection, 1 female. Japan: Shimbara Gulf, Imperial University, Tokyo, 6 males, 3 females. MErASUREMENTS.—Males, 24-41 mm.; females, 25-41 mm.; ovig. females, 27-36 mm.; juv., 20 mm.; Sacculina infected male, 37 mm. Remarxks.—The third maxilliped shows slight iridescence on the last three segments of the endopodite and also on the forwardly directed portion of the fourth. This feature, presumably of some specific importance, has not been noted previously. DistripuTion.—Madagascar and East Africa to Japan, Philip- pines, and Australia. The Portunus longispinosus Complex Crosnier (1962) has suggested that two and possibly three different species have been given the name longispimosus by Rathbun (1906), by Sakai (1939), and by Stephenson and Campbell (1959), respec- tively. He also notes that Amphitrite vigilans Dana 1852, which has been synonymized with A. longispinosa Dana 1852, is probably a distinct species and is possibly identical with his own P. dranjae. In the opinion of the present authors, it will prove impossible to relate specimens to one or another of Dana’s species unless the types can be discovered. The differences in anterolateral teeth lie within the range of a single species, and neither of Dana’s figures of carapace ornamentation conform exactly to those of known species. Apart from the above species (ie., P. longispinosa (Dana), P. vigilans (Dana), and P. iranjae Crosnier, the remaining closely NO. 3656 PORTUNID CRABS—-STEPHENSON AND REES 29 related species are: P. macrophthalmus Rathbun 1906, P. longi- spinosus bidens Laurie 1906, P. emarginatus Stephenson and Camp- bell, 1959, and P. tenuicaudatus Stephenson, 1961a. Disregarding synonymy problems, most of the undamaged males in the present collection could be referred to one or another of the above species without great difficulty. Problems arose over two dissimilar sized males (10 mm. and 20 mm. carapace breadths) from the same collection (Palau, Sta. 124-1183). Although undoubtedly belonging to the same species, they differed in certain features which have been relied upon heavily in past descriptions, viz the length and acuteness of the lateral frontal teeth, and the proportions and shapes of the ultimate and penultimate segments of the male abdomen. In the smaller specimen the frontal teeth are low and rounded (cf. P. emar- ginatus); the male abdomen is short and broad, with only slightly concave outer borders of the penultimate segment and with an ultimate segment as long as broad (cf. P. longispinosus Stephenson and Campbell). In the larger specimen the frontal teeth, while low and rounded, project farther (as in P. longispinosus Stephenson and Campbell), and the male abdomen, although moderately long and with distinctly concave borders to the penultimate segment (as in P. emarginatus), has the ultimate segment slightly longer than broad. P. longispinosus Stephenson and Campbell and P. emarginatus show many points of similarity, including an absence of tuberculate eleva- tions upon the granulated areas of the carapace and quite similar pleopods. Tentatively they are treated as synonyms, and specimens are listed under P. emarginatus. While, as indicated, frontal teeth and male abdomens may have less diagnostic importance than previously thought, they still have some value. Itis believed, however, that the details of ornamentation of the dorsal surface of the carapace offer the best opportunities for separating the species, although here again there is variation. In the mesobranchial areas of the carapace both of the present specimens of P. iranjae bear elevated tubercular projections that were not present in the specimens submitted by Crosnier to the senior author. Stephenson’s comments are quoted in Crosnier’s description. Recently Described Species in P. longispinosus Complex 16 Upper surface of palm of chelipeds with a single distal spine. P. longispinosus bidens (Laurie) 1906 Upper surface of palm of cheliped with two distal spines or with a spine carol G2} yopuatiitoraam EINE 4 5 5 6 GO oO-d Ob oO 6 On OO OOo 2 2. (1) Outer frontal teeth low and rounded (granular eminences of carapace without tubercles). P. emarginatus Stephenson and Campbell, 1959 Outer frontal teeth triangular, projecting, and typically sharp. . . 3 30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 3. (2) Cheliped very long and slender, 24 times the carapace length. P. lengispinosus Sakai, 1939 Chelipeds of moderate length and robustness, less then 2% times the carapaceilengthny. 2 0. scien cosets ce tenet tot a Semmens Mae os M4 4. (3) Carapace with conspicuous anoreule elevations within the granular Patches" wwe4 es MSs ee SHES Tenet) Bere eee qogtar Carapace Gun) Peon dung tianse: tubercular elevations with- in the granular patches . . . . . P. longispinosus Rathbun, 1906 5. (4) Carapace strongly embossed, with spiniform elevations in cardiac and mesobranchial areas (lateral postcardiacs distinct and separate). P. tenuicaudatus Stephenson, 196la Carapace moderately strongly embossed with tuberculate but not spiniform elevations in the cardiac and mesobranchial areas. . . 6 6. (5) Metagastric area with two tubercles, lateral postcardiac granular patch not distinctly recognizable . . . P. macrophthalmus Rathbun, 1906 Metagastric area with short ridges but without tubercles, lateral post- cardiac granular patch distinct and separate. P. iranjae Crosnier, 1962 Portunus emarginatus Stephenson and Campbell Portunus emarginatus Stephenson and Campbell, 1959, pp. 107-108, figs. 2H, 3H, pls. 2 (fig. 4), 4H, 5H.—Crosnier, 1962, pp. 66-68, figs. 108, 112-114, 116, 120-121. ?Portunus longispinosus Rathbun.—Stephenson and Campbell, 1959, pp. 104- 106, figs. 2F, 3F, pls. 2 (fig. 2), 4F, 5F. Martertau.—Palau: Sta. 124-1183, Yos Passage, W. of Kasao Reef, 34 mi. SSE. of Ngaremdiu, E. of Urukthapel, sand, 15-18 fm., Aug. 24, 1955, 2 males (10, 20 mm.). Portunus iranjae Crosnier Portunus iranjae Crosnier, 1962, pp. 61-65, figs. 107, 110-111, 115, 118-119, pl. 4 (fig. 2). MareriAu.—Philippines: Reef at Tileg, Lubang I., July 11, 1908, Paul Bartsch, Alb., 1 male (19 mm.). Tuamotus and Societies: Papetoai Bay, Moorea, poison sta. shore reef, 3-20 ft., Apr. 30, 1957, J. Randall, 1 male (23 mm.). Portunus macrophthalmus Rathbun Portunus macrophthalmus Rathbun, 1906, p. 871, fig. 31, pl. 12 (fig. 15). Marertau.—Philippines: Sta. 5218, Anima Solo I., between Burias and Luzon, 20 fm., Apr. 22, 1908, Alb., 1 ovig. female (11.5 mm.). Remarks.—In comparison with the various species examined by the authors, the present specimen comes closest to P. tenuicaudatus but differs in having more protruding and sharper lateral frontal teeth, these being distinctly acute; in having the carapace less strongly embossed, this applying particularly to the cardiac eminences and the lateral postcardiacs, which are only indistinctly recognizable; in possessing two small tubercular eminences instead of rows of gran- NO. 3556 PORTUNID CRABS—STEPHENSON AND REES ik ules in the metagastric region; and in having longer and more slender chelipeds. These four points of difference seem to be confirmed by examina- tion of Rathbun’s plate 12, figure 5. Portunus tenuicaudatus Stephenson Portunus tenuicaudatus Stephenson, 196la, pp. 114-116, figs. 2C, 3H, pls. 3 (fig. 2), 4H, 5C. Mareriat.—Philippines: Sta. 5160, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°12’40’N., 119°55’10”E., 12 fm., sand, Feb. 22, 1908, Alb., 1 male (14 mm.). Remarxs.—This species is very close to P. longispinosus bidens (Laurie) as described and figured by Sakai. Both species have spiniform tubercles on the dorsal surface of the carapace, and many of them are identically situated. (These are not mentioned by Laurie, 1906.) However, P. tenuwicaudatus lacks the tubercles of the posterior mesobranchial regions described by Sakai, and in the holotype (but not in the present specimen) there is a tubercle in each median postcardiac region. The median frontal teeth in P. longispinosus bidens appear much larger than those in P. tenuicau- datus. It also possesses a spiniform tubercle at the end of the central carina of the upper surface of the hand which has no described coun- terpart in P. longispinosus bidens. There is marked similarity but not absolute identity in the pigmentation of recently collected speci- mens. Portunus Species Matertat.— Marshalls: 4707, Rongelap Lagoon, 2 mi. W. of Busch I., 20 fm.; June 21, 1946, Taylor, 1 male (18 mm.). Remarks.—This specimen lacks the ultimate and penultimate segments of the abdomen and is either P. longispinosus Rathbun or P. longispinosus Sakai. Its chelipeds, although long and slender, are less so than Sakai described. Portunus Species Materrtau.—Mearshalls: EL-160, Arno Atol, received 1952, Hiatt, 1 damaged male (10 mm.). Remarxks.—This specimen cannot be identified with certainty because the dorsal surface of the carapace, which bears tuberculate elevations on some of its granular patches, is damaged. Portunus orbitosinus Rathbun FIGuRE 6 Portunus (Amphitrite) gladiator de Haan, 1837, only p. 65, pl. 18 (fig. 1) (not Portunus gladiator Fabricius, 1798). Portunus (Achelous) orbitosinus Rathbun, 1911, p. 205, pl. 15 (fig. 11). 32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Neptunus (Achelous) orbitosinus (Rathbun).—Gordon, 1938, pp. 182-185, figs. 5a-g, 6c, 6d (including N. octodentata). Neptunus (Achelous) orbitospinus (Rathbun).—Sakai, 1939, p. 396, pl. 81 (fig. 2). Portunus orbitosinus Rathbun.—Stephenson and Campbell, 1959, pp. 113-114, figs. 2L, 3L, pls. 3 (fig. 4), 4L, 5L.—Stephenson, 196la, pp. 108-109.— Crosnier, 1962, pp. 55-57, figs. 88, 90-91, 93.—Miyake, Sakai, and Nishi- kawa, 1962, p. 128 (record only). b Ficurr 6.—Portunus orbitosinus Rathbun, male pleopods and abdomens, respectively. Three variants of form A: a and e, Philippines, Mariveles, No. 27, Reese; b and f, Philip- pines, Sta. 5158, Alb.; c and g, same asa ande. Form B: d and h, Philippines, Labuan Blanda I., 4/b. (Scales=a-d, 1 mm.; eh, 5 mm.) MarrrraL.—Two forms are recognized in adult males, called forms A and B, respectively. These are listed separately as are females. MALES (FORM A) Philippines: Subig Bay, China Sea off southern Luzon, shore, NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 33 seine, sand, Jan. 7, 1908, Alb., 2 males; Alongapo, Luzon, shore, Jan. 7, 1908, Alb., 1 male; Port Binanga, ship’s side, 6 fm., Jan. 8, 1908, Alb., 2 males; Sta. 5158, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°12’N., 119°54’30”E., 12 fm., Feb. 21, 1908, Alb., 1 male; Sta. 5169, Sibutu Is., 4°32’15"N., 119°22’45”E., 10 fm., Feb. 27, 1908, Alb., 1 male; Guijulugan, Negros, sta. shore, sand, gravel, mud, Apr. 2, 1908, Alb., 1 male; Capurmypugan Point, Mindanao, 150 ft. seine, May 9, 1908, Alb., 1 male; Pomt Jarnelo, Luzon, 150 ft. seine, July 13, 1908, Alb., 1 male; Hinunangan Beach, Leyte, seine, July 30, 1909, Alb., 1 male; No. 27, Mariveles, Luzon, June 27, 1913, A. M. Reese, 6 males; no. 33, other data as last, 13 males; near Mari- veles, Luzon, 1913, A. M. Reese, 2 males; Cebu, 1929, Eclipse Exped., 2 males. MALES (FORM B) Philippines: Labuan Blanda J., Buton Strait, marginal coral, Dec. 13, 1909, Alb., 2 males. FEMALES Philippines: Subig Bay, China Sea, off southern Luzon, shore seine, sand, Jan. 7, 1908, Alb., 2 ovig. females; Sta. 5159, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°11’50”N., 119°54’E., Feb. 21, 1908, Alb., 1 ovig. female; Labuan Blanda I., Buton Strait, marginal coral, Dec. 13, 1909, Alb., 1 female; no. 27, Mariveles, Luzon, June 27, 1913, A. M. Reese, 3 ovig. females; no. 33, other data as last, 3 females, 1 ovig. female; near Mariveles, Luzon, 1913, A. M. Reese, 1 female, 3 ovig. females. Marshalls: Sta. VI, BikinilI., outer reef, Aug. 5, 1947, F. M. Bayer, 1 female. MEASUREMENTS.—Males form A, 11-26 mm.; males form B, 12, 13 mm.; females, 17-24 mm., ovig. females, 13-21 mm. Remarxs.—Gordon (1938, pp. 179-185, figs. 4 and 5) discussed Neptunus orbitosinus and particularly an unusual specimen with eight instead of nine anterolateral teeth and with a more robust male pleopod than in one of Rathbun’s cotypes. Stephenson and Campbell (1959, p. 144) supported Gordon’s suggestion that she was dealing with an undescribed species (Neptunus octodentata) and stated: ‘‘It is inconceivable that the variation in pleopod structure within P. orbitosinus could encompass the structures she figures (figs. 4c, 4c!).” Actually Gordon figured the pleopod of a further variant of a “male near to orbitosinus from Admiralty Is. (‘Challenger’ Colln.),” which was distinctly longer and thinner than any of the remainder. In the present collection, there is a very considerable variation in pleopod structure between males which are indistinguishable in 221-520-673 34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 general facies. Four main variants were noted and are figured (fig. 6). In the first, the pleopod is short and stout, little curved and naked on the center of its outer surface. In the second, the appendage is relatively thinner, more curved, and with hairs in the center of its outer surface. In the third, the appendage is even more curved but otherwise resembles the second form. In the fourth, the appendage is very long and slender, slightly curved, and resembles Gordon’s specimen from the “Challenger” collection. The first three variants show a certain amount of intergrading and are concluded to belong to the same morphological entity, whether a species, subspecies, or “form.” What Stephenson and Campbell considered “inconceivable variation” apparently occurs. On the other hand, there seems to be a distinct hiatus between these and the fourth variant. Following the pattern established for P. argentatus, the first three variants are designated form A and the fourth, form B. Crosnier (1962) figured a male pleopod of P. orbitosinus from Madagascar, greatly resembling figure 6 (c). Most of the present specimens differ from Crosnier’s in having a larger first anterolateral tooth well separated from the second tooth. All differ in having smaller ninth teeth pointing less directly outward. DistTRiBuTION.—Seychelles to Japan, and Australia. Portunus pelagicus (Linnaeus) Ficures 12c,d, 17b Cancer pelagicus Linnaeus, 1766, p. 1042. Neptunus pelagicus (Linnaeus).—Alcock, 1899, p. 34.—Sakai, 1939, pp. 387-388, pl. 49. Lupa pelagica (Linnaeus).—Barnard, 1950, pp. 152, 153-154, fig. 27. Portunus pelagicus (L.).—Stephenson and Campbell, 1959, pp. 96-98, figs. 2A, 3A, pls. 1 (fig. 1), 4A, 5A.—Sankarankutty, 1961a, pp. 103-104.—Crosnier, 1962, pp. 43-44, figs. 58, 61, 67.—Miyake, Sakai, and Nishikawa, 1962, p. 128 (record only). Dell, 1964a, pp. 303-304, 2 figs. Mareriau.—Malay Archipelago: Tachalorn, Siam, July 24, 1923, H. M. Smith, 1 male; Bangkok, Siam, July 24, 1923, H. M. Smith, 1 female; off Paknam, Gulf of Siam, Sept. 8, 1923, 1 female; Gulf of Siam, Lang Suen, Sept. 17, 1923, H. M. Smith, 1 female; Bandon Bight, Siam, September 1923, H. M. Smith, 1 male, 1 ovig. female; Chantabun River, at Lem Sing, Siam, May 7, 1927, H. M. Smith, 1 female; Singapore, Mar. 14, 1934, Herre, 1 female; Sta. 1004, purchased from Thailand, received Aug. 24, 1955, R. E. Elbel, 2 males, 1 female; Sta. 27, east coast of Gulf of Thailand, S. of Trat Bay off Lam Son Village, near Cambodian border, 11°57/00”N., 102°44’45”E., 10 m., shrimp trawl, Oct. 29, 1957, GVF, 1 male; Thailand, Songkhla, channel between Songkhala City and Goh Gnu I., Nov. 3, 1957, GVF, 1 female. | NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 35 China: U.S. Nav. Med. Res. Unit 2, Taipei, Formosa, purchased, received Feb. 13, 1958, Kuntz, 1 female. Philippines: Grande I., Subic Bay, Luzon, October 1907, Dr. J. C. Thompson, 1 ovig. female; Manila Bay, Dec. 9, 1907, Alb., 1 male; Manila Harbor, Dec. 30, 1907, Alb., 1 female; Sta. 5346, Cliff I., Malampaya Sound, Palawan Is., 10°50’30”N., 119°22/20”E., 7 f{m., Dec. 26, 1908, Alb., 1 female; Iloilo, Mar. 24, 1929, H. C. Kellers, 2 males; Iloilo, May 5, 1929, H. C. Kellers, 1 male, 1 female; Sandakan Harbor, British North Borneo, June 28, 1929, Herre, 1 male, 2 females; Manila Bay, Dec. 22, 1933, Herre collection, 1 male. Australia: Lake Macquarie, N.S.W., M. Filmer, Henry J. Brown, 1 male; Umba Kumba, south side of Little Lagoon, northeast end of Groote Eylandt, Gulf of Carpentaria, between Apr. 6 and May 18, 1948, R. R. Miller, 2 males, 1 female, 3 juvs.; Little Lagoon, northeast end of Groote Eylandt, Gulf of Carpentaria, 1948, R. R. Miller, 8 juvs. MEASUREMENTS.—Males, 26-129 mm.; females, 38-125 mm.; ovig. females, 108-118 mm., juvs. 13-40 mm. DistripuTion.—Kast Africa to Tahiti, including Japan, Philippines, Australia, and New Zealand. Portunus pubescens (Dana) Lupa pubescens Dana, 1852a, pp. 274-275, pl. 16 (fig. 9); 1852b, p. 84. Portunus pubescens (Dana).—Rathbun, 1906, p. 870, pl. 14 (fig 1) Edmondson, 1954, pp. 237-238, figs. 12, 18 —Stephenson and Campbell, 1959, pp. 99-101, figs. 2c, 3c, pls. 1 (fig. 3), 4C, 5C.—Stephenson, 1961a, p. 111. Neptunus pubescens (Dana).—Sakai, 1934, p. 303; 1939, pp. 388-389, pl. 80 (fig. 1). Martertau.—China: AT—-51, Namru-2-Taipei, Formosa, 12 mi. S. of Tau Hsui, seashore, Lighthouse Beach, 1957, R. E. Kuntz, 1 male (19 mm.). Philippines: Port San Pio, in small stream near mouth, 20 ft. seine, Noy. 11, 1908, Alb., 1 female (17 mm.). Remarxs.—Crosnier (1962) in redescribing P. convexus de Haan has drawn attention to its close similarity to P. pubescens; however, male P. converus are easily distinguished by their sinuous pleopods (Crosnier, 1962, figs. 64-66). DiIstTRIBUTION.—EHastern and Western Australia, Japan, and Hawaii. Portunus pulchricristatus (Gordon) FIGURE 7 Neptunus (Hellenus) spinipes Alcock, 1899, pp. 31-32, 39-40. Neptunus (Hellenus) pulchricristatus Gordon, 1931, p. 534, figs. 8, 10a. Not Neptunus (Amphitrite) spinipes Miers, 1886, p. 178, pl. 15 (fig. 1). Marerrau.—Philippines: Sta. 5097, Corregidor Lt., China Sea, off southern Luzon, 14°19’15”N., 120°33’52”E., 30 fm., Jan. 2, 1908, 36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Alb., 1 female; Sta. 5100, Corregidor Lt., China Sea, off southern Tuuzon, 14°17/15"N., 120°32’40"E., 35) fm’, Jan. 2, 1908, Albi male; Sta. 5104, China Sea, off southern Luzon, 14°45’48’N., 120°12’20”E., Jan. 8, 1908, Alb., 5 males, 1 female, 2 ovig. females, 2 juvs., 1 Sacculina infected female, 1 Sacculina infected unsexable specimen; Sta. 5105, Suesti Point Light, China Sea, 25 fm., 14°43’55”N., 120°12’50”E., Jan. 8, 1908, Alb., 1 male, 3 females, 1 juv., 1 unsexable Sacculina infected specimen; Sta. 5181, Antonia Is., eastern Panay, 11°36’40”N., 123°26’35”E., 26 fm., Mar. 27, 1908, Alb., 1 ovig. female; Sta. 5192, off North Cebu Is., Jilantaguan Is., 11°09/15”N., 123°50’E., 32 fm., Apr. 3, 1908, Alb., 1 male; Sta. 5207, Badian Is., off western Samar, 11°38’05”N., 124°40'45”E., 35 fm., green mud, sand, Apr. 14, 1908, Alb., 1 male, 1 female; Utara Point, Bongo I., 7°21’N., 124°07’E., May 22, 1908, Alb., 2 females; Sta. 5342, Endeavour Point (S.), 10°56’55”N., 119°17'24”E., 12-25 Ficure 7.—Portunus pulchricristatus (Gordon), male, Philippines, Sta. 5100, Alb.: w pleopod; b, pleopod tip, upper surface; c, abdomen. (Scales=a, 1 mm.; b, 0.5 mm.; ¢, 5 mm.) fm., Dec. 23, 1908, Alb., 5 males, 1 female, 1 ovig. female; Sta. 5442, west coast of Luzon, South Fernando Point Lt., 16°30’367N., 120°11’06”E., 45 fm., coarse sand, May 11, 1909, Alb., 16 males; Sta. 5448, South Miguel Point, east coast Luzon, 13°23’10’N., 123°45/19”E., 47 fm.; June 4, 1909, Alb., 3 males; Sta. 5644, Makassar I., Buton Strait, 5°27’24”S., 122°38’00’E., 22 fm.; Dec. 16, 1909, Alb., 1 male; Sta. 5646, North I., Buton Strait, 5°31/30"S., 122°22'40”E., 456 fm., green mud, Dec. 16, 1909, Alb., 1 male. China: All from China Sea, vicinity Hong Kong, Alb., 1908. Sta. 5302, 21°42’N., 114°50’E., 38 fm., Aug. 9, 4 males, 4 females, 1 ovig. female; Sta. 5303, 21°46’N., 114°47’E., 34 fm., black mud, Aug. 9, 5 males, 2 females; Sta. 5304, 21°46’N., 114°47’E., 34 fm., black mud, Aug. 9, 4 males, 3 females, 1 ovig. female; Sta. 5305, 21°54’N.., NO. 3556 PORTUNID CRABS—STEPHENSON AND REES an 114°46’E., 37 fm., Oct. 24, 2 males; Sta. 5308, 21°54’N., 115°42’E., 62 fm., Nov. 4, 1 male. MrasurEMENTS.—Males, 15-29 mm., females, 14-23 mm., ovig. females, 16-22 mm., juvs. 6-10 mm., Sacculina infected specimens, 14-18 mm. Remarxs.—In addition to the distinctions between this species and P. spinipes as listed by Gordon, it lacks the very fine tapering point on the male pleopod (compare figs. 7c and 14c). Distrinution.—Madras, Andamans, Gulf of Martaban, Arakan Coast, and Muscat. Portunus rubromarginatus (Lanchester) Achelous rubromarginatus Lanchester, 1900, pp. 746-747, pl. 46 (fig. 8). Neptunus (Amphitrite) rubromarginatus (Lanchester).—Shen, 1937, p. 104. Portunus rubromarginatus (Lanchester).—Stephenson and Campbell, 1959, pp. 112-113, figs. 2K, 3K, pls. 3 (fig. 3), 4K, 5K. MareriaAu.—Philippines: Jolo, ship’s side, electric light, 14 fm., Feb. 8, 1908, Alb., 1 female; Jolo anchorage, electric light, Mar. 5, 1908, Alb., 1 female; Sta. 5561, Tutu Bay (Jolo), coral and sand, Sept. 19, 1909, Alb., 3 males, 1 female; Labuan Blanda I., Buton Strait, marginal coral, Dec. 13, 1909, Alb., 1 female. MEASUREMENTS.—Males, 11-16 mm.; females, 16-29 mm. RemMarKs.—The anteroexternal angle of the merus of the third maxillipeds is produced laterally but not strongly so (see Stephenson and Campbell, 1959, pl. 5K). This leads to uncertainty in the key (above authors p. 90, couplet 16). The species is possibly closest to P. curvipenis Stephenson (1961a, pp. 106-108, figs. 1G, 3E; pl. 2, fig. 3; pls. 4E, 5B). DistripuTion.—Malay Archipelago, Hong Kong, South China Sea, and Australia. Portunus cf. rubromarginatus PLATE 3A MaTERIAL.—Japan: Sta. 5081, off Omai Saki, 35°14’N., 138°05’E., surface, Oct. 19, 1906, Alb., 1 incomplete female (13 mm.). Remarks.—Only the carapace and two chelae are present. The differences compared with typical P. rubromarginatus are more spiniform frontal teeth (pl. 34), more rounded granular eminences in the cardiac region of the carapace, more spiniform first eight anterolateral teeth, and longer ninth tooth. Portunus rugosus (A. Milne Edwards) Ficures 8-11; PLates 3p, 4, 5 Neptunus rugosus A. Milne Edwards, 1861, p. 335, pl. 33 (fig. 3).—Hess, 1865, p. 139.—A. Milne Edwards, 1873, pp. 156-157.—Miers, 1886, pp. 176-177 38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 (partim).—de Man, 1887, pp. 70-73 (under WN. andersoni).—Henderson, 1893, p. 869 (under N. andersoni). Portunus rugosus (A. Milne Edwards).—Guinot, 1957, p. 480 (under P. mariez).— Stephenson, 1961la, pp. 111-114, figs. 2B, 3G, pls. 3 (fig. 1), 4G, 5E. InrRopuUcTION.—Original sorting of specimens gave seven different forms resembling P. rugosus. Differences between them concerned shape of frontal teeth, length of last anterolateral tooth, acuteness or otherwise of posterior-posterolateral junction of the carapace, oc- currence of a paried or single metagastric granular elevation of the carapace, curvature and spinulation of the pleopod, and relative length of penultimate segment of male abdomen. Later it was shown that some of these features varied with size of specimens within a single collection (e.g., length of last anterolateral tooth), while others varied between adjacent collections (e.g., acuteness or roundedness of frontal teeth). ‘To separate most of the remaining POST - ORBITAL POST FRONTAL MED. GASTRIC PROTOGASTRIC MESOGASTRIC ANT. ANTEROLAT, POST. ANTEROLAT. EPIBRANCHIAL ANT, MESOBRANCHIAL METAGASTRIC CENT. MESOBRANCHIAL CARDIAC POST. MESOBRANCHIAL LAT. POSTCARDIAC POSTEROLATERAL MED. POSTCARDIAC Ficure 8.—Portunus rugosus (A. Milne Edwards): Diagram of elevated granular areas of carapace. forms involved reliance upon a single character (e.g., paired or single metagastric elevation), and it was concluded that two forms, here designated I and II, were present. These were compared with the Western Australian material upon which Stephenson’s (1961a) redescription of the species is based. This material belongs to form I. The features common to all variants are listed below, followed by accounts of the variation within forms I and II and of the distinctions between them. ‘These are followed by a brief report upon Milne Edwards’ material. Marertau.—Form I: Philippines: Jolo, ship’s side, Feb. 8, 1908, Alb., 1 female (dried); Jolo anchorage, electric light, Feb. 13, 1908, Alb., 1 male; Sta. 5156, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°12’50”N., 119°55’55”E., Feb. 21, 1908, Alb., 1 male; Sta. 5157, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°21’307N., 119°55’50”E., 18 fm., fine sand, Feb. 21, 1908, Alb., 5 males, 3 NO. 3556 PORTUNID CRABS—-STEPHENSON AND REES 39 females, 1 ovig. female, 1 juv; Sta. 5158, Tinakta I., Sulu Archipelago, Tawitawi, 5°12’N., 119°54’30”E., 12 fm., Feb. 21, 1908, Alb., 1 juv; Sta. 5159, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°11’50’N., 119°54’E., Feb. 21, 1908, Alb., 1 male; Sta. 5182, Antonia I., eastern Panay, 11°30’40’N., 123°23’20’E., Mar. 27, 1908, Alb., 1 juv. pean x1 d ane yeaaihas ~ Ficure 9.—Portunus rugosus (A. Milne Edwards). Male, abdomens, form I: a, Philippines, Sta. 5159, Alb.; b, W. Australian Mus. Reg. No. 47-60; form II: c¢, Philippines, Sta. 5159, Alb., 30mm. Fronts, form I: d, Philippines, Sta. 5158, 4/b., 8 mm. juv.; ¢, Philip- pines, Port Uson, Alb., 16 mm. ovig. female; f, Philippines, Sta. 5159, Alb.; g, W. Australian Mus. Reg. No. 51-60; form II: h, Philippines, Sta. 5169, Alb.; 1, Philippines, Sta. 5159. Alb., 30 mm. male. (Scale=approximately 5 mm., 2 lines indicate range.) male, 1 female, Cataingan Bay, Masbete, electric light, Apr. 19, 1908, Alb., 1 juv.; Busin Harbor, Burias 1., electric light, Apr. 22, 1908, Alb., 2 males, 1 juv. female; Looc, Lubang L., electric light, July 18, 1908, Alb., 1 female; Port Uson, W. of Pinas I., electric light, Dec. 17, 1908, Alb., 1 ovig. female; Endeavour Strait, electric light, Dec. 23, 1908, Alb., 2 juv. females. 40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Form II: Philippines: Sta. 5159, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°11’50”N., 119°54’E., Feb. 21, 1908, Alb., 2 males (ca. 21, 30 mm.), female (23 mm.), ovig. female (25 mm.); Sta. 5160, Tinakta JI., Sulu Archipelago, Tawitawi Group, 5°12’40’N., 119°55/10”E., 12 fm., sand, Feb. 22, 1908, Alb., 2 females (17, 18 mm.); Sta. 5169, Sibutu I., 4°32715”N., 119°22’45”E., 10 fm., Feb. 27, 1908, Alb., 2 females (both 16 mm.). MEASUREMENTS.—Form I: Males, 16-36 mm.; females, 15-38 mm.; ovig. females, 16, 32 mm., juvs. unsexed, 8-17 mm.; juv. male, 17 mm.; juv. females, 12-15 mm. FreaTuRES IN comMoNn.—Front: 3-toothed, middle tooth usually the smallest and least projecting. Cheliped: Robust but elongate. Posterior border of arm with single spine, anterior border with three spines, distal being widely separate from remainder. Ventrodistal extremity of arm bearing boss with small spine. Hand with upper surface bearing two granular carinae of which only inner ends in a subterminal spine; outer surface Ficure 10.—Portunus rugosus (A. Milne Edwards): Male abdomen, New Caledonian specimen from Muséum National d’Histoire Naturelle. (Scale=1 mm.) with two upper carinae clearly distinguishable and granular, but with lowermost merging into granulation of under surface; under surface with regular rows of granules giving a squamiform effect; inner surface with central row of carina of rounded granules (except in one juvenile). Anteroexternal angle of merus of third maxilliped: Produced forward. Ventral surface of thorax: Bearing conspicuous beaded granules. Dorsal surface of carapace: With hairy pile, through which project conspicuously elevated granular regions. These are more numerous than those figured by Stephenson and Campbell (1959, fig. 1b), and a modified terminology is adopted as shown in figure 8. Additional granulated areas are a pair of postfrontals and pair of post orbitals typically divided into three indistinct subareas. Gastric region with distinct median gastric patch with protogastrics and mesogastrics on each side, and metagastric behind. Protogastrics small, and typically NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 41 indistinctly separate from mesogastrics, which themselves typically bear ridgelike anterior terminations. Metagastric area sometimes bears two granular areas separated in midline, and sometimes a single continuous area with central ridge or tubercle. Two conspicuous anterolateral patches, anterior being the larger. Three pairs of mesobranchials, the anterior often subdivided into larger anteroexter- nal and smaller posteromedian patch. Central mesobranchial either oval or rectangular; posterior either short line of granules or small tubercle formed of fused granules. Cardiacs conspicuously elevated, usually triangular, and with ridgelike or tubercle-like anterior termina- tion. Lateral postcardiacs large and oval, median postcardiac small and with ridgelike or tuberculate anterior termination. Postero- laterals forming an are. Anterolateral teeth: Fourth and sixth the smallest, occasionally one or the other rudimentary or absent. Posterodistal border of merus of fifthlegs: Finely serrated. VARYING FEATURES.—Features that vary within a form are de- scribed below under each form respectively. FORM I Frontal teeth: Median tooth usually shorter than laterals, but sometimes projecting forward slightly farther. Lateral teeth varying from acute to bluntly rounded. Length of last anterolateral tooth: Varying within a collection depending on size of specimen, e.g., Sta. 5157 specimens less than 20 mm. carapace breadth with long tooth (equivalent in length to three or four preceding teeth), and specimens greater than 35 mm. breadth with short teeth (equivalent in length to one and a half preceding teeth). Also varying from collection to collection. Overall from lengths of one and a half to six preceding teeth. Posterior-posterolateral junction of carapace: Sometimes distinctly acute, sometimes slightly more acute than a right angle, typically obtusely angled, sometimes obtusely rounded. Acuteness varies within a collection (acuter teeth in smaller specimens) and also between collections. General granulated areas of carapace: Typically coarsely granular and conspicuously elevated. In some collections more finely granular and less conspicuous, in others more conspicuous and tending to form elevated tubercles. Male abdomen: Length of penultimate segment varying from slightly longer than broad to conspicuously so. Length of ultimate segment also varying. 42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Male pleopod: Varying in curvature with size of specimen being slightly curved in small and conspicuously curved in larger specimens. Subterminal bristles on inside varying from zero to nine. Subter- minal bristles on center of under surface varying from sparse to dense. ba} .* ————__., eel — —d. 5 5 ltl Renny ane ee * Ficure 11.—Portunus rugosus (A. Milne Edwards), male pleopods and pleopod tips, re- spectively. Form I: a and e¢, Philippines, Sta. 5159, Alb.; b and f, W- Australian Mus. Reg. No. 51-60; d, Philippines, Sta. 5157, Alb., largest male. Form II: ¢ and g, Philippines, Sta. 5159, 4/b. (Scales=a—d, 1 mm.; e-g, approximately 0.5 mm., 2 lines indicate range.) The most conspicuous variation is the presence of a single meta- gastric elevated area instead of two, as shown by the following: Endeavour Strait, 2 specimens; Looc, 1 specimen. One specimen (Port Uson, Dec. 17, 1908) possesses acute frontal teeth and conspicuously elevated central mesobranchial areas. Be- | NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 43 cause it is distinctly small for an ovigerous female (16 mm.), it is possible it should be separated from the remainder. FORM II Only eight specimens from three stations are available, but even so variation between them is less than might be expected from a study of form I. In smaller specimens the lateral frontal teeth are relatively shorter, less projecting, and less outwardly inclined. In two specimens (Sta. 5169) there is a single instead of a pair of meta- gastric granulated elevations. Unexpectedly constant features in form II comprise: a. Anterolateral teeth always long, equivalent in length to about five of the preceding teeth. b. Posterior-posterolateral junction of carapace always near to a right angle and moderately sharp. ec. General granulated areas of carapace relatively inconspicuous amongst the pile of hairs, not very coarsely granular, and without any tendency to be tuberculate. MitnE Epwarps’ MATERIAL.—Through the good offices of Mme. Daniéle Guinot, the following specimens were received from the Muséum National d’Historie Naturelle, Paris: 1 male (ca. 14 mm.), 1 ?female, labelled “Neptunus rugosus A. M. Edw. Auct. det. Coll. A. Milne Edwards 1903 (Marie) Nouvelle-Calédonie—Iles des Pins, 1873, p. 156. Ancienne collection séche. D. Guinot 1963.” Un- fortunately, when received they were so damaged that the central portions of the carapace could not be studied, and one specimen could not be measured at all accurately. Remaining details are: Frontal teeth: Acutely rounded, median almost as long as laterals. Similar to those of form I. Last anterolateral tooth: Length equivalent to a little more than three preceding teeth, sinuous, and directed slightly backward. Similar to those of form I in length and form II in shape. Carapace granulation: On undamaged areas, granulation much coarser than form IJ. Visible areas conspicuously elevated, as in form I. Male abdomen: Ultimate segment as broad as long with rounded tip. Penultimate segment broader than long (breadth 1.37 times length), with sinuous convex sides. (See fig. 10.) In general, these specimens are closest to form I, except for the shapes of both ultimate and penultimate segments of the male ab- domen, which are closer to form II. They bridge some of the gap between the two forms, and further collecting may result in a complete bridging. Meanwhile it is re- tained to emphasize the variability of the species. 44. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 DIFFERING FEATURES.—Features serving to distinguish between the two forms are tabulated as follows: lateral Soni last anterolateral granulated areas male abdomen male pleopod, tee teeth of carapace under surface Form I Directed Relatively Conspicu- Penulti- Row of forward straight ous, mate spinules or in- coarsely segment extending ward granular, either a long but some- slightly way back never times tu- or dis- from the outward; berculate tinctly tip tips longer acute or than bluntly broad rounded Form II Directed Slightly Relatively Penulti- Row of outward; curved or incon- mate spinules tips rec- sinuous; spicuous segment only ex- tangularly slightly and less as broad tending a rounded back- coarsely as long short dis- wardly granular, tance directed not tu- back from berculate the tip These features are not thought sufficiently weighty to merit specific separation for the following reasons: a. One of the smaller specimens of form II (Sta. 5169) (fig. 9h) has frontal teeth intermediate between those typical of this form (see fig. 97) and a specimen of form I from Western Australia (see fig. 9g). b. The proportions of the penultimate segment of the male abdomen vary in form J, and little extension of the range would encompass form IT. c. There are insufficient mature males of form II (only two) to establish that minor differences between the very similar pleopods in the two forms are not due to variability. d. Milne Edwards’ original description included some features of form I and some of form II, and seems to bridge most of the gap between them. Partial examination of his damaged specimens tends to confirm this bridging. The features of form I, including one or another of its variants, are: ‘‘Carapace trés-fortement bosselée” and ‘“‘Aneles postérieurs de la carapace aigus et spiniformes.” The fea- tures of form II are first, ““Cornes laterales trés-grandes, trés minces et légérement dirigées en arriére,’’ and second, obtuse lateral frontal teeth. It should be noted that Milne Edwards’ figures (pl. 33, figs. 3, 3a) show differences from both forms as follows: a. The posterior anterolateral granulated patch is shown as almost the same size as the anterior. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 45 b. The postorbital patch is shown more discretely than in any of the present specimens. Portunus sanguinolentus (Herbst) Figures 12a, 6 Cancer sanguinolentus Herbst, 1783, p. 161, pl. 8 (figs. 56, 57). Portunus sanguinolentus (Herbst).—Rathbun, 1906, p. 870.—Edmondson, 1954, pp. 236-237, figs. 12, 13—Stephenson and Campbell, 1959, pp. 98-99, figs. 2B, 3B, pls. 1 (fig. 2), 4B, 5B.—Sankarankutty, 1961a, p. 103.—Crosnier, 1962, pp. 45-47, figs. 59, 62-63, 68.—Miyake, Sakai, and Nishikawa, 1962, p. 128 (record only). Neptunus sanguinolentus (Herbst).—Alcock, 1899, pp. 32-33.—Sakai, 1939, p. 387, pl. 48 (fig. 1). Lupa sanguinolentus (Herbst).—Barnard, 1950, pp. 152, 154-155. Matertau.—Malay Peninsula: Gulf of Siam, beach, North Singora, Oct. 5, 1923, H. M. Smith, 2 females; Benkoelen, Sumatra, November Ficure 12.—Fronts. Portunus sanguinolentus (Herbst): a, Japan, Sta. 4920, Alb., 19 mm. male; b, Malay Peninsula, Sta. 53, G.V.F., 35 mm. male. P. pelagicus (Linnaeus): c, Australia, Umba Kumba, Miller, 22 mm. male; d, same as c, 30 mm. male. (Scale=5 mm.) 1925, Kellers, 1 male; Sriracha, southeast Siam, Apr. 20, 1934, H. M. Smith, 1 juv; Thailand, Songkhla Channel between Songkhla City and Goh Gnu I., GVF Sta. 53, Nov. 3, 1957, 1 male; Thailand, Songkhla Channel next to city, Songkhla Province, purchased in fish market, 07°12/07’N., 100°37'15”E., GVF Sta. 44, Nov. 4, 1958, 5 males, 5 females, 1 juv. (damaged); Thailand, near rock jetty, S. of Chol Buri, between Bangsaen and Chol Buri, GVF Sta. 84, Dec. 1, 1957, R. Rofen, 1 female. China: Kowloon, electric light, Sept. 14, 1908, Alb., 1 juv.; AT-51, Namru-2-Taipei, Formosa, 12 mi. S. of Tau Hsui, seashore, Lighthouse Beach, 1957, R. E. Kuntz, 4 males, (2 damaged), 2 females, 2 juvs. Philippines: Manila Bay, Dec. 7, 1907, Alb., 1 female; Manila Bay, Dee. 8, 1907, Alb., 1 female; San Vicente Port, Luzon Beach, Nov. 15, 1908, Alb., 1 female; Linapacan I., Malcochin Harbor, electric light, Dec. 18, 1908, Alb., 1 male; Nakochin Harbor, Linapacan I., seine, Dec. 19, 1908, Alb., 1 female; near Sta. 5344, Malampaya, Palawan, 46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 130 ft. seine, Dec. 26, 1908, Alb., 1 male (dried); San Miguel Bay, Colssi Point, east coast of Luzon, June 14, 1909, Alb., 1 juv; Sta. 5539, Apo I., between Negros and Siquijor, 9°03’20’N., 123°24’45”K., Aug. 19, 1909, Alb., 1 juv; Philippine Is., E. A. Mearns, 2 males, 1 female; near Mariveles, Luzon, 1913, A. M. Reese, 1 female; Manila, Feb. 25, 1929, Kellers, 1 male; Iloilo, Apr. 20, 1929, H. C. Kellers, 2 males, 41 juvs., 2 damaged specimens; Iloilo, May 9, 1929, H. C. Kellers, 2 females; no. 363, Pandan, Antique, from trap at mouth of tidal stream, Apr. 7, 1946, D. G. Frey, 1 female. Australia: M48-28, Port Bradshaw near Cape Arnhem, July 25, 1948, R. R. Miller, 1 female. Marianas: 112-x-65, Ajayan River, Guam, June 19, 1945, D. H. Johnson, 1 female; Agfayan Bay, Guam, 145/R.H.B. 150, July 7, 1945, R. H. Baker, 1 male. Japan: Sta. 4920, Kusakaki Jima, 30°34’N., 129°22’K., surface, Aug. 13, 1906, Alb., 1 juv; Sta. 5081 off Omai Saki, 34°14’ N., 188°05’ E., surface, Oct. 19, 1906, Alb., 6 juvs. MEASUREMENTS.—Males, 21-43 mm.; females, 20-121 mm.; juvs., 9-20 mm. Remarks.—In many specimens the red marks on the carapace are indistinct or unrecognizable because they have faded after prolonged preservation. In such cases the species can be recognized by frontal teeth (figs. 12a, b) that are more equal in size and more evenly spaced than in P. pelagicus and by the merus of the third maxilliped, which is hairy as opposed to the smoothness in P. pelagicus. Possibly P. madagascariensis (Hoffmann) is a synonym and based upon a faded specimen (see Crosnier, 1962, p. 47).? DistriputTion.—Kast Africa to Hawaii, including Japan and Australia and also from the Adriatic. Portunus spiniferus, new species FicurRE 13; PuatTE 6a Marteriau.—Philippines: Varadero Bay, Mindanao, 10:30-11:30, electric light, July 22, 1908, Alb., 1 male (23 mm., holotype, cat. no. 112094); Bolinao Bay, west coast of Luzon (on label, ‘“‘Bohiao’’), electric light, May 9, 1909, Alb., 1 female (24 mm.); Batananan L., Dyn., June 13, 1909, Alb., 1 female (28 mm.). Sta. 408, Dinagat, Surigao, light plankton, May 22, 1946, D. G. Frey, 1 male (30 mm.). Description.—Front: 3-lobed, somewhat protruding. All lobes roundedly triangular, median about one-half size of laterals. Two conspicuous notches in upper border of orbit, lower border with stout, rounded tooth. 2 Later studies indicate it belongs to Callinectes. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 47 Anterolateral teeth: Nine, first stout, fourth and sixth quite small, eighth larger than ninth, ninth the longest, stoutest, and very projecting. Carapace: Much broader than long (ca. 2.2 times), with posterior- posterolateral junction forming an acute upturned spine, covered throughout by very fine, dense pubescence, through which small conspicuously raised granular areas are evident, several of these being spiniform. Beneath hairs, most of carapace smooth and shining. (Granulated areas showing through pile correspond with Q g N Ficure 13.—Portunus spiniferus, new species, male holotype: a, pleopod; b, pleopod tip, upper surface; c, abdomen. (Scales=a, 1 mm.; b, 0.1 mm.;c, 5 mm.) those in Portunus rugosus complex and the terminology of figure 8 is employed.) Postfrontals relatively inconspicuous; median gastric elongate; protogastrics small and inconspicuous; mesogastrics termi- nated anteriorly by tubercle-like elevation; metagastrics each a small tubercle; postorbitals each composed of four reasonably distinct patches, the two posterior patches with ridgelike anterior termina- tions; anterolaterals large and conspicuously elevated; epibranchials conspicuous; anterior mesobranchial two discrete patches, both elevated, one much smaller than the other; central mesobranchial, AS PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 linear patch; posterior mesobranchial, short line or patch; cardiacs conspicuously elevated with tuberculate or spiniform tips; lateral postcardiacs resolvable into three or four tubercle-like clusters of eranules; median postcardiac conspicuously elevated ridge or spine; posterolaterals, forming curved arc, terminated anteriorly with forwardly directed spinelike tubercle. In one case two spines on posterolaterals, also two on lateral postcardiacs, and one on posterior mesobranchial. In another case approximately four spines on pos- terolaterals and one on posterior mesobranchial. Chelipeds: Right larger. With well-developed spines, covered by fine pile overlying squamiform markings. Posterior border of arm bearing two sharp spines, anterior border with three unusually large sharp spines, middle the largest. Wrist with large inner and outer spines. Upper surface of hand with two spines (excluding that at wrist articulation), outer near the finger articulation, inner a little distance back. Spines better developed on smaller left chela. ‘Two granular carinae on upper surface of hand, two less developed ones on outer surface, and one barely recognizable on inner surface. Spaces between carinae, also under surface, with conspicuous squamiform markings. Fifth leg: Merus very short (length almost exactly equalling breadth) with finely denticulate posterior border. Third maxilliped: Anterior portion of merus produced strongly forward but not laterally. Male abdomen: Penultimate segment elongate and with concave borders, ultimate segment, cordiform, slightly more than half length of penultimate. Male first pleopod: Short, stout, regularly curving to pointed tip. ReEMARKS.—This species keys out in Stephenson and Campbell (1959) with P. tweediet, P. tenuipes, P. mariei, and P. alcocki but differs from all in the elevated spiniform areas on the posterior por- tion of the carapace. It is clearly separable from P. rugosus by two spines on the posterior border of the arm. Portunus spinipes (Miers) Figure 14 Neptunus (Amphitrite) spinipes Miers, 1886, pp. 178-179, pl. 15 (figs. la—c). Neptunus (Hellenus) spinipes Miers—Gordon, 1931, p. 534 (under N. pulchri- christatus), figs. 9, 10a. Not Neptunus (Hellenus) spinipes Alcock, 1899, pp. 31-32, 39-40 (=P. pulchri- cristatus (Gordon)). Marertau.—Philippines: Sta. 5164, Observation I., Sulu Archi- pelago, Tawitawi Group, 18 fm., green mud, Feb. 24, 1908, Alb., 2 NO. 3556 PORTUNID CRABS—-STEPHENSON AND REES 49 males (wet), 3 males, 1 female (dried); Sta. 5181, Antonia I., eastern Panay, 11°36’40’N., 123°26’35”E., 26 fm., Mar. 27, 1908, Alb., 1 Sacculina infected male; Sta. 5642, Buton Strait, 4°31’40”S., 122°49’42”K., 37 fm., grey mud, Dec. 14, 1909, Alb., 5 males, 6 females; Sta. 5644, Makassar I., Buton Strait, 5°27’24”S., 122°38’00’E., 22 fm., Dec. 16, 1909, Alb., 1 male (damaged). MEASUREMENTS.—Males, 16-26 mm.; females, 18-24 mm.; Sacculina infected male, 15 mm. Remarxks.—Gordon has shown that Alcock’s Neptunus spinipes differs from Miers’ and has listed the distinctions as well as giving figures of the carapaces and male pleopods of the two species. Her figure of the carapace of one of the cotypes differs noticeably from Figure 14.—Portunus spinipes (Miers), male, Philippines, Sta. 5642, Alb.: a, pleopod; b, pleopod tip, upper surface; c, abdomen. (Scales=a, 1 mm.; b, 0.1 mm.;c, 5 mm.) Miers’ figure in having a much less prominent front. The present specimens resemble her figure. The male pleopods are refigured. The species is incorrectly placed in the key of Stephenson and Campbell (1959) because the posterior- posterolateral junction of the carapace is obtuse. Distrisution.—Apparently only known previously from Miers’ material, from the Philippines, and close to Albatross Sta. 5181, waz 11°37'00" N.,.123°31/007E. Portunus tenuipes (de Haan) Figure 15 Amphitrite tenuipes de Haan, 1835, p. 39, pl. 1 (fig. 4). Neptunus tenuipes (de Haan).—A. Milne Edwards, 1861, pp. 335, 339. 221-520—67—4. 510) PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Neptunus (Hellenus) tenuipes (de Haan).—Alcock, 1899, p. 42.—Shen, 1937, p. 104, figs. 4, 8. Neptunus (Amphitrite) tenuipes (de Haan).—Sakai, 1939, pp. 389-390, pl. 80 fig. 2). Sahin ae (de Haan).—Guinot, 1957, p. 480 (under P. mariei).—Stephen- son and Campbell, 1959, pp. 103-104, figs. 2H, 3E, pls. 2 (fig. 1), 4H, 5E. MareriaL.—Philippines: Subig Bay, shore, seine, Jan. 7, 1908, Alb., 4 males (1 Sacculina infected), 2 females, 3 ovig. females; Port Binanga, ship’s side, Jan. 8, 1908, Alb., 2 females; Tachloban Anchor- age, about ship, hand dredge, 3 fm., Apr. 12, 1908, Alb., 1 ovig. female; Busin Harbor, Burias Is., electric light, Apr. 22, 1908, Alb., 2 males; Surigao, Mindanao, 150 ft. seine, May 8, 1908, Alb., 1 male, 1 female; Davao, Mindanao, 150 ft. seine, May 16, 1908, Alb., 1 male, 1 female; near Sta. 5459, Cabugao, east coast of Luzon, electric light, June 9, 1909, Alb., 1 male, 1 female; no. 27 Mariveles, Luzon, June 27, 1913, A. M. Reese, 1 male; no. 33 Mariveles, Luzon, June 28, 1913, A. M. Reese, 5 males, 1 ovig. female; near Mariveles, Luzon, 1913, A. M. Ra i Ficure 15.—Portunus tenuipes (de Haan), fronts: a, Philippines, Mariveles, No. 33, Reese, 32 mm. male; b, Philippines, Surigao, 4/b., 36 mm. male; c, same as a, 22 mm. male; d, Philippines, Pt. Binanga, Alb., 16 mm. female. (Scale=5 mm.) Reese, 1 male; Waigin, tide pool, June 8, 1929, Herre collection, 1 male. MEASUREMENTS.—Males, 14-36 mm.; females, 16-35 mm.; ovig. females, 22-25 mm. Remarks.—As specimens become larger, the frontal lobes become more acute and prominent (see fig. 15). Distribution—Andamans to Japan and the Philippines, including Australia. Portunus trituberculatus (Miers) Figures 16, 17a Portunus pelagicus de Haan (not Linnaeus) 1835, p. 37, pls. 9, 10. Neptunus trituberculatus Miers 1876b, p. 222; 1886, p. 172.—Sakai, 1934, p. 303; 1936, p. 129, pl. 37. Portunus trituberculatus (Miers).—Rathbun 1902, p. 26.—Stephenson and Camp- bell, 1959, p. 90 (in key).—Miyake, Sakai, and Nishikawa, 1962, p. 128 (rec- ord only). Neptunus (Neptunus) trituberculatus (Miers).—Sakai, 1939, p. 388, pl. 50. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES ol Materiau:—Kast Africa: Ras Banas, Red Sea, 1950, Sozon Vatikiotis, 1 male, 3 females.* Malay Peninsula: S.1166, Singapore, “China,” Arthur de Sowerby, 1 male. | Figure 16.—Portunus trituberculatus (Miers), male: a, pleopod, Malay Peninsula, Singa- pore, de Sowerby; b, pleopod tip, upper surface, same specimen; c, abdomen, E. Africa, Ras Banas, Vatikiotis. (Scale=a, 10 mm.; b, 1 mm.; c,5 mm. Identity of specimen now doubtful.) a Ficure 17.—Fronts. Portunus trituberculatus (Miers): a, East Africa, Ras Banas, Vati- kiotis, 56 mm. female (identity of specimen now doubtful). P. pelagicus (Linnaeus): b, Philippines, Sta. 5346, Alb., 66 mm. female. (Scale=5 mm.) 3 Doubts have arisen concerning the identity of these specimens, which may be reported on in a later publication. o2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 China: S$.1165, China, 1925-1927, Arthur de Sowerby, 1 female. (Probably like the last specimen, this came from Singapore.) MEASUREMENTS.—Males, 45-180 mm.; females, 45-90 mm. Remarks.—This species is very close to P. pelagicus, and the differ- ence in the fronts is shown better by figures than by words (fig. 17). The male pleopods are also very similar although the overall curvature in P. trituberculatus departs farther from the circular than with P. pelagicus (see Stephenson and Campbell, 1959, fig. 2A). Additional material will be required to determine whether the details of terminal armature do differ slightly in the two species. DistriputTion.—China and Japan (Sakai, 1939). Figure 18.—Poriunus tuberculosus (A. Milne Edwards), male, Philippines, Sta. 5159, Alb.: a, pleopod; b, pleopod tip, upper surface; c, abdomen. (Scales=a, 1 mm.; b, 0.5 mm.; c, 5 mm.) Portunus tuberculosus (A. Milne Edwards) Figures 18; PLATE 6B Neptunus tuberculosus A. Milne Edwards, 1861, pp. 333-334, pl. 31 (fig. 5). Neptunus (Amphitrite) tuberculosus A. Milne Edwards.—Miers, 1886, p. 176. Neptunus (Hellenus) tuberculosus A. Milne Edwards.—Alcock, 1897, pp. 42-43. Portunus tuberculosus (A. Milne Edwards).—Stephenson and Campbell, 1959, p. 89 (in key).—Crosnier, 1962, pp. 69-71, figs. 97-98, 124-127. MareriAu.—China: Sta. 5309, China Sea, vicinity Hong Kong, 21°53/N., 115°51’E., 62 fm., green mud, Nov. 4, 1908, Alb., 1 female. Philippines: Sta. 5104, China Sea off southern Luzon, 14°45'48’N., 120°12’20” E., Jan. 8. 1908, Alb., 1 male, 1 female, 1 ovig. female; NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 53 Sta. 5159, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°11’50” N., 119°54’E ., Feb. 21, 1908, Alb., 1 male; Varadero Bay, Mindanao, 10:30-11:30, electric light, July 22, 1908, Alb., 1 female; Sta. 5442, west coast of Luzon, South Fernando Point Light, 16°30’36’N.., 120°11’06”E., 45 fm., coarse sand, May 11, 1909, Alb., 7 males, 1 female, 2 ovig. females; Sta. 5478, Tacbuc Point, (Leyte), 10°46’24” N., 125°16’30’E., July 29, 1909, Alb., 1 female; Sta. 5642, Buton Strait, 4°31’40"S., 122°49’42”K., 37 fm., grey mud, Dec. 14, 1909, Alb., 1 male. Palau: Sta. 125-1187 Palau I., about 134 mi. NE. of Ngabadongu, 7°17'36"N., 134°21/42”E., 17 fm., Aug. 24, 1955, GVF., 1 female. MeAsurREMENTS.—Males, 18-49 mm.; females 15-30 mm.; ovig. females, 21-26 mm. Remarxs.—Originally it was suspected that two different species were present, some being more strongly embossed and_ possessing spiniform tubercles on the cardiac and lateral postcardiac regions of the carapace. Later intergrades were found. The following, the only adequate description in English, amplifies Crosnier’s (1962) French description. Description.—Front: Projecting, four round, approximately equal lobes, medians the more prominent. Anterolateral teeth: Irregularly sized and spaced, first and second almost confluent, fourth and sixth small and depressed, third and fifth with enlarged granular bases, ninth the longest. Carapace: Moderately broad (B/L.=1.6-1.7). Regions varying from moderately to very strongly embossed, and carrying beaded granules or occasionally tubercles. Strongly depressed posterior to protogastric and mesogastric areas. Main granular patches as fol- lows: Protogastrics subdivided into two; mesogastrics, typically two oblique patches; metagastrics, transverse ridge with anterior prolonga- tion; cardiacs either an elevated patch, or pair of patches, or pair of elevated granular tubercles; lateral postcardiacs elevated and some- times bearing tubercles; median postcardiac present; anterolaterals diffuse; epibranchials with marked change in curvature near middle and interrupted on cervical groove to form discrete patch; three meso- branchials; diffuse posterolateral. Chelipeds: Elongate, right slightly the stouter. Upper surfaces cov- ered with beaded granules and lower surfaces, particularly of hands, bearing squamiform markings. Anterior border of arm bearing two spines typically followed by a tubercle, occasionally by a third spine. Posterodistal border with single spine. (Crosnier describes two teeth, the second ‘“méritant 4 peine le nom de dent.) Wrist normal. Upper surface of hand bearing three granular carinae of which inner- most terminates in distal spine or spiniform tubercle. Conspicuous 54 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 carinae on outer surface of hand, but inner carina represented only by double row of enlarged granules. Fifth leg: Merus short (L/B=1.2). Posterior border composed of rounded granules or spinules. Third maxilliped: Anterior border of merus produced forward and a little laterally. Male abdomen: Penultimate segment about 1.3 times as long as broad and with gently curving sides. Ultimate segment about 2.5 times as long as broad. Male first pleopod: Sharply curved near middle and slightly curved behind tip which is blunt, and bears only microscopic spinules (see fig. 180). The present material differs from Crosnier’s (1962) figure (fig. 124) in having blunter first anterolateral teeth, epibranchial ridge of cara- pace often ending in a swollen granulated patch, and posterolateral angle of the carapace generally much sharper. In the fragmented specimen from Sta. 5642, Dec. 14, 1909, Alb., the left anterior branchial region is swollen as if from a parasite. The species is clearly separable from P. macrophthalmus Rathbun, P. emarginatus Stephenson, and P. tenuicaudatus Stephenson by dif- ferences in the detailed ornamentation of the carapace. Distrinution.—Madagascar, Persian Gulf, Ceylon, Hawaii. Portunus tweediei (Shen) Fiaure 19 Neptunus (Hellenus) tweediet Shen, 1937, p. 109, fig. 6. Portunus tweediei (Shen).—Stephenson and Campbell, 1959, p. 90 (in key). Marerrau.—Malay Peninsula: Thailand, entrance to Trat Bay, Gulf of Thailand, flat mud bottom, 5 m., 11°58’30”N., 102°44’05”E., GVF, Sta. 26, Oct. 29, 1957, 1 fragmented female. Philippines: Manila Bay, off Eremita, June 5, 1907, Alb., 1 male, 2 ovig. females; Manila Bay, Dec. 7, 1907, Alb., 1 male; Manila Bay, outside of breakwater, mud, small rocks, Dec. 12, 1907, Alb., 41 males, 23 females (3 ovig.), 1 juv; Little Harbor at Lunela, Manila Harbor, Dec. 12, 1907, Alb., 1 male, 1 female; Tacloban Anchorage, about ship, hand dredge, Apr. 12, 1908, Alb., 1 male; Sta. 5346, Cliff I., Malampaya Sound, Palawan Is., seine, gravel, rocks, Dec. 29, 1908, Alb., 1 ovig. female. MEASUREMENTS.—Males, 8-21 mm.; females, 7-16 mm.; ovig. females, 15-25 mm.; juv., 10 mm. Remarks.—The posterior-posterolateral junctions of the carapace are possibly more spinous than indicated by Shen; frontal teeth are a little more rounded; the postcardiac granular patch is lozenge NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 00 shaped instead of triangular; and the mesobranchial granular patches are arranged slightly differently. The male abdomen is longer and thinner (particularly so in the smaller specimens) with penultimate segment tapering from halfway along its length, and ultimate almost twice as long as broad and with a rounded tip. The male pleopod is more robust centrally and more pointed distally than figured by Shen. However, details of terminal armature coincide with Shen’s figure 8d. The present specimens, which include some obviously mature, are all much smaller than Shen’s. Ficure 19.—Poriunus tweedei (Shen), male, Philippines, Tacloban, A/b.: a, pleopod; b; pleopod tip, upper surface; c, abdomen. (Scales=a, 1 mm.; }, 0.1 mm.;c, 5 mm.) DistripuTion.—Previously known only from Singapore (Shen, 1937). Genus Scylla de Haan, 1833 Scylla serrata (Forskal) Cancer serratus Forsk&l, 1755, p. 90. Scylla serrata (Forsk&l).—Stephenson and Campbell, 1960, pp. 111-115; fig. 2N, pls. 4 (fig. 4), 5N, 6C.—Crosnier, 1962, pp. 72-73, figs. 128, 129.— Miyake, Sakai, and Nishakawa, 1962, p. 128 (record only).— Dell, 1964b, pp. 59-62. 56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 * Marertau.—EHast Africa: Strait at Changamive, 3% mi. from Mombasa, Nov. 21-30, 1909, E. A. Mearns, 2 males. Malay Peninsula: Menam at Paknam, Siam, July 5, 1923, H. M. Smith, 1 juv; Chao Phya River, Paknam, Siam, June 3, 1927, H. M. Smith, 1 male, 1 juv; Bangpa Kong River, Siam, June 4, 1928, H. M. Smith, 3 males, 1 female; Sta. 10002, from Thailand, purchased, received Aug. 24, 1955, R. EH. Elbel, 1 male. China: U.S. Nav. Med. Res. Unit 2, Taipei, Formosa, purchased, received Feb. 13, 1958, R. E. Kuntz, 1 male. Philippines: Port San Pio Quinto, China Sea, vicinity of Batanes, in small stream near mouth, 20 ft. seme, Nov. 11, 1908, Alb., 1 male. Australia: U42-728 Ironstone Reef about 3 mi. W. of Yirrkala, Aug. 27, 1948, R. R. Miller, 1 ovig. female; Little Lagoon and vicinity northeastern end Groote Eylandt, Gulf of Carpentaria, 1948, R. R. Miller, 2 females. Japan: OPM958, Hentonia, Okinawa, Sept. 19, 1945, O. A. Muen- nink, 1 female. MrASuUREMENTS.—Males, 29-88 mm.; females, 26-123 mm.; ovig. female, 170 mm.; juvs., 26-28 mm. DistriputTion.—Hast Africa to Tahiti. Recorded from New Zealand by Miers (1876), with the record repeated by later workers including Chhapgar (1957) and Stephenson (1961b). Hutton (1882) suspected this record, but it has been confirmed by Dell (1964b). Genus Thalamita Latreille, 1829 See Stephenson and Hudson, 1957, p. 361, for synonymy. Thalamita admete (Herbst) Fiaure 20 Cancer admete Herbst, 1803, pp. 40-41, pl. 57 (fig. 1). Thalamita admete (Herbst).—Latreille, 1829, p. 33 (fide Miers, 1886).—A. Milne Edwards, 1861, pp. 356-357.—Heller, 1865, p. 28.—Miers, 1884, p. 230.—Ortmann, 1893, p. 83.—Borradaile, 1900, p. 579.—Calman, 1900, p. 23.—Lenz, 1905, p. 362.—Rathbun, 1907, p. 63.—Nobili, 1907, p. 383.— Rathbun, 1911, p. 208—Edmondson, 1923, p. 1550; 1925, p. 37; 1954, pp. 255-256, figs. 30a,b, 3la-e.—Sakai, 1939, pp. 414, 421-422, pl. 85 (fig. 1).—Ward, 1942, p. 80.—Barnard, 1950, p. 176, fig. 33¢.—Holthius, 1953, p. 7.—Stephenson and Hudson, 1957, pp. 320, 324-326, figs. 2I, 31, pls. 1 (fig. 1), 7A, 10A.—Stephenson, 1961a, p. 117.—Forest and Guinot, 1961, p. 30, figs. 19a,b.—Sankarankutty, 1961a, p. 106; 1961b, p. 122.—Crosnier, 1962, pp. 96-97, figs. 154, 157, 162-164, 168. Thalamita savignyi A. Milne Edwards, 1861, pp. 357-358, 367.—Ortmann, 1894, p. 46. Thalamita admeta (Herbst).—Alcock, 1899, pp. 82-84.—Tweedie, 1950a, p. 84, fig. 2b. Thalamita admete var. edwardsi Borradaile, 1900, p. 579.—Nobili, 1907, p. 383. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 57 Thalamita admeta var. A admeta Borradaile, 1902, pp. 202-203. Thalamita admeta var. B edwardsi Borradaile, 1902, pp. 202-203. Thalamita admeta var. C savignyi Borradaile, 1902, pp. 202-203. Thalamita edwardsit Borradaile-——Rathbun, 1906, p. 873.—Edmondson, 1925, p. 37; 1954, pp. 254-255, figs. 28b, 29a-d.—Forest and Guinot, 1961, p. 32, figs. 20a, b.—Crosnier, 1962, p. 98, fig. 158.—Guinot, 1962, pp. 2-3. Thalamita admeta var. savignyi Borradaile.—Nobili, 1906a, pp. 202, 206-208. Thalamita dispar Rathbun, 1914, p. 657, pl. 1, fig. 4. Thalamita admete var. savignyi Borradaile-—Laurie, 1915, pp. 440-441. Thalamita ?edwardsi Borradaile.—Holthius, 1953, p. 8. ?Thalamita spiceri Edmondson, 1954, pp. 258-260, fig. 33a-e. Mareriau: China: Nau Wan, Formosa, China Sea, Jan. 27, 1910, Alb., 1 male. Philippines: Subig Bay, China Sea off southern Luzon, shore, seine, sand, Jan. 7, 1908, Alb., 1 female; Tataan, Simaluc, shore, Feb. 19, 1908, 1 female; Guijulugan, Negros, station shore, sand, eravel, mud, Apr. 2, 1908, Alb., 1 ovig. female; Makinog, Camiguin I., between Leyte and Mindanao, tide pool, Aug. 3, 1909, Alb., 5 males, 1 female, 4 ovig. females; Sta. 5561, Tutu Bay (Jolo), coral and sand, Sept. 19, 1909, Alb., 1 female; Great Tobea I., tide pool, sand, coral, Dec. 15, 1909, Alb., 3 males, 2 females, 1 ovig. female; Makasser I., Buton Strait, 5°27'24”S., 122°38’00”E., tide pools, Dec. 16, 1909, Alb., 1 male, 1 female; Philippine Is., E. A. Mearns, 1 male. Australia: Reef at Yirrkala, July 12, 1948, R. R. Miller, 1 male; M.48-21, ironstone reefs (not coral) and rock pools at Yirrkala, NW. of Cape Arnhem, Aug. 6, 1948, R. R. Miller, 2 males; C-7 Yurkala, NW. of Cape Arnhem, coral reef, Aug. 25, 1948, R. R. Miller, 1 ovig. female; Little Lagoon and vicinity, northeastern end of Groote Eylandt, Gulf of Carpentaria, 1948, R. R. Miller, 1 male. Melanesia: Malaluva reef, Fiji, June 19, 1922, State University of Iowa, 1 male, 1 ovig, female; Ovalau I., Fiji, Mar. 17, 1929, Herre, 1 male, 1 female; New Georgia, Jan. 10, 1945, W. A. Bartos, 4 males, 3 females, 1 ovig. female. Marianas: No. 83/X-48, Piti Bay, Guam, June 30, 1945, McElroy and Baker, 1 ovig. female; 145/RHB-150, Agfayan Bay, Guam, July 17, 1945, R. H. Baker, 1 male; Saipan, in coral heads, 1945, A. H. Banner, 2 males, 1 female, 1 juv; Y-128G, from under rocks, tidal flats, Tomil Harbor, 1952, Hiatt, 1 male; Y-193 B, C, from under rocks, tidal flats of Balabat, Yap Is., 1952, Hiatt, 3 males, 1 female; 177—G-—1, Ifaluk, Caroline Is., 1953, F. M. Bayer, 9 juvs. Japan: Tanego Shima, 1906, Alb., 1 female. Marshalls: Eniwetok, reef, low tide, February 1940, Ziesenhenne, 2 males; no. 4331, Sta. 6, one-third mi. W. of Southeast Point, Bikini I., Apr. 23, 1946, J. P. E. Morrison, 1 female; no. 4351, Sta. 4, one- third mi. SW. of southeastern point, Bikini I., 3% fm., forams, 58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 sand, Apr. 23, 1946, 1 female; no. 49, Bikini I., outer reef flats, July 22, 1947, F. M. Bayer, 1 male. Hawaii: Black Point, Haunama Bay, Oahu, 1937, L. R. Woodward, Ficure 20.—Thalamita admete (Herbst), male pleopod tips, upper surface: a, Hawaii, Oahu, Army Med. Mus.; b, Australia, Yirrkala, M48-21, Miller, 21 mm.; ¢, Samoa, Apia, June 27, 1902, Alb., 21 mm.; d, Philippines, Makasser I., 4/b.; e, Samoa, Apia, June 27, | 1902, Alb., 23 mm.; f, Samoa, Apia, June 1, 1902, Alb.; g, China, Nau Wau, Alb.; h, Samoa, Pago Pago, Alb., 13 mm. (Scale=0.5 mm.) | S. F. Light, 1 male, 2 females; Mokuleia, Oahu, shallow water, reef, 1938, Degener, 2 males, 1 female, 1 juv; Waikiki Marine Lab., under stones, near shore, Feb. 9, 1942, G. S. Mansfield, 1 male; Waikiki) NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 59 Marine Lab., Feb. 12, 1942, G. S. Mansfield , 1 male, 1 female; Waikiki Marine Lab., Mar. 6, 1942, G. S. Mansfield, 2 males; Waikiki Marine Lab., Mar. 13, 1942, G. S. Mansfield, 1 male, 1 female, 2 ovig. females; Waikiki Marine Lab., Mar. 22, 1942, G. S. Mansfield, 3 males; Waikiki Marine Lab., Mar. 23, 1942, G. S. Mansfield, 2 males, 1 female, 1 ovig. female; Waikiki Marine Lab., Mar. 28, 1942, G. S. Mansfield, 2 females; Waikiki Marine Lab., Apr. 6, 1942, G. S. Mansfield, 1 male, 1 female, 1 ovig. female, 2 juvs; Waikiki Marine Lab., Apr. 24, 1942, G. S. Mansfield, 6 males, 2 females; off Waikiki Marine Lab., Apr. 30, 1942, G. S. Mansfield, 1 female; Oahu, 1945, Army Medical Museum, 1 male, 2 ovig. females; Hanauma Bay, Oahu, July 18, 1945, Army Medical Museum, 1 male. Tuamotus and Societies: Society Is., summer 1945, J. M. Clements, 2 males; remainder Bredin Exped. 1957; Sta. 3-57, Papeete Harbor, Tahiti, anchor chain, April 8, 3 males, 2 females, 2 ovig. females; Sta. 14a-57 Tikahau Atoll, Tuamotus Is., from Holothuria atra, ocean reef, April 12, 7 males (1 damaged), 2 females, 2 ovig. females; Sta. 51-57 Bora Bora, three-eighths mi. S. of Farepiti Point, fringing reef 2-3 ft. in algae in sand, April 23, 1 juv; Sta. 56-57, 57-57, Bora Bora, dredged, April 25, 1 male; Sta. 62-57, Bora Bora, inner edge of outer reef, April 25, 2 males; Sta. 64-57 Motu Tapu Is., Bora Bora Is., along northwestern shore, shallow water, April 26, 2 males, 2 females; Sta. 75-57, E. of dock at Uturoa, Raiatea, from coral boulders in muddy water, April 28, 1 female; Sta. 86-57, Baie de Bourayne, Huahine, sandy reef, May 1, 1 male, 1 ovig. female; Sta. 106-57, Moorea, northwestern Motu Fareme I., fossil coral along shore, May 8, 1 male; Sta. 127-57, Moorea, Nuarei Bay, coral in shallow sandy beach, May 11, 2 males. Samoa: Apia, at mouth of river, June 1, 1902, Alb., 1 male; Apia, outer reef, June 27, 1902, Alb., 3 males, 1 female; Apia, outer coral reef at low tide, July 1, 1902, Alb., 1 female; Pago Pago, August 1902, Alb., 4 males, 4 females (1 Sacculina infected), 3 ovig. females; Canton I., 1941, C. A. Ely, 7 males, 8 females, 3 ovig. females; Canton I., Lagoon West, in coral head near Flemings, January 1942, C. A. Ely, 2 females, 3 ovig. females; Tafuna, Tutuila, Jan. 9, 1949, L. Zachowski, 1 female. Fanning Is.: Inner Lagoon, Dec. 16, 1913, Fred and Charlotte Baker, 3 males, 2 females. MrAsuREMENTS.—Males, 6-38 mm.; females, 6—-ca. 28 mm.; ovig. females, S-29 mm.; juvs. 5-8 mm. Remarxks.—A few specimens from Makinog (Cumigium I., between Leyte and Mindanao, tide pool, Aug. 3, 1909, Alb.) have one side of the front bilobed. One of the females has only third and fifth antero- lateral teeth on the right side. 60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 A female from Black Point (Haunama Bay, Oahu, T.H., 1937, L. R. Woodward, S. F. Light) lacks the second anterolateral tooth on the right. In 1957, Stephenson and Hudson gave an extensive synonymy for this species. A further extension to include 7. dispar has been given by Stephenson (1961a). Since then Forest and Guinot (1961) have revived JT. edwardsi Borradaile as a separate species on the basis of three specimens from Tahiti which included a single mature male. The bases of separation of T. edwardsi from T. admete concern the general facies, and also, according to Forest and Guinot, the structure of the male pleopods. T. edwardsi is supposed to be distinguishable on general facies by the following: Cardiac ridges of carapace absent; posterior mesobranchial ridges absent; hands of chelae smoother and with ill-developed carinae; and fourth anterolateral tooth either absent or rudimentary. Forest and Guinot distinguish the male pleopods of T. edwardsi by large subterminal bristles of outer surface of irregular size and the row not extending so far (relatively) backward from the tip, and by large subterminal bristles of inner surface slightly more numerous (relatively) and extending farther backward from the tip. As regards general facies, Stephenson and Hudson (1957) on the basis of 152 specimens stated (pp. 31-32): . . . the only dubious point appears to be the status of Borradaile’s ‘‘var. ed- wardsi,’’ which is typified by the carination and granulation of the outer surface of the hand of the chelipeds. The existence of gradations between smoothness and carination, and the absence of correlated characters which would serve for distinction, suggests that this ‘“‘variety”’ should not be retained. Since then, field examination of several hundred further specimens has confirmed the gradations in cheliped structure and failed to reveal features of color, habitat, or behavior, which merited a de- tailed reinvestigation of the situation. As regards pleopod structure, figures already published cast doubt upon the distinguishing criteria of Forest and Guinot. Thus the outer surface of the figure of Stephenson and Hudson (1957, fig. 31) resembles their 7. admete, while the inner surface resembles their T. edwardsi. To obtain unequivocal data, 25 males were selected at random from the present collection. Since specimens with car- inated chelae predominated, five males with smoother chelipeds (the first five encountered) were added. The total was examined for seven of the supposedly distinguishing features of 7. edwardsi from 7. admete, and the results given in table 1 (p. 113) are arranged with T. edwardsi-like specimens first and JT. admete-like specimens last. In certain features, particularly those of the pleopods, it was difficult NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 61 to place a given specimen in a definite category, but wherever possible this was attempted. The results show that there is every gradation from specimens which might be categorized ‘6/7 edwardsi and 1/7 admete’’ to those which are “7/7 admete,”’ and that there is a lack correspondence be- tween any two features. Several pleopods are figured (fig. 20). These show intermediates between the two figured by Forest and Guinot and extend the range beyond their limits. A further extension is provided by Stephenson and Hudson (1957, fig. 31). Following the publications of Crosnier (1962) and Guinot (1962), the entire collection was reinvestigated separately by each of the present authors who agreed that, if two forms were present, the only hope of separation lay in the carination of the outer surface of the chelipeds. Investigation then showed that in some specimens both upper and middle carinae are strongly developed and granular. In others the upper carina is strongly granular and the middle one, while strongly developed, is without granules. In yet others the upper carina of the smaller cheliped is granular but that of the larger, nongranular. In others there is a distinct but smooth lower carina, while the upper one is barely distinguishable or indistinguishable. Finally in some, the upper carina has disappeared, and the lower one is distinguishable with difficulty or not at all. In other words there is a complete gradation between extremes. T. spiceri, based upon three females, remains a doubtful synonym. Edmondson compares it with 7. pilumnoides but not with T. admete, which it appears to resemble closely. DistrisuTion.—Red Sea and East Africa to Hawaii and Tahiti, including Japan and Australia. Thalamita auauensis Rathbun Figures 21, 22 Thalamita auauensis Rathbun, 1906, p. 874, pl. 12 (fig. 1)—Edmondson, 1951, p. 222, fig. 24b; 1954, pp. 257-258, figs. 32a—d.—Stephenson and Campbell 1957, pp. 319, 320 (in key). MarteriAu.—China: Sta. 5321, Ibugos Is., China Sea, Formosa, 20°19’30” N., 121°51’15” E., 26 fm., white sand, coral, broken shells, Nov. 9, 1908, Alb., 1 male. Philippines: Sta. 5557, Sept. 18, 1909, Alb., 1 ovig. female; Great Tobea I., tide pool, sand, coral, Dec. 15, 1909, Alb., 1 female. Marianas: 148/R.H.B.—-155, Ritidian Point, Guam, July 12, 1945, R. H. Baker, 1 ovig. female; Guam, October-November 1945, J. L. Gressitt, 1 ovig. female; Saipan, in coral heads, 1945, A. H. Banner, 1 juv; 177-G-1, Ifaluk, Caroline Is., 1953, F. M. Bayer, 2 juvs. 62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 (also 7 juvs. of a species near this but too small to identify positively). Samoa: U.01096, 1 ovig. female. MEASUREMENTS.—Male, 17 mm; female, 21 mm; ovig. females, 11-20 mm; juvs., 4-5 mm. Remarxks.—Very close to 7. admete in its general facies and also in its pleopod structure and distinguished by: Carapace being more convex and typically more hirsute; granulation between two lower- most ridges on outer side of chelae; more inclined borders of penulti- mate segment of male abdomen and more elongate ultimate segment; and by marked reduction in spinulation of male pleopods (see figs. Ficure 21.—Thalamita auauensis Rathbun, male, China, Sta. 5321, Alb.: a, pleopod; b, pleopod tip, upper surface; c, abdomen. (Scales=a, 1 mm.; b, 0.5 mm.; c, 5 mm.) Zia.b): Only four slightly recurved stout bristles of more or less uniform size are present on each. Evidently the size of the fourth anterolateral tooth varies as in T. admete. Rathbun (1906) describes this tooth as rudimentary, while Edmondson (1954) implies that it is either very small or absent. Edmondson comments on the bright pink color of specimens. The species is also close to 7. granosimana Borradaile and was | listed as a queried synonym of this species by Stephenson (1961a, pp. 119-121). It differs in the armature of the male*pleopods, in the penultimate segment of the male abdomen being less parallel sided . | | | | NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 63 and ultimate segment longer, and in having a more convex carapace. The front varies in the curvature of both inner orbital lobes and frontal lobes. Edmondson (1951, fig. 24b; 1954, fig. 32a) does not show the conspicuous frontal ridges which are clearly visible in Rathbun’s (1906) plate 12, figure 1. In his figures he shows what appears to be a ridge which continues behind the orbit; in the present specimens this is a groove, not a ridge. DistrisutTion.—Previously known only from Hawaiian waters. Figure 22.—Thalamita auauensis Rathbun, fronts: a, China, Sta. 5321, Alb.; b, Philippines, Great Tobea I., A/b. (Scale=5 mm.) Thalamita beuvieri Nobili Thalamita bouviert Nobili, 1906b, p. 262; 1907, pp. 384-385, pl. 2 (fig. 2).— Rathbun, 1911, p. 210.—Balss, 1938, p. 34.—Crosnier, 1962, pp. 119-121, figs. 201-204, pl. 10 (fig. 2). Thalamita inhacae Barnard, 1950, p. 179, fig. 33g.—Stephenson and Hudson, 1957, pp. 337-339, figs. 2L, 3L, pls. 3 (fig. 2), 7H, 10E.—Stephenson, 1961a, ps zie Materran.—Details of sex, size, and spinulation of the chelipeds are given with each specimen. Australia: Dredged off Bottle and Glass Reef, 2 fm., Melbourne Ward, 1 female, 7mm. (Right cheliped only present, with 2 spines, 1 spmniform tubercle, 1 indistinct tubercle.) Marshalls: Eniwetok Lagoon, 3 mi. N. of Jieraru I., dredged 15-20 fm., June 5, 1946, Taylor, 1 ovig. female, 10 mm. (Both chelipeds with 2 spines, 1 spiniform tubercle, 2 tubercles.) Rongelap Lagoon 2 mi. W. of Busch I., 20 fm., June 21, 1946, Taylor, 1 male, 6 mm. (Both chelipeds with 3 spines, 2 tubercles.) Samoa: Pago Pago, August 1902, Alb., 1 female, 12 mm. (Left cheliped only present with 3 spines, 2 tubercles.) Remarxs.—The above specimens are conspecific with material from both east and west Australia which was identified as T. inhacae 64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 (Stephenson and Hudson, 1957; Stephenson, 1961a). In the single small male, the pleopod differs slightly from that figured in Stephenson and Hudson (1957) in possessing fewer subterminal bristles. On the inner side, only five stout spines are present, while on the outer side there are only five bristles, two of which are large. Barnard’s description of 7. inhacae from Inhaca Bay in South Africa did not include comparison with T. bowmert. The distinctions between the two species seem to be solely in the ornamentation of the hand of the cheliped. In 7. inhacae, five blunt spines and one outer ridge are described by Barnard, while there are three spines and three ridges in 7. bouviert. Throughout the genus Thalamita, spines on the upper surface of the cheliped are invariably borne upon partial or complete carinae, and the presence of five spines necessarily implies more than one carina, with a suspicion of the usual two on the upper surface. These are certainly present in the Australian and present material. Spines are invariably subject to wear and capable of being reduced to obscure tubercles. Evidence of this is given in the details of the present collection. Crosnier (1962) has examined female syntypes of T. bowert and provided the final confirmation of the synonymy. DistRiputTion.—South Africa, Amirantes, Australia, Gilbert, and Tuamotou Is. Thalamita chaptalii (Audouin) Portunus chaptalit Audouin, 1826, p. 83 [figs. in Savigny, 1809, pl. 4 (fig. 1)]. Thalamita chaptalit (Audouin).—Alcock, 1899, pp. 80-81. Thalamita chaptali (Audouin).—Stephenson and Hudson, 1957, pp. 327-328, figs. 2F, 3F, pls. 1 (fig. 3), 7C, 10B.—Forest and Guinot, 1961, p. 34, figs. 21A, B—Sankarankutty, 196la, p. 106.—Crosnier, 1962, pp. 111, 118, figs. 184, 189, 191. Mareriau.—Philippines: Tara I., Mindoro Strait, 130 ft. seine, Dec. 15, 1908, Alb., 1 male, 1 female, 1 ovig. female; Mariveles, Luzon, June 27, 1913, A. M. Reese, 1 male. Melanesia: Florida I., Solomon I., Oct. 28, 1944, Lt. R. E. Kuntz, 1 male, 1 female. Marshalls: S-46-44, dredged 180-200 ft., Bikini Lagoon, Mar. 29, 1946, L. P. Schultz, 2 females. | MEASUREMENTS.—Males, 10-24 mm.; females, 10-18 mm.; ovig. | j female, 13 mm. | DistripuTion.—Madagasear and Red Sea to Australia and Tahiti. | | Thalamita coeruleipes Jacquinot, 1852, pl. 5 (figs. 6-10).—Jacquinot and Lucas, | 1853, pp. 53-54.—Edmondson, 1954, pp. 265-267, figs. 38a-f, 39a.—Stephen- | son and Hudson, 1957, pp. 329-331, figs. 2P, 3P, pls. 2 (fig. 1), 7D, 9B.— | | ) Thalamita coeruleipes Jacquinot | | i fe NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 65 Forest and Guinot, 1961, pp. 32-33.—Crosnier, 1962, pp. 128-130, figs. 219 bis a-b, pl. 11 (fig. 2). Materiau.—Philippines: Sta. 5159, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°11’50” N., 119°54’ E., Feb. 21, 1908, Ald., 1 male, 1 female. Australia: Great Barrier Reef, Apr. 8-May 29, 1952, John K. Howard, 1 male. Marianas: Saipan I., in coral heads, 1945, A. H. Banner, 1 female; Aganta Bay, Guam, Sta. 85-x-44, May 20, 1945, R. H. Baker, 1 female; Agfayan Bay, Guam, 135/RHB 150, July 7, 1945, R. H. Baker, 1 female; Y259, Gillifitz, Yap I., sand, rocky coral covered reef, received 1952, Hiatt, 1 male. Samoa: Apia, at mouth of river, June 1, 1902, Alb., 2 females, 1 ovig. female; Apia, outer reef, June 27, 1902, Alb., 2 males, 1 ovig. female; Apia, outer reef, June 27, 1902, Alb., 1 juv; Apia, coral reef, July 1902, Alb., 1 female; Apia, outer coral reef at low tide, July 1, 1902, Alb., 2 males; Pago Pago, no. 10, August 1902, Alb., 8 males, 2 females, 3 ovig. females; Pago Pago, August 1902, Alb., 2 males. Fanning Is.: Inner Lagoon, Fanning Is., Dec. 16, 1913, Fred and Charlotte Baker, 1 ovig. female. Tuamotus and Societies: Sta. 84a-57 Huahine, head of Baie de Maroe, from branching coral, Apr. 30, 1957, Bredin Exped., 2 males. MEASUREMENTS.—Males, 10-40 mm.; females, 18-53 mm.; ovig. females, 26-46 mm. Remarks.—The fourth anterolateral tooth is smaller than the fifth in small specimens, which hence key out as 7. prymna. ‘They are distinguished from JT. prymna by the form of the front and inner orbital angles. In the Australian specimen the second of the normal anterolateral teeth is missing, and the first is greatly enlarged. DistRisuTION.—Mauritius to Australia and Central Pacific Is. Thalamita corrugata Stephenson and Rees FIGURE 23 Thalamita cooperi Stephenson and Hudson, 1957, pp. 331-332, pls. 1 (fig. 4), 10C [in part, some specimens of 7’. demani Nobili, q.v.]. Thalamita corrugata Stephenson and Rees, 1961, pp. 421-425, figs. 1A, C, EH, F, 2A-C.—Guinot, 1962, p. 9. Not Thalamita cooperi Borradaile, 1902, pp. 206-207, fig. 37.—Sankarankutty, 1961a, p. 122, fig. 113. MareEriAL.—Tuamotus and Societies: Sta. 10a-57, Tickahau | Lagoon, Apr. 11, 1957, Bredin Exped., 2 males (6, 8 mm.), 2 females (5, 6 mm.). _ Remarxs.—In general facies, the present material agrees in all respects with the females from Queensland, described by Stephenson and Hudson (1957) as 7. cooper. | 221-520—67 5 66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 The male pleopod and abdomen (figs. 23a—-c) are described in Stephenson and Rees (1961, pp. 423-424). This pleopod shows con- siderable resemblances to that of 7. trilineata Stephenson and Hudson (1957)(=T. demani Nobili), the differences being extra subterminal bristling on the outer side in 7’. trilineata where there is a relatively close packed elongate patch or double row. The species as a whole is close to 7. demani but differs in the relatively narrower lateral frontal lobes (one-fifth instead of one-third breadth of the medians) and in the median cardiac ridge of the carapace being separated from the mesobranchials. Ficure 23.— Thalamita corrugata Stephenson and Rees, 8 mm. male, Tuamotus and Societies, Sta. 10a—57, Bredin: a, pleopod; b, pleopod tip, upper surface; c, abdomen. (Scales=a, 1 mm.; d, 0.5 mm.; c, 1 mm.) DistTRIBUTION.—Queensland and the Gilbert Is. Thalamita crenata (Latreille) Portunus crenatus Latreille, 1829, fide H. Milne Edwards, 1834, p. 461. Thalamita crenata (Latreille)—Alcock, 1899, pp. 76-77.—Sakai, 1939, pp. 413- 415, pl. 84 (fig. 3)—Edmondson, 1954, pp. 267-269, figs. 39b, 40a-f.— Stephenson and Hudson, 1957, pp. 332-334, figs. 2Q, 3Q, pls. 2 (fig. 3), 7F, 9C.—Sankarankutty, 196la, pp. 106-107.—Crosnier, 1962, pp. 130-132, figs. 220, 226-227, 232-233. Materiau.—Malay Penninsula: Allfrom Siam: Taleui I., Sept. 27, 1923, 1 male (front damaged, median lobes confluent), 2 ovig. females; Koh Chang, Jan. 14, 1924, 1 dried male (left median and lateral NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 67 frontal lobes fused); Koh Chang, Apr. 5, 1924, H. M. Smith, 1 male; Koh Chang, west side of Gulf of Siam, July 15, 1926, 1 ovig. female; Hualap I., Sept. 26, 1926, H. M. Smith, 1 female; Sri Raja, Feb. 5, 1927, H. M. Smith, 1 male; Sri Raja, Feb. 5, 1927, 1 male, 1 female; Spiracha, southeastern Siam, rocks, May 6, 1929, 1 female; Gulf of Siam, Apr. 30, 1934, H. M. Smith, 1 female; Mayor Stream on north side of Goh Chang I., east side of Gulf of Siam, Oct. 28, 1957, Fehlmann, 1 female. Philippines: Grand I., Subig Bay, Luzon, October 1907, Dr. J. C. Thompson, 1 female; Panabutan Bay, shore, Feb. 6, 1908, Alb., 1 ovig. female; Tataan, Simaluc, shore, Feb. 19, 1908, Alb., 1 male, Tataan Tawitawi, shore, Feb. 20, 1908, Alb., 2 males; Borrgoa I., Tawitawi, shore, Feb. 23, 1908, Alb., 1 male, 1 ovig. female; Sandakan, Borneo, seine, Mar. 2, 1908, Alb., 6 females; Jolo, Jolo I., shore, Mar. 6, 1908, Alb., 1 male; reefs opposite Cebu, Apr. 7, 1908, Alb., 1 male; east side Tagbilaran Strait, Bohol I., shore, Apr. 9, 1908, Alb., 4 males, 5 females; San Miguel Harbor, Ticao I., between Burias and Luzon Is., Apr. 21, 1908, Alb., 1 fragmented male; shore about Hoilo River, 130 ft. seine, June 2, 1908, Alb., 2 males, 1 female; Point Jarnelo, Luzon River, 150 ft. seine, July 13, 1908, Alb., 1 male, 2 females; Sta. 5292, July 23, 1908, Alb., 1 male, 1 female; Port San Pio, in small stream near mouth, 20 ft. seine, Nov. 11, 1908, Alb., 1 male; Luzon shore, San Vicente Harbor, seine, Nov. 11, 1908, Alb., 1 male; Luzon shore, San Vicente Harbor, seine, mud, sand, sticks, and leaves, Nov. 13, 1908, Alb., 1 male, 1 female; Port Uson, W. of Pinas I., electric light, Dec. 17, 1908, Alb., 2 males; Nakochin Harbor, Linapacan I., seine, Dec. 19, 1908, A/b., 1 male, 1 female; Ulugan Bay, near mouth of Caiholo River, Palawan I., seine, Dec. 28, 1908, Alb., 1 male, 1 female; Buena Vista, Guimaras I., Hloilo Strait, seine in mouth of river, Jan. 14, 1909, Alb., 1 male; Alimango River, Burias I., Mar. 5, 1909, Alb., 1 male (Saceulina infected), 1 female; San Pascual, Burias 1., tide pool, Mar. 8, 1909, Alb., 1 female; Caumahala Bay, Ragay Gulf, mouth of small stream, Mar. 11, 1909, Alb., 1 male (front damaged), 4 ovig females; Cebu Market, Mar. 26, 1909, Alb., 1 male; Cuyo I., 130 ft. seine, Apr. 9, 1909, Alb., 1 male, 1 female; Nato River, east coast Luzon, San Bernadino Strait to San Miguel Bay, June 18, 1909, Alb., 1 female; Fishermen Makinog, Camiguin I., between Leyte and Mindanao I., tide pool, Aug. 3, 1909, Alb., 1 female; Labuan Blanda I., Dec. 14, 1909, Alb., 2 males, 3 females (2 ovig.); Great Tobea I., tide pool, sand, coral, Dec. 15, 1909, Alb., 1 male; Iloilo, Panay I., Apr. 6, 1929, H. C. Kellers, 1 female; Jolo, Sulu, mud, sand, sticks, and leaves, July 13-19, 1929, Herre collection; 1 male, 1 female (Philippine Is.; EK. A. Mearns, 3 males, 2 females). 221-520 676 68 PROCEEDINGS OF .THE NATIONAL MUSEUM VOL. 120 Australia: Little Lagoon and vicinity northeastern end of Groote Eylandt, Gulf of Carpentaria, 1948, R. R. Miller, 1 fragmented male; Yirrkala, 1948, Arnhem Land Exped., 2 fragmented males; Umba Kumba, south side of Little Lagoon, northeastern end Groote Eylandt, Gulf of Carpentaria, taken between Apr. 6 and May 18, 1948, R. R. Miller, 1 male, 1 female; Ironstone Reef at Rocky Beach +3 mi. S. of the point E. of Yirrkala and NW. of Cape Arnhem, N.T., July 18, 1948, R. R. Miller and F. M. Setzler, 1 male, 1 female (soft and fragmented). Palau Is.: GVF, High I., Atoll Project, Sta. 12, 1955, H. A. Fehl- mann, S. Pierce, R. Harry, 2 males; GVF, High Atoll Project, Sta. 60, W. of Ebadul’s Pier, north end of Koror IJ., sand and eel grass flat, 1011—Crustacea, Aug. 5, 1955, Fehlmann and Harry, 1 male. Marianas: Ajayan River, Guam, 112X-65, June 19, 1945, D. H. Johnson, 1 male, 2 females; Oca Point, Guam (Alupat I.), July 5, 1945, L. P. McElroy, 1 female; Y114A, from under stones in holes in dead coral blocks, intertidal zone, Tomil Harbor, received June 23, 1952, 2 males, 2 females, 7 juvs. Samoan Is.: Apia, outer reef, June 22, 1902, Alb., 1 fragmented male; Apia, holes along Vailele River above tidewater, July 1902, Alb., 1 juv; Pago Pago, no. 10, August 1902, Alb., 1 male. Tuamotus and Societies: Sta. 26-57, Maiai I., Tikahau Atoll, lagoon side, off beach, Apr. 13, 1957, Bredin Exped., 4 males; Sta. 27, Maiai I., Tikahau Atoll, Tuamotu Is., lagoon side, along shore, seining marly clay bottom, Apr. 14, 1957, Bredin Exped., 1 female. MEASUREMENTS.—Males, 17-91 mm.; females, 19-63 mm.; ovig. females, 40-63 mm.; juvs., 8-16 mm. ReMARKS.—In juveniles narrower than 10 mm., the fourth antero- lateral tooth is smaller than the fifth, and the specimens key out in the wrong direction at couplet 3 in Stephenson and Hudson (1957, p. 316). In the males ‘89 mm. Port Uson, West of Pinos I., Dec. 17, 1908, Alb.,” and ‘91mm. Cebu market, Mar. 26, 1909, Alb.,” the mesogastric ridges are concave anteriorly instead of being straight, and the basal antennal joint is more crestlike than usual. Possibly these features are related to size, these being the largest specimens so far examined by the senior author. The single Sacculina infected male ‘34 mm, Alimango River, Burias I., Mar. 5, 1909; Alb.,” in addition to having the characteris- tically broader abdomen, has an unusually convex carapace. The male “47 mm, Koh Chang, Siam, Jan. 14, 1924,” has the sub- median and lateral frontal lobes fused on the left side. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 69 A male ‘60 mm, Sta. 26-57, MaiaiI., Apr. 13, 1957,” is abnormal in possessing a small additional frontal lobe between the lateral frontal lobe and the supraorbital angle on the left side. The right pleopod of a male “67 mm, G.V.F., High I. Atoll Project, Sta. 12; 1955” is grossly malformed, and bears what is probably a crustacean parasite. DistriBpuTION.—Durban and Red Sea to Japan, Australia, Hawaii, and Society Is., and now Tuamotus. Thalamita dakini Montgomery Figure 24 Thalamita dakini Montgomery, 1931, pp. 432-433, pls. 23 (fig. 3), 28 (fig. 4).— Stephenson and Hudson, 1957, pp. 334-335.—Stephenson, 1961la, pp. 118- 119, figs. 2D, 3J, pls. 3 (fig. 3), 41, 5F. Thalamita medipacifica Edmondson, 1954, pp. 260-262, figs. 34a,b, 35a. Materisau.—Marianas: Ritidian Point, Guam, Sta. 148—-R.H.B. 155, July 12, 1945, Baker, 1 male, 1 ovig. female. Figure 24.—Thalamita dakini Montgomery, male, Hawaii, Waikiki Marine Lab., Mar. 28, 1942, Mansfield: pleopod tip, upper surface. (Scale=0.5 mm.) Gilbert Is.: Howland I., 1926, C. H. Edmondson, 1 male, 1 ovig. female. Hawaii: Black Point, Haunama Bay, Oahu, 1937, L. R. Woodward, S. F. Light, 1 female, 1 ovig. female, 1 juv; Waikiki Marine Lab., Honolulu, Mar. 28, 1942, G. S. Mansfield, 1 male; coast near Kawailoa, Oahu, under growth of coralline algae, Apr. 4, 1942, G. S. Mansfield, 3 females (2 ovig.); Waikiki Marine Lab. from coral, Apr. 24, 1942, 5 males, 1 ovig. female, 1 juv. Tuamotus and Societies: Society Is., summer 1925, J. M. Clements, 1 male, 1 ovig. female; Bora Bora, inner edge of outer reef, Sta. 62-57, 2 males, 3 females (2 ovig.), 1 juv; Nuarei Bay, Moorea, Sta. 127-57 70 PROCEEDINGS OF THE NATIONAL MUSHUM VOL. 120 coral in shallow, sandy beach, May 11, 1957, Bredin Exped., 1 male, 1 ovig. female. MEASUREMENTS.— Males, 7-28 mm.; females, 10-15 mm.; ovig. females, 10-21 mm.; juv., 6 mm. Remarks.—In previous papers (Stephenson and Hudson, 1957; Stephenson, 1961a) Edmondson’s species was included as a queried synonym. Doubt arose from minor differences between the male pleopod as figured by Edmondson (1954, figs. 34g, h) and those of the single male available from Australia. Edmondson showed two terminal spinules on the inner surface, which had no counterpart in the Australian specimen, and five instead of three subterminal spinules on the outer surface. The present material shows variability in armature of male pleopods (see table 2, p. 114). DisrrisutTion.—Western Australia and Hawaii (Edmondson, 1954; Stephenson, 1961a). Thalamita danae Stimpson Figures 25a-e, 26a—c Thalamita crenata Dana, 1852a, pp. 282-283, pl. 17 (figs. 7a,b) (not Latreille, 1829). Thalamita danae Stimpson, 1858, p. 37.—A. Milne Edwards, 1861, pp. 366-367, pl. 36 (figs. 1, la-c).—de Man, 1902, pp. 644-645, pl. 21 (fig. 28) —Stimpson, 1907, p. 85, pl. 11 (figs. 1,la)—Shen, 1934, pp. 52-53, figs. 15, 16a—c.— Sakai, 1939, pp. 4138, 415, pl. 85 (fig. 3).—Stephenson and Hudson, 1957, pp. 335-337, figs. 2N, 3N, pls. 3 (fig. 1), 7G, 10D.—Crosnier, 1962, pp. 135-1386 (under 7’. forestz), figs. 228, 232-233. Nore ON SYNONYMY AND NOMENCLATURE.—Crosnier (1962), in describing 7’. foresti, revised the synonymy of TY. danae. To his synonymy, Sakai (1939) is now added. MarrriaL.—Two forms are recognized in adult males, called forms A and B, respectively, distinction being by male abdomens and ple- opods. ‘These are listed separately as are females and juveniles. MALES (FORM 4A) Malay Peninsula: Hualap I., Siam, Sept. 26, 1926, H. M. Smith, 1 male. Philippines: Guijulugan, Negros, shore, seine, Apr. 2, 1908, Alb., 1 male; reefs opposite Cebu, Apr. 7, 1908, Alb., 2 males; Dumurug Point, Masbate, shore, Apr. 19, 1908, Alb., 8 males (1 Sacculina infected, 1 with broad abdomen); Busin Harbor, Burias I., between Burias and Luzon, Apr. 23, 1908, Alb., 1 male; Little Santa Cruz L., Zamboanga, Mindanao, soft coral, marginal reefs, May 26, 1908, Alb., 1 male; Masinloc Bay, Tacubales Province, Nov. 22, 1908, Alb., 1 male; Caumahala Bay, Ragay Gulf, mouth small stream, Mar. 11, 1909, Alb., 1 male; Canino I., near Daet, June 15, 1909, Alb., 1 male; NO. (8056 PORTUNID CRABS—STEPHENSON AND REES 71 Batan I., July 22, 1909, Alb., 2 males; near Mariveles, Luzon, 1913, Albert M. Reese, 1 male; mangrove swamp, Zamboanga, Mindanao, Mar. 8, 1914, F. Baker, 1 male. Figure 25.—Male pleopods and tips. Thalamita danae Stimpson: a, c, d, form A, Philip- pines, Dumurug Pt., Alb.; b, e, form B, China, Nau Wau, Alb. T. foresti Crosnier: f, China, Sabtan I., 4/b. (Scales= a, b, 5 mm.; c, 0.5 mm.; d-f, 1 mm.) 72 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Australia: Sta. 11B, Sydney, Mar. 21, 1948, F. D. McCarthy, 1 male; reef at Yirrkala, July 12, 1948, R. R. Miller, 1 male; ironstone reefs (not coral) and rock pools at Yirrkala, NW. of Cape Arnhem, part of M48-21, emulsifiable rotenone, Aug. 6, 1948, R. R. Miller and natives, 1 male; Arnhem Land Exped., M48-21, reef at Yirrkala, NW. of Cape Arnhem, Aug. 12, 1948, R. R. Miller, 1 male. Palau: No. 10-814 west end Koror I., Madalai district, 1 {t., July 8, 1955, GVF, 1 male. Marianas: Y-128G from under rocks, tidal flats, Tomil Harbor, received 1952, Hiatt, 1 male. MALES (FORM B) China: Nau Wau, Formosa, China Sea, Jan. 27, 1910, Alb., 1 male. Philippines: Gubat Bay, tide pool, June 23, 1909, Alb., 1 male (right lateral frontal lobe missing). FEMALES AND JUVENILES Philippines: Guijulugan, Negros, station shore, sand, gravel, mud, Apr. 2, 1908, Alb., 2 females, 9 juvs; Dumurug Point, Masbate, shore, Apr. 19, 1908, Alb., 6 females, 2 juvs; Busin Harbor, Burias I., between Burias and Luzon, Apr. 23, 1908, Alb., 1 ovig. female; Point Jarnelo, Luzon, 150 ft. seine, July 13, 1908, Alb., 1 ovig. female; Tivanao I., near Palawan Is., reef, Dec. 31, 1908, Alb., 1 ovig. female; San Pascual, Burias I., tide pool, Mar. 8, 1909, Alb., 1 ovig. female; Caumahala Bay, Ragay Gulf, mouth small stream, Mar. 11, 1909, Alb., 2 females; Batan I., July 22, 1909, Alb., 1 female, 3 ovig. females; Nasigit, Mindanao I., tide pools, Aug. 1, 1909, Alb., 1 female; Mactan I., tide pools, Aug. 31, 1909, Alb., 1 female; Great Tobea L., tide pool, sand, coral, Dec. 15, 1909, Alb., 1 female, 1 ovig. female; Dumaguete, Negros Is., July 11, 1931, Herre collection, 1 female. Australia: Sta. 11B, Sydney, Mar. 21, 1948, F. D. McCarthy, 1 female, 1 ovig. female. MEASUREMENTS.—Males form A, 22-73 mm.; males form B, 36, | 46 mm.; females, 18-53 mm.; ovig. females, 42-56 mm.; juvs., 9-18 © mm. Remarks.—Form A males have a relatively long ultimate segment of the male abdomen (almost as long as penultimate) with sides straight rather than concave (fig. 26a). In form B the ultimate abdominal segment is shorter (about four-fifths the length of penulti- mate) with sides markedly concave (fig. 26c). In this concavity it resembles T. foresti, but this species is distinguished by the broader, distally swollen penultimate segment, and the more elongate ultimate segment (fig. 26d). NO. 3556 PORTUNID CRABS—STEPHENSON AND REES (3) The pleopods of form A males are relatively stout and end abruptly (figs. 25a, c, d). Subterminally on the outer side there are approxi- mately six very short, stout, tubercle-like, laterally directed spines, overlapped and succeeded by about six bristles which are mostly forwardly directed. Subterminally on the inner side there are a few Al Bi AA Ficure 26.—Male abdomens. Thalamita danae Stimpson: a, form A (normal), Philip- pines, Dumurug Pt., 4/b.; b, form A (abnormal), Philippines, Dumurug Pt., Alb.; c, form B, China, Nau Wau, Alb. T. foresti Crosnier: d, China, Sabtan I., Alb. (Scale= 5 mm.) sparsely arranged hairs. In form B, the pleopods are thin, gradually tapering, and distinctly curved near the tip. Subterminally on the outer side there are about six stout, elongate, forwardly directed bristles. On the inner side there is an extended row of elongate, forwardly directed spines. 74 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 The pleopods of form B differ markedly from those of form A and related species such as 7. coerulerpes, T. crenata, T. prymna, and T. stimpsoni. But for the similarity in general facies, it would have been placed in a different “group” within the genus (see Stephenson and Hudson, 1957, pp. 320, 362). Form B is very close to T. foresti in the form of the male abdomen and in possessing a slender curved pleopod. It differs in having stouter pleopod bristling, in not having a recurved pleopod tip, and in the form of the front. In one of the form A males (fig. 266) the abdomen is much broader than long, but the pleopods are normal. DistRIBUTION.—Mozambique and Red Sea to Marshalls, Gilberts, and Fiji, including south Japan and north Australia. Recorded from New Zealand by Miers (1876) and by Heller (fide Doflein, 1904) and repeated by later workers, e.g., Stephenson (1961b). As Hutton (1882) has shown, this is almost certainly a mistaken locality. Thalamita demani Nobili Thalamita demani Nobili, 1905, p. 402; 1906a, pp. 209-210.—Crosnier, 1962, pp. 124-125, figs. 200, 208-209. Thalamita invicta de Man, 1895, p. 565, pl. 13, (figs. 11, 11a). Thalamita cooperi Stephenson and Hudson, 1957, pp. 331-332 (in part). Thalamita trilineata Stephenson and Hudson, 1957, pp. 359-360, figs. 2E, 3H, pls. 6 (fig. 4), 85, 10L.—Stephenson, 196la, p. 124, pl. 4 (fig. 2L). ?Thalamita invicta Thallwitz, 1891, pp. 46-47, fig. 11. Not Thalamita cooperi Borradaile, 1902, pp. 206-207, fig. 37—Sankarankutty, 196la, p. 122, fig. 113. MareEriau.—Philippines: Bataan, Simaluc, shore, Feb. 19, 1908, Alb., carapace, two chelae, and one walking leg only of a female (7 mm.). ReEMARKS.—Crosnier (1962) has clearly shown that T. trilineata Stephenson and Hudson (1957) is a synonym of 7. demani Nobili. Reexamination of the holotype of 7. trilineata (Aust. Mus. Reg. No. P.2863) and also specimens from Western Australia (W. Aust. Mus. Reg. No. 84-60) confirms this synonymy. In addition, two specimens in the Australian Museum (Aust. Mus. Reg. Nos. P.7546 (9 mm. female) and P.12754) reported as T. cooperi by Stephenson and Hudson (1957, pp. 331-332) belong to this species, as does a third (Aust. Mus. Reg. No. P.7546, 10 mm. female). Distrisution.—Red Sea, Madagascar, and both east and west Australia. Thalamita foresti Crosnier Figures 25f, 26d Thalamita danae de Man, 1887b, pp. 78-79, pl. 4 (figs. 8, 9). Thalamita forestt Crosnier, 1962, pp. 132-136, figs. 221-223, 229-231, pl. 13 (fig. 1). NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 75 ?Thalamita danae Alcock, 1889, pp. 77-78.—Barnard, 1950, p. 174. Not Thalamita danae Stimpson, 1858, p. 39. Marerriau.—Malay Peninsula: Koh Pipedon, Pocket Bay, Benjal, Siam, Mar. 10, 1925, Hugh M. Smith, 1 female. China: Sabtan I., China Sea, vicinity Hong Kong, Nov. 8, 1908, Alb., 1 male. Philippines: Nogas Point, Panay, shore seine, Feb. 4, 1908, Alb., 1 ovig. female; base reef, Gulf Boni, Dec. 17, 1909, Alb., 1 female. M8ASUREMENTS.—Male, 41 mm.; female, 56 mm.; female, damaged, ca. 37 mm.; ovig. female, 36 mm. Remarks.—Crosnier (1962) distinguishes this species from the very similar 7. danae on the basis partly of adult male features and partly on general facies as follows: a. Penultimate segment of the male abdomen is much wider than long, diverging in its proximal two-thirds, then narrowing con- spicuously toward its end, while the ultimate segment has markedly concave borders. In the single male in the present collection, the penultimate segment (fig. 26d) is broader than figured by Crosnier. It should be noted that form B of 7. danae approaches 7’. foresti in the form of the male abdomen. (Compare figure 26¢ with Crosnier’s figure 223.) b. Male pleopod is elongate, tapering with a recurved tip bearing small spinules and very different from that of 7. danae (form A). However, forms A and B of 7. danae differ markedly themselves. ce. According to Crosnier the basal antennal joint in T. forests bears a more prominent crest carrying sharper granules. In the present collection of 7. danae (form A), there are specimens which cannot be distinguished from 7’. foresti on this feature. d. Different form of the front. Here again examples of 7. danae (form A) show some of the points, for example prominence of the median frontal lobes, which Crosnier lists as diagnostic features of T. foresti. However, typical specimens do differ, and in all cases the median frontal lobes in 7’. foresti are relatively smaller and more rounded than in 7’. danae. A further small distinction lies in the infraorbital lobe which is more ventrally inclined in 7. foresti, the difference being most obvious in ventral view. DistriBuTIoN.—Previously Madagascar and possibly Andaman I. and Mozambique. Thalamita gatavakensis Nobili Thalamita pilumnoides var. gatavakensis Nobili, 1906b, p. 262; 1917, p. 384. Thalamita pilumnoides gatavakensis Nobili.—Forest and Guinot, 1961, pp. 34-86, figs. 23-25. 76 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Thalamita gatavakensis Nobilii—Crosnier, 1962, pp. 106-108, figs. 156 bis a-c, e, 177 bis a-b. Thalamita granosimana Stephenson, 1961a, pp. 119-121, figs. 2H, 4A, pls. 4J, 5G [not Thalamita granosimana Borradaile]. Materiau.—Philippines: Sta. 5165, Observation I., Sulu Archi- pelago, Tawitawi Group, 4°58’20”N., 119°50’30”E., 9 fm., coral, Feb. 24, 1908, Alb., 3 males. Tuamotus and Societies: Sta. 3-57, Papeete Harbor, Tahiti, anchor chain, Apr. 8, 1957, Bredin Exped., 3 males, 2 females, 2 ovig. females; Sta. 62-57, Bora Bora, inner edge of outer reef, Apr. 25, 1957, Bredin Exped., 3 males. MEASUREMENTS.—Males, 4-8 mm.; females, 7, 8 mm.; ovig. females, 9, 13 mm. RemMarKs.—The present material greatly resembles that described by Forest and Guinot, including Nobili’s, from the Tuamotus. The only differences are that the mesobranchial ridges of the carapace are more distantly separated from the median cardiac, the mesogastric are less interrupted in the midline, and the dactyl of the walking legs is slightly shorter than the propodus. Minor differences also existed between Crosnier’s single specimen and Nobili’s type, and between Nobili’s syntypes (see Crosnier, 1962, fig. 156 bis a-c, e). None of the males bore pleopods; these were smaller than the lectotype figured by Forest and Guinot. Distinctions from J. pilumnoides Borradaile are given under that species; a fuller description of the present species, albeit misidentified, is given in Stephenson (1961a). DistrRipuTION.—Madagascar, Western Australia, and Tuamotus. Thalamita gloriensis Crosnier Thalamita gloriensis Crosnier, 1962, pp. 98-100, 102-103, figs. 155, 156 bis d, 159-160, 165-167, 169. Marerru.—Melanesia: No. 4, Bougainville, Jan. 10, 1945, W. Bartos, 1 ovig. female (10 mm.). Marianas: Saipan, in coral heads, 1945, A. H. Banner, 1 male (6 mm.). Marshalls: East Rigili I., Eniwetok Atoll, rock flats, May 30, 1946, J. P. E. Morrison, 1 ovig. female (18 mm.). Hawaii: Waikiki Marine Lab., from seaweed, Feb. 1, 1942, G. S. Mansfield, 1 juv. (4 mm.). Tuamotus and Societies: Huahine, off Point Teffaao Sta. 90a—57, May 2, 1957, Bredin Exped., 1 ovig. female (6 mm.). Remarxs.—Only the following minor differences with Crosnier’s description were noted: NO. 3556 PORTUNID CRABS—STEPHENSON AND REES ri a. Marshall Is. specimen, 3rd anterolateral tooth not much smaller than the other teeth. b. Above plus Huahine specimen, one or two well-developed spines on wrist behind that at the articulation; wrist more spiniform than in Crosnier’s figure 165. DistriBuTION.—Previously only from Glorious Is., Madagascar. a Figure 27.—Thalamita granosimana Borradaile, male, Philippines, Caiholo R., Alb.: a, pleopod; b, pleopod tip, upper surface. (Scales=a, 1 mm.; 5, 0.5 mm.) Thalamita granosimana Borradaile FIGURE 27 Thalamita granosimana Borradaile, 1902, pp. 202-203.—Crosnier, 1962, pp. 103- 106, figs. 171-172, 175-177, pls. 8 (fig. 2), 13 (fig. 3). Not Thalamita granosimana Stephenson, 196la, pp. 119-121, figs. 2E, 4A, pls. 3 (fig. 4), 43, 5G (= Thalamita gatavakensis Nobili). Materi.t.—Philippines: Sta. 5158, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°12’N., 119°54’30”E., 12 fm., Feb. 21, 1908, Alb., 78 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 2 males; Sta. 5159, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°11/50”N., 119°54’E., Feb. 21, 1908, Alb., 10 males, 6 females, 4 ovig., females, 1 Sacculina infected specimen; Sta. 5160, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°12’40”N., 119°55’10”E., 12 fm., sand, Feb. 22, 1908, Alb., 1 female, 3 ovig. females; Tomindao I., anchorage, electric light, Feb. 26, 1908, A/b., 1 female; Sta. 5169, Sibutu i., 4°3215”"N., 119°22’45”E., 10 fm., Feb. 27, 1908) 2Alb. males, 1 ovig. female; San Miguel Harbor, Ticao Is., between Burias and Luzon, Apr. 21, 1908, Alb., 1 male; Sta 5218, Anima Solo Is., be- tween Burias and Luzon, 20 fm., Apr. 22, 1908, Alb., 2 males, 1 ovig. female; Caiholo River, Ulugan Bay, seine, Dec. 29, 1908, Alb., 4 males (1 damaged), 1 female, 1 ovig. female. MEASUREMENTS.—Males, 8-24 mm.; females, 7-16 mm.; ovig. fe- males, 9-18 mm.; Sacculina infected specimen, 13 mm. REMARKsS.—Crosnier’s (1962) identification and redescription of this species is based upon comparison with Gordon’s refiguring of Bor- radaile’s type, and there are no doubts regarding Stephenson’s (1961a) misidentification. Male pleopods of the present material (fig. 27) bear fewer subter- minal spines than those figured by Crosnier (1962, figs. 175-177) but evidently belong to the same species. DistripuTION.—Madagascar, Maldive-Laccadive Archipelago. Thalamita imparimana Alcock FiIGuRES 28; PLATE 7A Thalamita imparimanus Alcock, 1899, p. 87.—Aleock and Anderson, 1900, pl. 47 (figs. 3,3a). Marerrau.—China: Sta. 5304, China Sea, vicinity Hong Kong, 21°46’/N., 114°47’E., 34 fm., black mud, Aug. 9, 1908, Alb., 1 male (8 mm.). Philippines: Sta. 5131, Island off Panabutan Point, 27 fm., Feb. 6, 1908, Alb., 1 male (11 mm.). Remarxs.—Alcock’s brief diagnosis serves to identify this species. It resembles 7. investigatoris in its long walking legs and in the form of the chelipeds. In these the upper surface bears rounded granules; there are no carinae, and only two spines are present on the upper surface (including the one at the wrist articulation). The species is characterized by the considerable overlap between median and sub- median frontal lobes and by the absence of spinules on the posterior borders of the propodite of the fifth leg. The male abdomen and pleopod appear not to have been described; both are highly characteristic. Penultimate segment of abdomen with markedly convex lateral borders (fig. 28¢c). Pleopod with re- NOs 3556 PORTUNID CRABS—STEPHENSON AND REES 79 markable double curvature: tip flared and swollen, surmounted with a crest of long bristles; overall resembling a crested crane. Beyond basal lobes outer surface bare until just behind tip, here a single row of short hairs. Conspicuous crest on inner surface (which because of curvature becomes directed anteriorly) consisting of single row of about 30 elongate spines merging distally with about 15 short bristles. Tip of appendage bearing four stout bristles. DistTrRiBuTION.—Previously apparently only from Alcock’s material from Ganjan Coast, India. Ficure 28.—Thalamita imparimana Alcock, male, Philippines, Sta. 5131, 4/b.: a, pleopod; b, pleopod tip, upper surface; c, abdomen. (Scales=a, 1 mm.; b, 0.5 mm.; c, 1 mm.) Thalamita integra Dana Thalamita integra Dana, 1852b, p. 85.—Alcock, 1899, p. 85.—Sakai, 1939, pp. 414, 420-421, fig. 15, pl. 84 (fig. 2)—Edmondson, 1954, pp. 252, 253-254, figs. 27a-e, 28a.—Stephenson and Hudson, 1957, pp. 339-341, figs. 2H, 3H, pls. 38 (fig. 3), 71, 10F.—Sankarankutty, 1961la, p. 105; 1961b, p. 122.—Crosnier, 1962, p. 103, figs. 156, 161, 170. MarertAu.—Philippines: Alb., 1 female (no further data); Sta. S169, Sibutu 1., 4°32’715N”., 119°22’45”E., 10 fm.,Keb. 27, 1908, Alb., 1 male; Romblon, 150 ft. seine, Mar. 26, 1908, Alb., 1 ovig. female; Tilig, Lubang, beach, July 14, 1908, Alb., 1 ovig. female; SO PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 San Pascual, Burias I., tide pool with copper sulphate, Mar. 8, 1909, Alb., 2 males. Palau Js.: Sta. 12-832, Madalai district, west end Koror I., man- grove shore grading into mud and sand flat, July 9, 1955, GVF, 2 males. Marianas: Sta. 96/x-49, Oca Point, Guam, June 26, 1945, D. H. Johnson, 2 males. Japan: Okinawa, received Oct. 22, 1945, Maj. Wm. Thomas, 1 male. Samoa: Pago Pago, August 1902, 1 male. Hawaii: Pearl Harbor, Oahu, September 1920, Paul Bartsch, 2 males. MEASUREMENTS.—Males, 8-32 mm.; female, 18 mm.; ovig. females, 16-24 mm. Remarxs.—The males from Pearl Harbor have more terminal bristles on the outer side of the pleopod than described and figured by Stephenson and Hudson (1957). ‘The larger specimen (32 mm.) has about 18 bristles. Distrisution.—Hast Africa to Tahiti and Hawaii, including Australia. Thalamita mitsiensis Crosnier FIicuRE 29 Thalamita mitsiensis Crosnier, 1962, p. 127, figs. 212-213, 216-218. MateriAu.—Philippines: Sta. 5159, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°11’50”N., 119°54’E., Feb. 21, 1908, Alb., 2 males (both 12 mm.). Remarks.—the first anterolateral tooth is much the stoutest and the second to fourth subequal. Crosnier’s description states, ‘“Dents 1-3 subégales, dent 4 de taille légerément inférieure,” although in his figure (fig. 212) the first tooth is shown as the stoutest. The posterior border of the propodus of the fifth leg bears seven to eight spines, not six. DistrRIBUTION.—Previously only from Mitsio Is., Madagascar. Thalamita multispinosa, new species PLATE 7B Mareriat.— Hawaii: Off Waikiki Marine Lab., Honolulu, Apr. 30, 1942, G. S. Mansfield, 1 female (21 mm., holotype, cat no. 112206). Description: Front: 6-lobed, protruding slightly beyond supra- orbital angles. Median lobes rounded, deeply separated from each other and from the intermediates which are about 1% times their breadth. Lateral frontal lobes small with shallow separation from intermediates. Supraorbital angles short, sharply arched. Two conspicuous indentations in upper border of orbit. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES Sl Anterolateral teeth: Five. First stoutest and bluntest, remainder sharp with fourth the smallest. Carapace: Relatively long (B/L=1.3), covered with fine pile through which coarsely granular ridges are evident. Frontals, small elevated eranular patches; protogastrics curved, slightly irregular; mesogastrics concave anteriorly, distinctly separated. Conspicuous patches oppo- site second and fourth anterolateral teeth. Epibranchials smoothly curving, terminating abruptly with small coarsely granular subtermi- nal patch. Short continuous metagastric. Two broadly separated eardiacs. One pair of mesobranchials slightly in front of cardiacs. Posterior border of carapace apparently smooth, actually micro- scopically granular. Ficure 29.—Thalamita mitsiensis Crosnier, male, Philippines, Sta. 5159, 4/b.: a, pleopod; b, pleopod tip, upper surface; c, abdomen. (Scales=a, 1 mm.; b, 0.5 mm.; c, 1 mm.) Basal antennal joint: Shorter than width of orbit (0.8 times) and bearing short, steeply elevated crest, covered with small, rounded, close-packed granules. Chelipeds: Left only present. Spinous, coarsely granular, and hirsute with squamiform markings on under surface of hand. An- terior border of arm with two long distal spines and shorter proximal spine. Wrist with three usual spines on outer surface and longer than usual spine at inner surface. Outer border of hand with two distinct coarsely granular carinae. Upper surface bearing four sharp spines excluding that at wrist articulation, and without obvious carinae. Inner surface with central carina granular, remainder coarsely granular. Fingers long, sharp, strongly carinated. 82 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Fifth leg: Merus about twice as long as broad, with usual sub- terminal spine on hinder surface; posterodistal border also spinous. Propodus with very numerous small spines (ca. 20). Remarks.—This specimen keys out in Stephenson and Hudson (1957) as T. picta; however, it is neither 7. picta, the related T. wakensis, nor T. philippinensis. It differs in the very large number of spines on the posterior border of the propodus of the fifth leg, hence the specific name. It is probably closest to J. picta but differs in the more rounded frontal and inner orbital lobes and in various details of the carapace ridging. The protogastric ridges are shorter than in 7. picta, the mesogastrics are more distinctly interrupted in the midline and are concave anteriorly. There is a broader interruption between the the metagastric and epibranchial ridges, and the cardiacs are composed of two short, widely separated ridges. The cheliped also differs in having longer and sharper spines on the wrist and hand and in having squamiform markings on the under surface of the hand. Thalamita oculea Alcock Thalamita oculea Alcock, 1899, pp. 91-92.—Alcock and Anderson, 1900, pl. 48 (figs. 3, 3a).—Sakai, 1939, pp. 424-425, text-fig., 18—Crosnier, 1962, pp. 109-111, figs. 173-174, 178-180, 193-194, pl. 9 (fig. 1). MareriAu.—Philippines: Sta. 5165, Observation I., Sulu Archi- pelago, Tawitawi Group, coral, 9 fm., 4°58’20”N., 119°50’30’E., Feb. 24, 1908, Alb., 1 female; Sta. 5561, Tutu Bay (Jolo), coral and sand, Sept. 19, 1909, Alb., 1 female 1 juv. (? female); Labuan, Blanda Is., Buton Strait, marginal coral, Dec. 13, 1909, Alb., 2 males, 1 female. MEASUREMENTS.—Males, 9, 10 mm.; females, 10-18 mm.;juv. 8mm. Distripution.—Madagascar, Seychelles, Japan, and Ceylon. Thalamita parvidens (Rathbun) Figure 30 Thalamonyx parvidens Rathbun, 1907, p. 62, pl. 5 (fig. 9). Thalamita parvidens (Rathbun).—Sakai, 1939, pp. 425-426, fig. 19.—Stephenson, 1961la, pp. 122-124, figs. 2F, 4B, pls. 4 (fig. 1), 4K, 5H.—Crosnier, 1962, pp. 113-115, figs. 182, 185-187, 190, pl. 9 (fig. 2). MareErtau.—Philippines: Sta. 5159, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°11’50’N., 119°54’E., Feb. 21, 1908, Alb., 1 male, 1 female; Tunimdao J. Anchorage, Sulu Archipelago, vicinity Sibutu I., electric light, Feb. 25, 1908, Alb., 1 male, 1 juv; Sta. 5169, Sibutu I., 4°32/15”N., 119°22’45”E., 10 fm., Feb. 27, 1908, Alb., 1 male; Pascao, Ragay Gulf, electric light, Mar. 8, 1909, Alb., 1 male; | Mariveles, Luzon, June 27, 1913, A. M. Reese, 3 males. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 83 Japan: Mogi, near Nagasaki, summer of 1900, Dr. D. S. Jordan and Mr. J. O. Snyder, 1 female. MEASUREMENTS.—Males, 7-25 mm.; females, 8-13 mm.; juv., 8 mm. REMARKS.—Stephenson (1961a) noted that Sakai’s (1939) figure 16 does not show the most distal bristles on the inner side of the male Ficure 30.—Thalamita parvidens (Rathbun), male, Philippines, Mariveles, Reese: a, pleopod; b, pleopod tip, upper surface; ¢, abdomen. (Scales=a, 1 mm.; b, 0.5 mm.; c, 5 mm.) pleopod. The specimen here figured is roughly intermediate between Sakai’s figure and Stephenson’s (1961a) figure 4B. Crosnier’s (1962) figures 185 and 186 show numerous long spines on both sides of the tip of the appendage, more than possessed by any of the present specimens. Crosnier’s figure (fig. 190) of the male abdomen shows a straight- sided ultimate segment. In the larger of the present specimens and 84 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 in the one figured by Stephenson (1961a, pl. 4, fig. K), the sides are distinctly concave. Minor differences in general facies as between the descriptions of different authors have been noted by Crosnier. Possibly later study will show the existence of several subspecies. Distripution.—Madagascar, Carolines, Japan, and Western Australia. Thalamita philippinensis, new species Ficure 31; PLATE 8a Matertau.—Philippines: Sta. 5163, Observation I., Sulu Archipel- ago, Tawitawi Group, 4°59’10”N., 119°51’E., 28 fm., coarse sand, Feb. 24, 1908, Alb., 2 ovig. females; Sta. 5165, Observation I., Sulu Archipelago, Tawitawi Group, 4°58’20”N., 119°50’E., 9 fm., coral, Feb. 24, 1908, Alb., 3 females (1 ovig.); Sta. 5218, Anima Solo I., between Burias and Luzon, 20 fm., Apr. 22, 1908, Alb., 5 males, 5 females (1 ovig.); Davao Bay, from pearl oysters, May 18, 1908, Alb., 1 male; Sta. 5249, Gulf of Davao, Lanang Point, 7°06’06”N., 125°- 40’08”E., 23 fm., May 18, 1908; Alb., 1 male (holotype—cat. no. 112238); Sta. 5251, Gulf of Davao, Linao Point, 7°05’12”N., 125°- 39'35”E., 20 fm.; May 18, 1908, Alb., 1 female; Sta. 5252 and Sta. 5253, Linao Point, Gulf of Davao, 7°04’48”N., 125°39’38”E., 28 fm., coral, May 18, 1908, Alb., 3 males, 2 females (1 ovig.); Sta. 5254, Gulf of Davao, Linao Point, 7°05’/12”N., 125°39’35”E., 20 fm., May 18, 1908, Alb., 1 male, 2 females; Sta. 5254, Gulf of Davao, Linao Point, 7°05’- 42”N., 125°39'42”E., 21 fm., sand coral, May 18, 1908, Alb., 2 males. Description.—Front: Six lobes, all rounded and distinctly sep- arated; medians on lower plane than remainder and approximately two-thirds width of submedians; laterals roundedly triangular. Inner orbital lobes moderately broad, almost straight, sloping slightly back- ward, and almost the length of submedian lobes. Anterolateral teeth: Fourth very much smaller than remainder, but still clearly visible and blunt. Remainder all sharp with first the stoutest, followed by second, with third and fifth fairly slender. Carapace: Covered with short pile and finely granular. All normal ridges present excepting anterior mesobranchial and cardiac. Meso- gastrics interrupted in midline, and epibranchial broadly interrupted at cervical grooves. Basal antennal joint: Shorter than major diameter of orbit (ca. 0.8 times in larger specimens) and bearing short acute ridge with rounded eranules. Chelipeds: Coarsely granular on upper surfaces and smooth below. Right much stouter than left. Anterior border of arm bearing three stout spines and one or more tubercles. Spine at inner angle of wrist NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 85 stout and well developed, three spines on outer surface, sometimes reduced to tubercles. Upper surface of hand with inner and outer spine near middle and with two normal distal spines often reduced to rounded tubercles. Outer surface typically with three carinae, lower- most running to immovable finger, distinct granular middle carina and indistinct granular upper carina. In larger chelae of male these carinae indistinct or unrecognizable. Fifth leg: Merus slender (L/B=ca. 3), smooth posteriorly except for usual spine; posterior border of propodus bearing four to eight spines. Male abdomen: Penultimate segment much broader than long (ca. Ficure 31.—Thalamita philippinensis, new species, male holotype: a, pleopod; b, pleopod tip, upper surface; d, abdomen. Philippines, Sta. 5218, Alb.: c, pleopod tip, upper surface. Philippines, Sta. 5254, Alb., 25 mm. male; e¢, abdomen. (Scales=a, 1 mm.; b, c, 0.5 mm.; d, e, 5 mm.) 1% times) with bulging sides. Ultimate segment rounded equilateral triangle. Male first pleopod: Fairly stout, smoothly curving to oblique tip bearing flared membrane. Both sides without armature until near tip. Here outer surface typically with five to seven stout subterminal spines directed obliquely forward and decreasing in size toward tip, and usually with two or three thinner bristles widely spaced and lying back from tip. In largest male single subterminal spine present. Inner surface bearing two to four stout spines opposite widest portion of flared membrane; these almost at right angles to appendage. MEASUREMENTS.—Males, 7-14 mm.; females, 8-12 mm.; ovig. females, 7-12 mm. 991-520—67——7 86 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Remarks.—This species keys out with T. wakensis Edmondson (1925) and T. picta Stimpson (1858). (See Stephenson and Hudson, 1957, p. 316.) It is closest to T. wakensis, sharing with it possession of a distinctly granular basal antennal joint and a male pleopod with stout anteriorly directed bristles on the outer side near the tip. It differs from T. wakensis in: a. More rounded median and submedian frontal lobes; see Edmondson (1925, fig. 7a; 1954, fig. 36a) and Sakai (1939, fig. 13a). b. Longer and less arched inner orbital lobes lying farther behind the front. Only in the two smallest specimens are the lobes relatively arched. c. Reduction of spines to tubercles upon the wrist and arm; see Edmondson (1925, fig. 7c) and Sakai (1939, fig. 13c). d. The absence of a distinct cardiac ridge on the carapace. e. The different shape of the male pleopod. In the present material, there is no subterminal broadening of the appendage as shown in Edmondson (1954, fig. 36c), and the numbers and shapes of the terminal bristles differ conspicuously from those figured either by Edmondson (1954, fig. 36d) or Sakai (1939, fig. 13b). A male (25 mm., Sta. 5254, May 18, 1908, Alb.) has the ultimate segment of its male abdomen much shorter and more rounded than the remainder (see fig. 31e). Thalamita picta Stimpson Thalamita picta Stimpson, 1858, p. 39.—Alcock, 1899, p. 79.—Sakai, 1939, pp. 4138, 417, pl. 51 (fig. 2)—Edmondson, 1954, pp. 253, 263-264, figs. 35b, 36e-h.—Stephenson and Hudson, 1957, pp. 344-346, figs. 2A, 3A, pls. 4 (fig. 2), 8K, 10I.—Forest and Guinot, 1961, pp. 33-34.—Crosnier, 1962, pp. 1388-189, figs. 237-240, pl. 12 (fig. 2)—Miyake, Sakai, and Nishi- kawa, 1962, p. 128 (record only).—Garth, 1965, pp. 12-14, figs. 7, 11, 12. Goniosoma lineatum A. Milne Edwards, 1861, p. 377, pl. 35 (figs. 1, la—b).— Richters, 1880, p. 153—Lenz, 1905, pp. 360-361, pl. 48 (figs. 5, 5a-e). Thalamita gardineri Borradaile, 1902, pp. 205-206, fig. 36—Rathbun, 1911, p. 209. Thalamita alcocki de Man, 1902, pp. 646-650.—Edmondson, 1954, pp. 264-265, figs. 37a, b. Charybdis lineata (A. Milne Edwards).—Balss, 1934, p. 505. Thalamita rooseveltti Schmitt, 1939, p. 16, fig. 2. ? Thalamita investigatoris Alcock, 1899, pp. 85-86.—Alcock and Anderson, 1900, p. 8, pl. 47 (figs. 1, la).—Barnard, 1954b, p. 124, fig. 4. Marrriau.—Philippines: Sta. 5593, Mount Putri, Borneo, 4°02’40”N., 118°11’20”E., 38 fm., fine sand, Sept. 29, 1909, Alb., 1 female. Marianas: Saipan, coral heads, 1945, coll. A. H. Banner, 1 male. Marshalls: Eniwetok Reef, low tide, February 1946, F. C. Ziesen- henne, 1 male; Sta. 4442, Yonsyaran I., Bikini Atoll, May 16, 1946, NO. 3556 PORTUNID CRABS—-STEPHENSON AND REES 87 coll. Johnson, 1 male; Sta. 4449, Eniwetok I., Eniwetok Atoll, under rocks, rock flats, east side of island, May 20, 1946, J. P. E. Morrison, 1 male; H1—-121, Arno Atoll, received 1952, 1 female; Jaluit Atoll, October 1960, Rehder, 1 female. Samoa: U-01015, Schultz, 1 female. Hawaii: Kilanoa Volcano House, Otto Degener, 1 ovig. female; following from Waikiki Marine Lab., Honolulu, G. S. Mansfield; under stones near shore: Feb. 9, 1942, 1 male; Mar. 13, 1942, 4 males; Mar. 28, 1942, 1 male. Tuamotus and Societies: Sta. 29a-57, outer reef Tikahau Atoll, Maiai I., near coral rampart, Apr. 14, 1957, Bredin Exped., 1 female. MEASUREMENTS.—Males, 9-27 mm.; females, 11-20 mm.; ovig. females, 14, 17 mm. Remarxs.—Crosnier (1962) has examined Milne Edwards’ type of Goniosoma lineatum and has shown that it belongs to 7. picta. DistRIBUTION.—Examination of specimens from Clipperton I., kindly forwarded by Dr. John S. Garth, confirms that this species extends farther east than others in the genus and that 7. roosevelti is a synonym (see Garth, 1965). Previously from Madagascar, Mozambique, and Red Sea to Tuamotus, including Japan and Aus- tralia (Stephenson and Hudson, 1957; Forest and Guinot, 1961). Thalamita pilumnoides Borradaile FIGURE 32 Thalamita pilumnoides Borradaile, 1903, p. 207, figs. 38, 38a.—Holthuis, 1953, p. 8 (record only).—Crosnier, 1962, pp. 150-151, fig. 253-256. Not Thalamita pilumnoides var. gatavakensis Nobili, 1906b, p. 262; 1907, p. 384. Not Thalamita pilumnoides gatavakensis Forest and Guinot, 1961, pp. 34-36, figs. 23-25. Matertau.—Marianas: Saipan, in coral heads, 1945, A. H. Banner, 1 male, 1 ovig. female. Tuamotus and Societies: Sta. 80-57 Raiatea, Uturoa, off northern end Taoru I., branching coral, depth 2-3 ft., Apr. 29, 1957, Bredin Exped., 1 female; Sta 90a—57 Huahine, off Point Teffaao, collected from dead coral, May 2, 1957, Bredin Exped., 2 males, 1 female, 3 ovig. females; Sta. 103-57, W. of Waroa Pass, Moorea, reef, May 7, 1957, Bredin Exped., 1 female. MEASUREMENTS.—Males, 3-5 mm.; females, 4 mm.; ovig. females, 4-5 mm. Remarxs.—Forest and Guinot (1961) pointed out that Thalamita pilumnoides var. gatavakensis Nobili differed from Borradaile’s species in a number of particulars and left open the question as to whether ‘it really belonged to another species. Two males in the present collection bear pleopods, and their marked difference from those 88 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 figured by Forest and Guinot indicates clearly that they were dealing with a different species. Independently Crosnier (1962) came to the same conclusion after reference to Borradaile’s type and has re- described the present species. Apart from the pleopods, the following obvious features separate the present specimens from T. gatavakensis: Upper surface of cheliped covered with sharp tubercles instead of small oranules; three spines on inner side of upper surface of the hand of the cheliped, instead of two; and relatively shorter basal antennal joint with about four as | against six granules. WA Ficure 32.—Thalamita pilumnoides Borradaile, male, Tuamotus and Societies, Sta. 90a-57, Bredin: a, pleopod; b, abdomen. (Scales=a, 0.5 mm.; b, 1 mm.) The male abdomen (fig. 325) of the present specimen has the penul- timate segment much broader than long, and the sides are less sinuous: than in Crosnier’s figure. The ultimate segment is broader than. long in the present specimen, against as long as broad in Crosnier’s. The male pleopod is smoothly curved, with two conspicuous bristles | on the outer surface some distance behind the tip. In addition, near the tip there is a small spinule, and on the tip there are two bristles. | These additional structures are absent from Crosnier’s figure (1962, fig. 256). The male (5 mm.) from the Marianas has three anterolateral teeth instead of the usual four on each side. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 89 DistrRiBuTION.—Madagascar, Maldive-Laccadive Archipelago, and Marianas. Thalamita poissonii (Audouin and Savigny) Portunus poissonii Savigny 1817, pl. 4 (figs. 3 [1-2], 5 [1-2].)—Audouin, 1825, p. 84 [fide Stephensen, 1945, and Crosnier, 1962]. Thalamita poissonit (Audouin and Savigny).—de Man, 1880, p. 181.—Alcock, 1899, p. 81.—Borradaile, 1903, p. 201.—Nobili, 1906b, p. 206.—Rathbun, 1911, p. 208.—Klunzinger, 1913, p. 263, pl. 4 (figs. 12a—b).—Balss, 1924, p. 4. Thalamita potssoni (Audouin and Savigny).—Crosnier, 1962, pp. 116-117, figs. 183, 188, 192, pl. 10 (fig. 1). Not Thalamita poissoniit Sakai, 1939, pp. 423-424, text-fig. 17, pl. 85 (fig. 2) (=T. pseudopoissoni, new species). MartertaAu.—East Africa: Ras Banas, Red Sea, 1950, Sozon Vatikiotis, 2 males (86 mm., 45 mm.). Marshalls: S-46-44, dredged 180-200 ft., Bikini Lagoon, Mar. 29, 1946, L. P. Schultz, 2 females (8 mm., 19 mm.). REMARKS.—See under T. pseudopoissone. DistripuTION.—Madagascar, Red Sea and Suez, Laccadive Is., and Ceylon. Thalamita prymna (Herbst) Cancer prymna Herbst, 1803, pp. 41-42, pl. 57 (fig. 2). Thalamita prymna (Herbst).—H. Milne Edwards, 1834, p. 461.—Alcock, 1899, pp. 78-79.—Sakai, 1939, pp. 413, 416, pl. 51 (fig. 1) —Stephenson and Hudson, 1957, pp. 346-349, figs. 2R, 3R, pls. 4 (fig. 3), 8L, 9E.—Sankarankutty, 196la, p. 107.—Crosnier, 1962, pp. 136-138, figs. 234-236.—Guinot, 1962, p. 3.—Miyake, Sakai, and Nishikawa, 1962, p. 128 (record only). Materrau.—Malay Peninsula: Koh Pipedon, Siam, Mar. 10, 1925, H. M. Smith, 1 male; Kok Gao, Siam, in coral, shoal water, Sept. 24, 1928, H. M. Smith, 1 juv. Philippines: Near Sta. 5184, between Panay and Negros, surface, Mar. 30, 1908, Alb., 2 males, 1 female; Little Santa Cruz I., Zam- boanga, Mindanao, soft coral, marginal reefs, May 26, 1908, Alb., 1 female; Point Jarnelo, Luzon, 150 ft. seine, July 13, 1908, Alb., 1 male, 2 females; Sablayan, Mindanao, shore work collection, tide pool, Dec. 13, 1908, Alb., 1 male, 1 female, 1 ovig. female; Batan I., tide pool, June 5, 1909, Alb., 2 males, 1 female; Canino I., near Daet, June 15, 1909, Alb., 3 males, 1 female, 1 ovig. female; Batan I., July 22, 1909, Alb., 1 male, 1 ovig. female; Pilas I., tide pool, Sept. 12, 1909, Alb., 1 male. Australia: Arnhem Land Expedition, M48-21, reef at Yirrkala, NW. of Cape Arnhem, Aug. 12, 1948, R. R. Miller, 1 ovig. female. Melanesia: Vila Efate, New Hebrides, Mar. 27, 1929, Herre col- lection, 1 male; Waigin, June 6, 1929, Herre collection, 1 female; Can “D,” presumably New Guinea, received December 1944, Meyer, 1 ovig. female; as above, Can “‘K,’’ 1 male. 90 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Palau Is.: Palau I., Sta. 125-1187, about 1% mi. NE. of Ngabadon- eu, 7°17/36’"N., 134°21’42"E., 17 fm., Aug. 24, 1955, GVF, 1 female. Marianas: Tumon Bay, Guam, Sta. 80/x-30, N.M.R.V. No. 2. June 27, 1945, 1 female; Agfayan Bay, Guam, 145/RHB-150, July 7, 1945, R. H. Baker, 1 male, 1 female; Tumon Bay, Guam, Sta. 147-RHB-153, July 10, 1945, R. H. Baker, 1 male; Tumon Bay, Guam, 174/RHB/153, July 10, 1945, R. H. Baker, 1 male; Guam, December 1945, D. G. Frey, 1 male; Oca Pomt, Guam, January 1946, D. G. Frey, 1 male. Japan: Akune, Alb., 1 male, 1 female; Tanego Shima, 1906, Alb., 2 males, 4 females; Okinawa, 1906, Alb., 2 males. Samoa: No. 14, Apia, outer reef, 1902; Alb., 1 male; Apia, at mouth of river, June 1, 1902, Alb., 1 female; Apia, outer reef, June 27, 1902, Alb., 5 males, 2 females, 2 ovig. females; Apia, outer coral reef at low tide, July 1, 1902, Alb., 1 ovig, female; Apia, outer reef, July 1902, Alb., 2 males, 1 female, 1 ovig. female; Pago Pago, no. 10, August 1902, Alb., 1 male, 1 female. M&ASUREMENTS.—Males, 14-62 mm.; females, 19-51 mm.; ovig. females, 36-54 mm.; juv., 19 mm. ReEMARKS.—In unworn specimens, the basal antennal joint bears about five sharp teeth or spines, but with progressive wear these appear to be reduced first to two teeth and eventually to none. The 2-toothed forms key out with Stephenson and Hudson (1957, p. 316, couplet 8) to T. tenuipes Borradaile, 1902, which is possibly a syno- nym. Forms without teeth may key out as T. picta, but in this case the basal antennal joint is never acute as in 7. picta and rarely com- pletely smooth. The nonovigerous female (42 mm. from Apia, Samoa, outer reef, July 1902) has only four anterolateral teeth on the right side, the second and third of those normally present being fused. | Disrripution.—Delagoa and Red Sea to Marshalls, including | Japan and Australia. | Thalamita pseudopoissoni, new species Figure 33; PLATE 8B Thalamita poissonii Sakai, 1939, pp. 423-424, fig. 17, pl. 85 (fig. 2). not Portunus poissonit Savigny, 1817, pl. 4 (figs. 3 [1-2], 5 [1-2]). Audouin, 1825, p. 84 [fide Stephensen, 1945]. | Not Thalamita poissoni Stephensen, 1945, pp. 131-136, figs. 32A—H. Mareriau.—Philippines: Sta. 5159, Tinakta I., Sulu Archipelago, | Tawitawi Group, 5°11'50”N., 119°54’E., Feb. 21, 1908, Alb., 1| male (8 mm.). Palau Is.: Sta. 125-1187, about 1% mi. NE. of Ngabadongu, — @ i486", 134°21’E., 17 fm., Aug. 24, 1955, GVF, 1 male (8 mm.— | NO. 3556 PORTUNID CRABS—STEPHENSON AND REES OQ] holotype—cat. no. 112-233), 2 ovig. females (6 mm., 7 mm.), 1 juv. (5 mm.). Uncertain locality: Bearing two contradictory labels as follows: “Nagasaki” and “Sta. 5254, Gulf of Davao, Linao Point, 7°05’42”N., 125°39'42”K., 21 fm., sand, coral, May 18, 1908, Alb.,” 1 male (10 mm.), 1 female (18 mm.). The male, which is the figured speci- men, would have been selected as a holotype except for locality uncertainty. Ficure 33.—Thalamita pseudopoissoni, new species, male paratype, Philippines, Sta. 5254, Alb.: a, pleopod; b, pleopod tip, upper surface; c, abdomen. (Scales=a, c, 1 mm.; b, 0.5 mm.) Description.—Front: Bilobed, median notch varying from small, but distinct (18 mm. female; 7, 6 mm. ovig. females) to barely detectable (10 mm. male, 6 mm. juv.). Near lateral margins each lobe slightly concave, then terminating abruptly. Inner orbital lobes short and barely arched. Anterolateral teeth: Five. First stoutest, fourth distinctly the smallest, and fifth sharpest and more protruding. Carapace: Central area typically smooth and shining, lateral and frontal areas pilose. All normal carapace ridges present excepting anterior mesobranchials; no central interruption in mesogastric; an indistinct interruption in cardiac. In 8 mm. male, posterior meso- branchials not visible. 92 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Basal antennal joint: Much shorter than orbit, bearing short inconspicuous crest composed of fused granules. Chelipeds: Typically equal in females and with right stouter than left in males. Anterior border of arm bearing two spines and tubercle, or three spines. Wrist with usual spines, that on inner margin being unusually long, about two-fifths length of palm. Hands coarsely granular on upper surface, under surface with squamiform markings (sometimes faint). Upper surface with indis- tinct inner and outer carinae, each bearing a spine somewhat distal to center and terminal tubercle. Outer surface with indistinct upper and distinct central and lower carinae, uppermost being the most granular. Fingers short, stout. Fifth leg: Merus short (length about twice breadth). Propodus without posterior spines. Male first pleopod: Short, stout, tip flared. No subterminal arma- ture on inner side (paratype) or only single sharp bristle showing from under row (holotype). Outer subterminal armature an_ elongate sparse row of long, thin, forwardly directed bristles concentrated just behind tip. Under surface with transverse row of two to four short sharp bristles just behind membrane, and general but sparse distribu- tion of long forwardly directed bristles continuous with those of outer side. Remarxs.—Differs from T. poissonit markedly in the form of the male pleopods. In T. poissonii the tip is recurved, and the outer subterminal armature consists of long stout spines almost at right angles to the appendage. The differences in the general facies are: Chelipeds are more granular and spinous with clearly recognizable carinae on outer surface of hand; frontal lobes have characteristic sinuous outlines; and propodus of the fifth leg is without spines. The male pleopod greatly resembles that of JT. sezlobata, but this species differs obviously in its 4-lobed front. DisTRIBUTION.—Previously Japan. Thalamita quadrilobata Miers Thalamita quadrilobata Miers, 1884, pp. 539-540, pl. 48 (figs. B, b). Alcock, 1899, pp. 84-85.—Stephenson and Hudson, 1957, pp. 349-350, figs. 2G, 3G, pls. 4 (fig. 4), 8M, OF. Marertau.—Philippines: Nearougas I., shore, coral head, Feb. 10, 1908, Alb., 1 female. Melanesia: An-ki, Malaita, Solomon Is., Apr. 23, 1929, Herre collection, 1 ovig. female. Palau Is.: Palau Is., Oct. 13, 1933, Herre collection, 1 ovig. female. NO. 3556 PORTUNID CRABS—STEPHENSON AND REES 93 Tuamotus and Societies: Stas. 56-57, 57-57, Bora Bora, dredged, Apr. 25, 1957, Bredin Exped., 1 female. MEAsuREMENTS.—Females, 27-33 mm.; ovig. females, both 20 mm. DistriBuTIoN.—Although widespread (Seychelles, Andamans, Gil- berts, and Australia), apparently not common. Thalamita sexlobata Miers Thalamita sexlobata Miers, 1886, pp. 196-197, pl. 16, figs. 2a, b, e-—Henderson, 1893, pp. 373-374.—Alcock, 1899, pp. 87-88.—Stephensen, 1945, pp. 136-137, figs. 32C, D.—Stephenson and Hudson, 1957, pp. 350-352, figs. 2B, 3B, pls. 5 (fig. 1), 8N, 10K.—Crosnier, 1962, pp. 117-118, figs. 195-198. Thalamita sexlobata var. plicatrifrons de Man, 1902, pp. 651-653. Marertau.—Philippines: Sta. 5159, Tinakta I., Sulu Archipelago, Tawitawi Group, 5°11’50”N., 119°54’E., Feb 21, 1908, I I ? >] fe} y (Laurie), male, Philippines, Guijulugan, Albatross. (Scale 1 mm.) PROC. U.S. NAT. MUS. VOL. 120 STEPHENSON AND REES—PLATE 3 A, Portunus cf. rubromarginatus, juv., Japan, Sta. 5081, Albatross. B Edwards), form I, ovig. female, Philippines, Sta. 5157, Albatross. (Scale 1 mm.) , P. rugosus (A. Milne PROC. U.S. NAT. MUS. VOL. 120 STEPHENSON AND REES—PLATE 4 Portunus rugosus (A. Milne Edwards), form I: A, female, Philippines, Sta. 5182, Albatross; B, male, Philippines, Sta. 5159, Albatross. (Scale 1 mm.) PROC. U.S. NAT. MUS. VOL. 120 STEPHENSON AND REES—PLATE 5 B Portunus rugosus (A. Milne Edwards): A. form I, juv., Philippines, Sta. 5158, Albatross; B, form IT, male, Philippines, Sta. 5159, Albatross, (Scale 1 mm.) PROC. U.S. NAT. MUS. VOL. 120 STEPHENSON AND REES—PLATE 6 A, Portunus spiniferus, new species, holotype. B, P. tuberculosus (A. Milne Edwards), Philippines, Sta, 5159, Albatross. (Scale 1 mm.) PROC. U.S. NAT. MUS. VOL. 120 STEPHENSON AND REES--PLATE 7 A, Thalamita imparimana Alcock, Philippines, Sta. 5131, Albatross. B, T. multispinosa, new species, holotype. (Scale 1 mm.) PROC. U.S. NAT. MUS. VOL. 120 STEPHENSON AND REES—PLATE 8 A, Thalamita philippinensis, new species, holotype. B, T. pseudopotssoni, new species, holotype. (Scale 1 mm.) PROC. U.S. NAT. MUS. VOL. 120 STEPHENSON AND REES PLATE 9 Thalamita spinimera, new species, holotype. (Scale 1 mm.) ee sy i Se nen Ri ome Proceedings of the United States National Museum SMITHSONIAN INSTITUTION . WASHINGTON, D.C. Volume 120 1967 Number 3557 CLASSIFICATION OF CULEX SUBGENUS CULEX IN THE NEW WORLD (DIPTERA: CULICIDAE)! By Ravpu A. Bram Department of Entomology A synopsis of the mosquitoes of the world by Stone et al. (1959) with supplements by Stone (1961, 1963) listed 63 species in the Culex subgenus Culex from the New World. The present study assigns 61 species to the subgenus and lists 3 unrecognized species. In this study four species are described as new, five species are revalidated, one subspecies is elevated to species rank, and 11 species are placed in synonymy. This study would not have been possible without the enthusiastic guidance and support received from several outs tanding culicidologists. I take pleasure in extending most sincere gratitude and thanks to the following persons: Dr. William E. Bickley, Professor and Head, Department of Entomology, University of Maryland; Dr. Alan Stone, Entomology Research Division, Agricultural Research Service, United 1 Modified from a doctoral dissertation, submitted to the University of Mary- land, June 1964, and supported in part by the University of Maryland, Agricultural Experiment Station, project H-80. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 States Department of Agriculture; and Dr. Oswaldo P. Forattini, Professor Associado, Departamento de Parasitologia, Universidade de Sao Paulo. I am indebted to the followmg individuals who ma- terially contributed to this study by supplying specimens for examina- tion or generously sharing their knowledge: Dr. P. F. Mattingly, Dr. T. H. G. Aitken, Dr. M. Bates, Dr. T. Belterman, Dr. O. H. Casal, Dr. E. I. Coher, Dr. R. R. Correa, Dr. P. Cova Garcia, Dr. L. Vargas, and Dr. J. F.G. Clarke. I wish to dedicate this study to my wife. The major portion of material examined during the course of this study is housed in the U.S. National Museum (USNM). Additional specimens from collections in the following institutions were also consulted: Trinidad Regional Virus Laboratory, Trinidad (TRVL); Instituut Voor Tropische Hygiene en Geographische Pathologie, Amsterdam (ITHGP); Instituto National de Microbiologia, Buenos Aires (INM); Servigo De Erradicagio da Malaria e Profilaxia da Doenca de chagas, SAo Paulo (SEMPDC); Division de Endemias Rurales, Maracay (DER); Universidade de Sio Paulo, Sao Paulo (USP); Centro Ecuatoriano de Investigaciones Entomologicas (lo- cated in the USNM); and Instituto de Salubridad y Enfermedades Tropicales, Mexico, D. F. (SET). Since this manuscript was originally submitted for publication, Forattini (1965) has published a monograph of the culicine mosquitoes of the new world utilizing, in part and with the author’s permission, the classification of Bram (1964). Unfortunately, Dr. Forattini was not aware that the present manuscript had not yet been published in accordance with the International Code of Zoological Nomenclature and, therefore, attributed Culex alani, O. bickleyi, C. covagarciai, and C. oswaldoi to Bram (1964). Since Forattini (1965) was the first author to publish these species in accordance with the Code, the names must be credited to him. However, since Forattini saw no specimens and did not designate holotypes or type localities for these species, the original material upon which the species were based by Bram (1964) should be recognized as the type series for the species. Simi- larly, synonymy listed as new in this publication was actually first published by Forattini (1965), but reasons for proposing the synonymy are included in this paper. New World synonymy is listed under each species. Complete references may be found in Stone et al. (1959) and Stone (1961, 1963). | Zoogeography Little is known about the zoogeography of the family Culicidae. | This is due mainly to reasons enumerated by Bates (1949), who NO. 3557 SUBGENUS CULEX BRAM 3 observed that (a) the few fossil mosquitoes that are known are of little help in understanding the past history of the group; (b) the various taxonomic categories are not sufficiently defined to form a sure foundation for generalization; and (c) a map showing distribution of various species is apt to reflect the distribution of the collectors more than the distribution of the insects. We should also note that, at least in the Neotropical region, sufficient biological data is not available in many instances to permit intelligent interpretations of distributional patterns. Despite these limitations, noteworthy contributions to mosquito zoogeography have been made by Lane (1943, 1944), Bates (1949), Mattingly (1962), and Belkin (1962). The dominant mosquito genera in the New World, as determined by number of species, are Culex and Aedes. In the Nearctic region Aedes assumes the dominant role with Culex being represented by only 26 species (15 of which are both Nearctic and Neotropical). In the Neotropical region Culex predominates and is represented by 239 species according to Stone et al. (1959). In no other region is the genus Culex so diversified. In the Ethiopian region there are 104 species in the genus. The Oriental region is next in number of species with 88, followed by 33 species in the Palearctic. Within the genus Culex the subgenus Melanoconion is dominant. It is indigenous to the New World and encompasses 117 species (only eight of which are found in the Nearctic). The subgenus Culex is cosmopolitan, but also exhibits a decided concentration in the tropical regions. Only three species of the subgenus, restuans, salinarius, and tarsalis, are indigenous to the Nearctic region. Forty-nine species are indigenous to the Neotropical region, and nine species are found in both Nearctic and Neotropical regions. The following discussion treats zoogeo- graphical patterns of the subgenus Culex in the New World. The three species of Culex (Culex) native to the Nearctic region present interesting patterns, since they are related to widely distrib- uted Neotropical species. Culex restuans is found throughout North America, ranging from the Gulf of Mexico into Canada and from California to the east coast (Carpenter and LaCasse, 1955). Studies of the male terminalia indicate that it has close affinity with C. laticlasper, a species described from the Panamanian Isthmus. Another species found in the complex, C. acharistus is distributed from Columbia south to Llanquihue, Chile and eastward to Argentina and Brazil. Culex salinarius, also indigenous to the Nearctic region, occurs in the eastern United States, southeastern Canada, extends westward to Utah, and dips into Mexico (Carpenter and LaCasse, 1955). Studies of male terminalia indicate that it has close affinities with C. archegus, which is found on the northwest coast of South America, 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 and C. dolosus and C. spinosus, species found in the central portion of South America. Culex tarsalis ranges from southwestern Canada through the western, central, and southern United States and into Mexico (Carpenter and LaCasse, 1955). This species demonstrates close affinity with C. abnormalis which extends from Colombia to central Brazil. All three of these species, indigenous to the Nearctic region, demon- strate considerable extensions into Mexico but none has been reported from Guatemala southward. Thus, it appears that all three species have close affinities with, and that their ancestors may have been part of, the Neotropical fauna. ‘The affinities of these three Nearctic species with the Neotropical fauna could be explained by Belkin’s (1962) hypothesis of the intercontinental origin of new types. This hypothesis suggests that the phylad originated on an island in the intercontinental area, dispersed to both hemispheres initially, and then continued evolving independently. The two principal intercontinental areas which Belkin (1962) con- sidered as primary centers of origin are the Indo-Pacific area between Eurasia and Australia, and the American Mediterranean between the North and South American continents (ibid., vol. 2, map on p. 18). It was his contention that these areas are the most likely places for the origin and evolution of new major types of mosquitoes. Both are characteristically unstable regions where major orogenic belts of adjacent continents meet, intersect, form arcs, or otherwise have complex relationships. Thus, in the course of island formation, great environmental stress would be imposed on the surviving populations and these would be greatly reduced and isolated. Under such con- ditions there would be an ideal opportunity for the quick fixation of new adaptive types of organisms. Examination of distributional patterns in the subgenus Culex indicates that the American Mediter- ranean is probably the primary center of origin. Forty-seven of the 61 species considered in this study are found within the boundaries of this area as delimited by Belkin. Several endemic species are found on the Antillean and Bahamanian islands. Both Culex scimitar and C. sphinx have been reported only from the Bahamas and both show strong affinities in the male termi- nalia to C. nigripalpus, which is distributed throughout the southern Nearctic and American Mediterranean areas. C. duplicator is endemic to the island of Hispaniola and demonstrates affinity with C. saltanensis, a species with a distribution range from Venezuela to northern Argentina. C. finlayi has been reported only from Cuba. Although the male terminalia is somewhat unique, there seems to be some affinity with the more widely distributed C. chidesterr. C. bahamensis, a unique species demonstrating affinity with the subgenus NO. 3557 SUBGENUS CULEX—BRAM 5 Melanoconion, is restricted to the Caribbean Islands and the southern tip of Florida. The Central American isthmus also possesses endemic species. Culex delys and C. laticlasper are known only from the Panama Canal Zone. C. stenolepis is known from the state of Veracruz, Mexico, and Costa Rica; it exhibits affinities with C. thriambus, a species which extends into Colombia. C. pinarocampa has been reported from four states in southern Mexico and from Panama; however, considering our present knowledge of distributions, none of these Central American forms can definitely be considered as truly endemic. In the subgenus Culex it appears that the intercontinental islands of the American Mediterranean area did not assume the dominating role as a center of origin but, in general, were the recipients of the South American fauna. Although somewhat reduced in number of species, the Trinidadian fauna is typical of the fauna of the adjacent South American continent. The continental fauna is also reflected to some degree in the fauna of the other Carribean islands. This tendency towards a South American fauna (in contrast to a Central American, North American, or endemic fauna) is readily explained by consideration of prevailing surface winds of the Atlantic Ocean and typical tracks of tropical storms. Darlington (1957, fig. 8) outlined the course of the prevailing winds. Examination of these figures revealed that the winds and storms originate in the Atlantic Ocean and develop in a circular, clockwise direction, touching the South American continent and proceeding through the Caribbean or towards the Central American isthmus. The winds would have probably served as the primary agents of dispersal. Lane (1943) reviewed the geographical distribution of the mosquito tribe Sabethini. In this study he recognized five centers of endemism and dispersal in the Neotropical region as well as two negative areas in which no Sabethines were found (ibid., map 8, p. 425). In a sub- sequent study, Lane (1944) suggested that the tribe Anophelini also adhered to these centers of endemism and dispersal. Belkin’s Ameri- can Mediterranean area encompasses Lane’s Central American, Incasic, and Cariba centers as well as the Caribbean islands. This area is considered the primary center of dispersal in the New World. In the subgenus Culex the most prominent secondary center of dispersal corresponds generally to the Patagonian area. Species included in this area are: Culex apicinus, a highly adapted species with rather independent tendencies; C. articularis, also somewhat unique; C. brethesi and C. acharistus, species showing affinities with the restuans- laticlasper complex; C. lahillei, which shows affinity with C. foliaceus; and C. tramazayguesi, a unique species of unknown affinity. Another secondary center of dispersal might well be Lane’s negative center 4. 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Species which appear to be endemic to this area include C. foliaceus, C. oswaldoi, and C. maxi, which extends through the central plateau to northern Argentina. Lane’s center 5 (the Tupi area) is represented by four endemic species: C. carcinoxenus, C. lygrus, C. renatoi, and OC. acharistus; however, each of these species is known only from the type locality. This limited distribution is probably a reflection of the close proximity of this area to culicidologists working at the University of Sao Paulo. The Central Plateau area (or Lane’s center 3) exhibits two endemic species, C. mauesensis and C. airozai, which are also known only from their type localities. Again, this does not necessarily represent a recognizable center of endemism. Several species of Neotropical Culex (Culex) have rather broad areas of distribution which are worthy of consideration. Culex chidesterr extends from Mexico southward to northern Argentina, being re- stricted to the western half of the continent. Interestingly enough, this species has been found on the island of Jamaica but has not been reported from any of the other Caribbean islands. C. corniger is distributed from Mexico throughout the northern half of South Amer- ica and southward to Argentina on the east coast. It is also found throughout the Caribbean. C. mollis extends from Mexico south to northern Argentina and is apparently a common species. In the Caribbean, however, it has only been reported from the island of Trinidad which typically exhibits a continental fauna. C. abnormalis and C. saltanensis seem to be restricted to a central, north-south band on the South American continent. With the exception of Central America and the Caribbean islands, C. surinamensis is distributed throughout the American Mediterranean area and extends into the central plateau. C. declarator ranges from southern Texas throughout the American Mediterranean, but is limited in the Caribbean to the Lesser Antilles. It also extends eastward to the state of Pernambuco, Brazil. A very closely related species, C. bidens, is more southern in distribution ranging from central Argentina northward through the east coast and central plateau areas and extending into Venezuela. It has not been reported from the west coast of South America and is also generally absent from the American Mediterranean. Martinez Palacios (1952) did report this species from six southern and central states of Mexico. The C. coronator complex of species presents an interesting distributional pattern, which will be discussed in detail under the various species. In general, however, C. coronator and C’. usquatus extend from Texas southward to northern Argentina, but are absent from the west coast of South America and the Caribbean islands. C. usquatissimus, C. ousqua, C. camposi, and C. covagarciat are distributed in varying degrees within the American Mediterranean. These patterns, then, do not indicate distinct centers of endemism NO. 3557 SUBGENUS CULEX—BRAM 7 and distribution in the subgenus Culex as was found by Lane (1943, 1944) for the tribes Sabethini and Anophelini. Rather, the dis- tributional patterns of the various species indicate several broad faunistic areas as suggested by Belkin (1962) for the entire family Culicidae. The American Mediterranean exhibits the greatest num- ber of species of any area, as well as the largest number of endemic species. Although suggestions of subpatterns seem to exist, the general area is considered the primary center of distribution. The Patagonian area was recognized by both Lane and Belkin (in this case as the Patagonian-South Chilean area) and in the subgenus Culex demonstrates a distinctively unique fauna. In this subgenus the Patagonian area would seem to represent a compromise between the concept of the areas presented by Lane and Belkin. The remain- ing area of South America seems best considered under Belkin’s broad Neotropical area. In this area suggestions of subpatterns also exist, but sufficient data is not presently available to confirm or delimit these possibilities. In summary, examination of distributional patterns reveals several broad faunal areas of New World Culex (Culex). The entire Nearctic region exhibits only three endemic species and possesses nine more species which are also found in the Neotropical region. The Neo- tropical region may be subdivided into three faunal areas: (1) the American Mediterranean, which is considered the primary center of origin and distribution; (2) the Patagonian, which is confined to the west coast and southern tip of the continent, and which exhibits a unique fauna; and (3) the Neotropical, which includes the rest of the South American continent. These faunal areas, modified after Lane (1943) and Belkin (1962), are outlined in figure 1. Additional subpatterns within these areas are suggestive, but more distribu- tional data is required to confirm and delimit these secondary features. Terminology MALE TERMINALIA.—The terminology followed in this study is basically that of Carpenter and LaCasse (1955). The one exception to this is in the male terminalia. A wide variety of terms has been used to describe the structures of the male terminalia and few authors agree on standard terms in the subgenus Culex. Table 2 lists the terminology used by eleven different authors to describe the struc- tures of the mesosome in the Culex subgenus Culex. This table is by no means complete since, as indicated by Freeborn (1924), it is impossible to produce an accurate synonym list for the nomenclature of the different parts without consulting the original descriptions of all species and all monographic works. In respect to male terminalia, 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 NEW WORLD FAUNAL REGIONS AND AREAS BASED ON DISTRIBUTION WITHIN THE CULEX SUBGENUS CULEX Nearctic Region American Mediterraneon Area Neotropical Fregion Neotropical Dis Area Potagonian Area Ficure 1.—New World faunal regions and areas based on distribution within the Culex | subgenus Culex. NO. 3557 SUBGENUS CULEX—BRAM 9 the nomenclature of Freeborn and Bohart (1951) is utilized through- out this study. Following is a brief discussion of major morpho- logical characters of the male terminalia utilized in this study. Each structure discussed is included in diagrams of each species. Tenth sternites (XS): These structures were regarded by Chris- tophers and Barraud (1923) as the cerci united with the anal lobe, but in general taxonomic nomenclature they are recognized as sternites. In the subgenus Culex the tenth sternites are crowned with a tuft of spines and it is this character that is considered im- portant. Three basic arrangements of tufts are found and can be readily identified: (1) spines sparse and weak; (2) spines dense and rather strong; and (3) spines dense and strong apically, but with the spines on the outer lateral margins spatulate and rather scalelike in appearance. It should be noted that all species exhibiting arrange- ment ‘‘(1)’’ do not possess a leaf on the apical lobe of the basistyle; however, the reverse is not true. Ventral cornu (VC): Stone (personal communication, 1962) con- sidered this structure to represent a modification of the apical tooth of the median process of the mesosome. ‘Two rather distinct con- figurations can be recognized. Most species possess a dentiform ventral cornu, although the structure may assume a variety of shapes and sizes in relation to the teeth of the median process of the mesosome. A second form exists where the ventral cornu is very large, rounded, and exhibits rugulose outer margins. Median process of the mesosome (MP): The precise number of teeth on the median process of the mesosome varies somewhat, even within a species. Three recognizable situations, however, seem to exist: (1) the teeth are absent or are, at most, one or two in number; (2) the teeth are distinctly separated and range in number from three to approximately 10; and (3) the condition in which a large number of conglomerate teeth is present. External process (EP): This structure issues from the center of the mesosome and assumes various shapes and sizes in relation to the median process of the mesosome. The structure is present in all species of the subgenus, but is difficult to distinguish when the termi- nalia is mounted in the everted position. Basal arm of the mesosome (BA): This structure generally issues from the mesosome as a short, straight, bluntly rounded limb. In some species the structure may be completely absent, and in other species it may be bent at an acute angle so as to be directed posteriorly. Apical lobe of the basistyle (AL): There is great variation in setal arrangement on the apical lobe, as well as variations in the conforma- tion of the lobe, itself. Lane (1953) conveniently divided the sub- genus into two groups, depending on whether or not the apical lobe 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 possessed a leaf-shaped seta; however, this division seems to represent an oversimplification of the complex interrelationships of terminalic structures. The basic setal arrangement on the apical lobe appears to be: three strong, basal rods followed by a leaf-shaped seta; a straight, pointed seta; and a curved, pointed seta. Variations to this basic pattern range widely from a simple group of straight, pointed setae in Culex usquatus to elaborate modifications and setal arrangements in C. apicinus. FEMALE CHARACTERS.—Historically, adult female mosquitoes have been extremely difficult to identify accurately. This is due mainly to the limited number of distinctive characters, intraspecific varia- tion of characters, and similarity of closely related species. In the subgenus Culex primary taxonomic characters are concerned with body ornamentation and scale color patterns. In this study 13 different colorational characters have been selected and are listed in table 1. For many of the species, positively determined material was not available (due to lack of associated, individual rearings) and in these cases reference was made to the original descriptions. These characters were not incorporated into a formal key because, for most species, data on intraspecific variation are not available at the present time. ‘Terminology used in describing these colorational characters is that of Carpenter and LaCasse (1955). LARVAL CHARACTERS.—In the larval stage many stable morpho- logical structures are available for use as taxonomic characters. As in the case of female characters, the terminology utilized in describing these characters is that of Carpenter and LaCasse (1955). Morpho- logical larval characters utilized in this study are as follows: Antenna: In all but two species examined the antennae are con- stricted near the outer third, beyond the insertion of the antennal tuft. Degree of spiculation of the antennal shaft is also utilized as a taxonomic character. Head hairs: The arrangement, length, and condition of the four primary head hairs are characteristic. The following head hairs are utilized as taxonomic characters: postclypeal head hair 4; upper frontal head hair 5; lower frontal head hair 6; and preantennal head hair 7. Mentum: This structure is the strongly sclerotized and pigmented, toothed plate of the labial-hypopharyngeal complex. It is character- istic in the number and conformation of the lateral teeth. Integument: The presence or absence of spicules on the thoracic and abdominal integument is frequently diagnostic. Comb: This structure is a row or patch of enlarged specialized spicules in the middle of each side of the eighth abdominal segment. NO. 3557 SUBGENUS CULEX—BRAM 11 The general arrangement of the comb scales is frequently utilized, as well as the number and conformation of the individual scales. Siphon: The air tube of the typical nonanopheline mosquitoes; morphologically includes part of the abdominal segments VIII and IX. Several features of the siphon are utilized as taxonomic characters. Siphonal index: The ratio between the length of the siphon and the basal width. Siphonal tufts: Pairs of setae inserted on the siphon. The number, location, and condition of the siphonal tufts are diagnostic. Pecten: A paired, comblike row of spiculate teeth along the ventro- lateral border of the siphon. The number, arrangement and con- formation of individual teeth is diagnostic. L/S ratio: The ratio of the length of the siphon to the dorsal length of the saddle. This ratio was originally proposed by Colless (1957) and is used as a supplementary character in the larval key. Keys In this study emphasis has been placed on characters of the male terminalia because of the scarcity of material, particularly larval skins associated with adults, and because many species are known only from the male terminalia. Of the 61 species recognized in this study, 59 have been included in the key to male terminalia. Culex delys is known only from the female and C. pseudojanthinosoma is known only from the female and a description of the larval stage. A key to New World subgenera of the genus Culex based on the male has been constructed utilizing many of the characters suggested by Foote (1954). Due to the scarcity of reliable data, the key to larvae is extremely tentative. In 18 species the larval stage is unknown. Descriptions of the larvae in many other species are incomplete or questionable. A thorough understanding of larval morphology and affinities awaits future individual rearings. Only by this technique can correlations between immature and adult stages be conclusively established. A list of larval characters separating the subgenus Culex from other New World subgenera of the genus was prepared instead of a formal key, because some difficulty was encountered in separating the subgenera Carrollia and Microculez. Morphological and colorational characters of adult females in the subgenus Culex are extremely difficult to differentiate. As pointed out by Lane (1953), a key utilizing adult females serves only to approximate species because the characters are variable. In this study few species were represented by a suitable sample of positively determined specimens, and the amount of material obtained from 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 individual rearings was very limited. Original descriptions of adult females are often vague and different authors emphasize different morphological and colorational aspects. Therefore, no attempt was made to construct a formal key incorporating adult female characters. However, a table of salient female characters has been prepared as table 1. Key to Subgenera of New World Culex Based on the Male 1. Tenth sternite crowned with a tuft of predominently pointed setae; spat- ulate setae may be present on the outer lateral margins... .... .2 Tenth sternite crowned with arow of spatulatesetae ......... .4 2-*"Tenth sternite without a basalvarm:, 0°... so. We i. he an eee Tenth sternite with a-basal arm’ present . 2°. 4)): .)). ee. a Gaolex 3. Ventral cornu present on themesosome. .......... . Lutzia Ventral cornu absent. . . . . Neoculex 4. Palpus longer than the enon ce ane proboscis by x Tere fee length of the last palpal segment ...... eR . . . . . Melanoconion Mochlostyrax Palpus no longer than the length of the proboscis ...........8 5. Abdomen without basolateral metallic spots ............ 2.6 Abdomen with basolateral metallic spots. ...... . . . Carrollia 6: Mesosomal plate without retrorse teeth -.). 9: - .). 5 3) 37S eee Mesosomal plate with a pair of retrorse teeth. . . .. . . Microculex 7. Apical lobe of the basistyle extended into a single, medially placed, thumb- like projection ... . . . . . .Eubonnea Apical lobe of the Sessile not as abone: divided ito a distal and proximal BeCtiON 2. ys aw. sae: By os. go ba ee ee ow. foul: plies ole Se ee CMR Key to Species of New World Culex (Culex) Based on Male Terminalia Ie Apical lobe of the basistyle without a broadened leaf ...... .2 Apical lobe of the basistyle with at least one broadened leaf. . . . 18 2. (1) Tenth sternite crowned with a dense tuft of strongspines .... .3 Tenth sternite crowned with a sparse tuft of finespines ... 4 oy) (2) Dististyle enlarged medially; apical lobe of the basistyle enieaded into a tuberculiform projection terminating in a strong apical rod and possessing a distal seta and a basal seta .. . . habilitator Dististyle not enlarged medially ; apical lobe of the basistyle bearing only three strong, obtuse rods and several normal setae . tramazayguesi 4. (2) Median process of the mesosome bearing 10 of more conglomerate Geeth eps. oe Pe ee Median process of the mesosome eaine ies hoy 10 distinctly sep- arated teeth. . ... ‘ Peewee) 5 (4) Apical lobe of the Beaistyle Santas Ae ppeare ana one hooteed rod, as well as additional setae; ae third of the dististyle minutely annulate. °. ... . . . . . bahamensis Apical lobe of the ecistayle noe as prone: apical third of the dististyle not annulates..) nie . ge 6G. (5) Ventral cornu pointed, Satan Hiei Outer escnaine . hae Ventral cornu rounded, rugulose on outer margins . . maracayensis 7. (6) Tenth sternite with pilosity on the lateral margins ....... .8 Tenth sternite without pilosity on the lateral margins . . bonneae NO. 3557 8. 10. on. 12. 13. 14. 15. 16. ie 18. no: (7) (10) (9) (13) (14) (18) (16) (1) (18) SUBGENUS CULEX—BRAM 13 Ventral cornu pointed and bent at the outer third so as to be directed posterolaterally; external process evenly tapered to an obtuse termination ... . . . . saltanensis Ventral cornu serrate commie exieneal mrodess broad at the basal %, then sharply indented on the lateral edge and tapered; endemic to the island of Hispaniola ... . . . . . . duplicator Ventral cornu more or less dentiform iad Shore, not distinctly sepa- rated from the teeth of the median process . . . 49S als es HO) Ventral cornu long and slender, directed laberally: end ciepmrele separated from the teeth of the median process (coronator group) .13 Apical lobe of the basistyle with two sections; the basal section with three rods, the distal section with approximately six setae. .... .11 Apical lobe of the basistyle with about six subequalsetae. . . . . .12 Ventral cornu clavate, located just above the teeth of the median process: . 4% . . . . . brevispinosus Ventral cornu deneieora leeated loser o the teeth of the median process: ») . . . surinamensis Ventral cornu ergo fal cieninet: medern THOUGRE bearing about four teeth; basal process directed posteriorly ... . . . . oswaldoi Ventral cornu small and dentiform; median Peauesa bearing about eight teeth; basal process directed anteriorly . . . se. Goines A group of short setae located at the apex of the Basicowie Sa ke 1 A group of long setae located at the apex of the basistyle; these setae reach to at least the midpoint of the dististyle . ....... 16 Apical lobe of the basistyle undivided. . .. . bee 5 ONS Apical lobe of the basistyle divided into a large aeesinnel lobe and a small distal tubercle; the proximal lobe possesses a group of approxi- mately 10 subequal setae; the distal tubercle possesses four or five subequal setae . . . . . . Ousqua Apical lobe of the peaiseele Boneeseine re or fthres rode basally fol- lowed by 5tol5setae ... . . coronator Apical lobe of the basistyle Entended cio! a pchumblike projection possessing the following appendicles: one short, very broad rod, two long prominently hooked rods, and several subequal, hooked setae. covagarciai Basistyle with a small tubercle located distad of the apical lobe and possessing from one to five setae .. . areas Pell L( Basistyle without a small tubercle distad of the apical ioe apical isbe of the basistyle possessing from 10 to 15 setae. . . . . usquatus Apical lobe of the basistyle possessing three rods and one seta; small tubercle distad of the apical lobe possessing four or five setae. usquatissimus Apical lobe of the basistyle possessing approximately 10 setae; small tubercle distad of the Bora lobe ao one or two strong Setaen 2) Le . . . .camposi Median process of them mesosome yprojected ito a erode extension upon which is located a sphincter-form ring. ....... . . finlayi Median process of the mesosome not asabove. . . . Be 8, OSV APG Ventral cornu absent; teeth of the median process spec or one reduced in size and number; external process very broad and promi- nent, slightly curved laterally. ... . aaa ee 0) ence cornu present, dentiform or paealonel ne pounded teeth of the median process not reduced in size; external process normal. . . 24 14 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34, (19) (20) (21) (22) (19) (24) (25) (24) (27) (28) (28) (30) (31) (27) (33) PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Basal arm of the tenth sternite long and curved. ....... .21 Basal arm or the tenth sternite short and straight. . . . . . pipiens Apical lobe of the basistyle with three rods, a sin a hooked seta, and a straight seta . . RS ee Apical lobe of the Dacicigle ah Aone deales a eae antl a straight Setaenrs soars Leo spe Drethest Apical third of ihe Aistatyle wachoue nee Sa den yh ER eee Apical third of the dististyle with minute annulations (this character is best seen under oil immersion or phase contrast) . . . acharistus Hooked accessory seta on apical lobe of the basistyle unusually strong and curved; median process of the mesosome with about two small, lightly sclerotized denticles . .. . ... . . .laticlasper Hooked accessory seta on apical lobe of {he basistyle not unusually strong, gently curved; median process of the mesOsome completely without teeth ... . . . .restuans Apical lobe of the soa (ea —_—_—— Bs Ficure 29.—C. saltanensis, Bahia, Brazil, USNM RB62 563: a, basistyle and dististyle; b, mesosome. C. scimitar, New Providence, Bahama Is., USNM RB62 720: c¢, basistyle and dististyle; d, mesosome. 221—521— 67 8 100 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 inserted on the siphon beyond the pecten. Pecten with about 12 teeth on the basal third of the siphon: each tooth with coarse barbs on one side. Anal segment completely ringed by the saddle, which is spiculate dorsolaterally. MATERIAL EXAMINED.—The type series and nine other adult males and associated terminalia from the Bahama Islands. DistriBuTION.—Reported from the Bahama Islands and Cuba. Culex (Culex) secutor Theobald, 1901 FIGURES 30a, b Culex (Culex) secutor Theobald, 1901, p. 321. Culex (Culex) lamentator Dyar and Knab, 1906b, p. 219. Culex (Culex) quasisecutor Theobald, 1907, p. 398. Culex (Culex) towert Dyar and Knab, 1907b, p. 13. SystemMatics.—This species exhibits a rather generalized mesosome in the male terminalia, but the arrangement and distribution of the appendicles on the apical lobe of the basistyle is diagnostic. SALIENT CHARACTERS.—Adult female: See table 1. Male terminalia: Basistyle conical, slightly longer than twice the basal width; clothed with fine setae in addition to the normal, long setal pattern. Apical lobe of the basistyle prominent, undivided, and also clothed with fine setae. Appendicles of the apical lobe as follows: basal rod strong, straight, and with a fine hook at the apex; median rod longer and broader than basal rod, prominently hooked; apical rod longer than, but not as broad as the median rod, also prominently hooked; two short, straight, fine setae next to the apical rod; leaf moderate in size, bluntly rounded, with a few prominent longitudinal striations in the center; two accessory setae present, one strong and prominently hooked, the other more slender, long, and gently curved. Dististyle normal. Tenth sternite crowned with a dense tuft of short, pointed spines; basal arm moderate in length, strongly curved termi- nally; three cercal setae present in a compact group. External process eradually tapering to a blunt point; reaching to about the ventral cornu. Ventral cornu dentiform, somewhat larger than the teeth of the median process. Median process with about five distinct, sharply pointed teeth. Basal process broad and sharply bent at the apical third so that the pointed apex is directed posteriorly. Larva: Antennal tuft located in a constriction near the outer third; antennal shaft spiculate basally. Postclypeal head hair 4 short, single; upper and lower frontal head hairs 5 and 6 4-branched, long, and barbed; preantennal head hair 7 multiple, long, and barbed. Mentum with about 21 teeth, the apical tooth broader and larger than the lateral teeth. Body glabrous. Comb with numerous scales in a patch; each scale rounded apically and fringed with subequal spinules. NO. 3557 SUBGENUS CULEX—BRAM 101 Siphonal index about 4.0; six multiple siphonal tufts on the siphon, the basal tuft inserted within the pecten. Pecten with about 13 teeth on the basal third of the siphon, the apical tooth removed from the other pecten teeth; each tooth with coarse barbs on one side. Anal segment completely ringed by the saddle, which is spiculate distally. MATERIAL EXAMINED.—Three adult males and associated terminalia from Puerto Rico and one from Jamaica. DIstTRIBUTION.—Reported from Jamaica (the type locality), His- paniola, Puerto Rico, the Lesser Antilles, and Venezuela. Culex (Culex) sphinx Howard, Dyar, and Knab, 1912 Figures 30c, d Culex (Culex) sphinx Howard, Dyar, and Knab, 1912 [1913], p. 134. Systrematics.—This species demonstrates close affinity with Culex nigripalpus but is distinctive in the arrangement and conformation of the appendicles on the apical lobe of the basistyle. SALIENT CHARACTERS.—Adult female: See table 1. Male terminalia: Basistyle conical, about two and a half times longer than the basal width; clothed with fine spicules in addition to the normal, long setal pattern. Apical lobe of the basistyle promi- nent in the area of the rods, but somewhat flattened in the area of the leaf and accessory setae; also clothed with fine spicules. Appen- dicles of the apical lobe as follows: basal rod straight, strong, and pointed; median rod broader and longer than the basal rod, termi- nating in a gentle hook; apical rod longer and broader than the median rod, also terminating in a gentle hook; two subequal, strong, straight, pointed accessory setae located between the apical rod and the leaf; leaf rather narrow and truncate, about as long as the accessory setae. Dististyle normal. Tenth sternite crowned with a dense tuft of short, pointed spines apically and six to nine spatulate, scalelike spines on the lateral outer margins; basal arm short and gently curved; about six cercal setae present in a compact group. External process some- what curved and bluntly rounded; not reaching the ventral cornu in length. Ventral cornu distinct from the teeth of the median process, truncate and serrate. Median process with about six distinct, sharply pointed teeth. Basal process reduced to a bulbous expansion of the mesosome; not distinct. Larva: Antennal tuft located in a constriction near the outer third; antennal shaft spiculate basally. Postclypeal head hair 4 short, single; upper and lower frontal head hairs 5 and 6 three or four branched, long, barbed; preantennal head hair 7 multiple, long, and barbed. Mentum with about 17 teeth, the apical tooth broader and larger than the lateral teeth. Thorax finely spiculate, abdomen glabrous. Comb with numerous scales in a patch; each scale rounded 102 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 apically and fringed with subequal spinules. Siphonal index about 3.0 to 3.5; three double or triple siphonal tufts inserted on the siphon. Pecten with about 20 fine teeth, restricted to the basal third of the siphon; each tooth with coarse barbs on one side. Anal segment spiculate, completely ringed by the saddle. WN My i Ay) Ni phen \\ // Wy. Ficure 30.—C. secutor, Mayaguez, P.R., USNM RB62 356: a, basistyle and dististyle; b, mesosome. C. sphinx, New Providence, Bahama Is., USNM RB62 381: ¢, basistyle and dististyle; d, mesosome. NO. 3557 SUBGENUS CULEX—BRAM 103 MATERIAL EXAMINED.—The lectotype male terminalia and _ six other male specimens from the Bahama Islands. DistriButTion.— Reported only from the Bahama Islands. Culex (Culex) spinosus Lutz, 1905 Figures 3la, b Culex (Culex) spinosus Lutz, 1905, p. 26. SystpeMatics.—This species is a member of the salinarius complex. The primary distinguishing characteristic is the large, distinctive basal process. The ventral cornu and external process are also useful diagnostic characters. SALIENT CHARACTERS.—Adult female: See table 1. Male terminalia: Basistyle conical, slightly longer than twice the basal width; clothed with fine setae in addition to the normal, long setal pattern. Apical lobe of the basistyle prominent, undivided, and also clothed with fine setae. Appendicles of the apical lobe as follows: basal rod strong, straight, and rounded terminally; median rod broader and longer than basal rod and terminating in a prominent hook; apical rod not as broad as, but longer than the median rod, also terminating in a prominent hook; leaf moderate in size, obovate, exhibiting distinct longitudinal striations; two accessory setae present, one shorter than the leaf and prominently hooked terminally, the other very long, pointed, and gently curved. Dististyle normal. Tenth sternite crowned with a dense tuft of short, pointed spines; basal arm moderate in length, gently curved; two or three cercal setae present in a compact group. External process gradually tapering to a blunt point; exceeding the ventral cornu in length. Ventral cornu dentiform, very similar in size and shape to the teeth of the median process. Median process with about four to six sharply pointed, distinct teeth. Basal process sharply pointed distally, acutely bent so as to be directed posteriorly; robust basally, but long and slender apically; reaching almost to the ventral cornu. Larva: After Lane (1953). Antennal tuft located in a constriction near the cuter third; antennal shaft spiculate. Postclypeal head hair 4 short, single; frontal head hairs 5, 6, and 7 multiple, long, barbed. Body glabrous. Comb with numerous scales in a patch; each scale rounded apically and fringed with subequal spinules. Siphonal index 4.5 to 6.0; four triple siphonal tufts inserted on the siphon beyond the pecten. Pecten with about 12 teeth on the basal third of the siphon; each tooth without coarse barbs on one side. Anal segment com- pletely ringed by the saddle, which is densely spiculate on the entire surface, 104 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 MATERIAL EXAMINED.—Three adult males and associated terminalia from Colombia and Venezuela. Distrisution.—Reported from Brazil. The author has seen speci- mens in the U.S. National Museum collection from Lagunillas, Venezuela, and from Colombia. Culex (Culex) stenolepis Dyar and Knab, 1908 FIGURES 3le, d Culex (Culex) stenolepis Dyar and Knab, 1908, p. 60. Systamatics.—Culez stenolepis demonstrates relationship to C. interrogator, O. peus, and C. thriambus. It may be distinguished from the first two species by the number and arrangement of the appendicles on the apical lobe of the basistyle. It is distinct from C. thriambus in the conformation of the external process. SALIENT CHARACTERS.—Adult female: See table 1. Male terminalia: Basistyle conical, about twice as long as the basal width; clothed with fine setae in addition to the normal, long setal pattern. Apical lobe of the basistyle somewhat flattened, undivided, and also clothed with fine setae. Appendicles of the apical lobe as follows: basal rod strong, straight, and pointed; median rod broader and longer than the basal rod and terminating in a gentle hook; apical rod longer than, but not as broad as the median rod, also terminating in a gentle hook; leaf moderate in size, obovate, with several prominent, longitudinal, centripetal striations; one long, pointed, gently curved accessory seta present. Dististyle normal. External process gradually tapering to a point, reaching to about the ventral cornu in length. Ventral cornu large, rounded, and rugulose. Median process with about seven sharply pointed, sub- equal teeth. Basal process short, straight, broad, and bluntly rounded. Larva: Antennal tuft located in a constriction near the outer third; antennal shaft spiculate basally. Postclypeal head hair 4 short, single; frontal head hairs 5, 6, and 7 multiple, long, and barbed. Mentum with about 23 teeth; the apical tooth broader and longer than the lateral teeth; the subbasal lateral teeth longer than the other lateral teeth. Body glabrous. Comb with many scales in a patch; each scale rounded apically and fringed with subequal spinules. Siphonal index about 3.5; five pairs of siphonal tufts inserted beyond the pecten, the basal four tufts double, the apical tuft multiple. Pecten with about 10 teeth, restricted to the basal third of the siphon; each tooth with several coarse barbs on one side. Anal segment completely ringed by the saddle, which is sparsely and finely spiculate distally. MATERIAL EXAMINED.—Seven adult males and associated ter- minalia from Mexico. The lectotype male terminalia was also studied. DistriBuTION.—Reported from Mexico and Costa Rica. NO. 3557 SUBGENUS CULEX—BRAM 105 a (ge AL 0.1mm. Figure 31.—C. spinosus, Lagunillas, Venezuela, USNM RB62 428: a, basistyle and disti- style; b, mesosome. C. stenolepis, Cordoba, Mexico, USNM RBé2 714: c, basistyle and dististyle; d, mesosome. Culex (Culex) surinamensis Dyar, 1918 Ficures 32a, b Culex (Culex) surinamensis Dyar, 1918b, p. 121. Systrematics.—This species belongs to the group which lacks a leaf on the apical lobe of the basistyle. It is distinguished from its nearest relative, Culex brevispinosus, by possessing a dentiform ventral cornu. SALIENT CHARACTERS.—Adult female: See table 1. Male terminalia: Basistyle conical, slightly longer than twice the basal width; clothed with a sparse covering of very fine spicules in 106 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 addition to the normal, long setal pattern. Apical lobe of the basistyle prominent, undivided, also clothed with fine spicules. Appendicles of the apical lobe as follows: basal, median, and apical rods subequal in length, very fine, pointed, and gently curved terminally; beyond the apical rod are three or four fine, straight, and pointed accessory setae; leaf absent. Dististyle normal. Tenth sternite crowned with a sparse tuft of short pointed spines; basal arm moderate in length, distinctly curved; four cercal setae present. External process gradually tapering to a blunt point; reaching to the ventral cornu in length. Ventral cornu dentiform, similar in shape, but slightly larger than the teeth of the median process. Median process with about five distinct, sharply pointed teeth. Basal process short, straight, and bluntly rounded. Larva: Antennal tuft located in a constriction near the outer third; antennal shaft spiculate basally. Postclypeal head hair 4 short, single: frontal head hairs 5, 6, and 7 multiple, long, and barbed. Mentum with about 11 very broad teeth, the apical tooth larger than the lateral teeth. Thorax and abdomen densely covered with rather long, fine spicules. Comb with numerous scales in a patch; each scale rounded apically and fringed with subequal spinules. Siphonal index about 4.5; five multiple siphonal tufts inserted beyond the pecten. Pecten with about 16 teeth on the basal fourth of the siphon; each tooth with coarse barbs on one side. Anal segment spiculate, com- pletely ringed by the saddle. MaTiERIAL EXAMINED.—Four adult males and associated terminalia from Brazil, one from Colombia, and one from Surinam. The lecto- type male terminalia has also been studied. DistrisuTion.—Reported from Surinam, French Guiana, Vene- zuela, Brazil, and Bolivia. One specimen in the U.S. National Museum collection from La Dorada, Colombia has been examined by the author. Culex (Culex) tarsalis Coquillett, 1896 FiauREs 32c, d Culex (Culex) tarsalis Coquillett, 1896, p. 43. Culex (Culex) willistont Giles, 1900, p. 281. Culex (Culex) kelloggit Theobald, 1903b, p. 211. Systematics.—Culex tarsalis demonstrates close affinity with C. abnormalis in structures of the male terminalia, but may be distin- guished by the arrangement and number of the appendicles on the apical lobe of the basistyle. SALIENT CHARACTERS.—Adult female: See table 1. Male terminalia: Basistyle conical, slightly longer than twice the basal width; clothed with fine setae in addition to the normal, long setal pattern. Apical lobe of the basistyle prominent, undivided, NO. 3557 SUBGENUS CULEX—BRAM 107 and also clothed with fine setae. Appendicles of the apical lobe as follows: two strong, straight, subequal, bluntly rounded rods followed by a straight, pointed seta; leaf very narrow, somewhat pointed at apex; two accessory setae present, one as long as leaf, the other broader and longer than the leaf. Dististyle normal. Tenth sternite crowned with a dense tuft of short, pointed spines apically and about seven spatulate, scalelike spines on the lateral, outer margins; basal arm rather long, distinctly curved; eight to 10 cercal setae present in a large patch on the lateral margin. External process rather narrow, eradually tapering to a blunt point; slightly exceeding the ventral cornu in length. Ventral cornu long and narrow, similar in size and shape to the external process. Median process with three to five strong, pointed, distinctly separated teeth. Basal process long, thin, straight, and bluntly rounded at the apex. Larva: Antennal tuft located in a constriction near the outer third; antennal shaft spiculate basally. Postclypeal head hair 4 short, single; frontal head hairs 5, 6, and 7 multiple, long, and barbed. Mentum with about 13 sharply pointed teeth; the apical tooth is longer and broader than the lateral teeth. Thorax glabrous. Comb with many scales in a patch; each scale rounded apically and fringed with sub- equal spinules. Siphonal index 4.5 to 5.5; five multiple siphonal tufts inserted in a line beyond the pecten. Pecten with 10 to 15 teeth on the basal third of the siphon; each tooth with one to five coarse barbs on one side. Anal segment completely ringed by the saddle, spiculate distally. MATERIAL EXAMINED.—A series of over 50 specimens from the United States and five adult males and associated terminalia from Mexico. Distripution.—Carpenter and LaCasse (1955) reported that this species ranges from southwestern Canada through the western, central, and southern United States and into Mexico. Culex (Culex) thriambus Dyar, 1921 FIGURES 33a, 6 Culex (Culex) thriambus Dyar, 1921a, p. 33. SysTEMATICS.— This species demonstrates affinities with Culex peus, C. interrogator, and C. stenolepis. It may be distinguished by the number and arrangement of appendicles on the apical lobe of the basistyle and by its characteristic and diagnostic external process. SALIENT CHARACTERS.—Adult female: See table 1. Male terminalia: Basistyle conical, slightly longer than twice the basal width; clothed with fine setae in addition to the normal, long setal pattern. Apical lobe of the basistyle prominent, undivided, 108 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 and also clothed with fine setae. Appendicles of the apical lobe as follows: basal rod strong, straight, and pointed; median rod longer and broader than the basal rod and terminating in a gentle hook; apical rod not as broad as, but longer than the median rod, also termi- nating in a gentle hook; leaf moderate in size, obovate, exhibiting distinct longitudinal striations; one long, pointed, gently curved accessory seta present. Dististyle normal. Tenth sternite crowned a Ficure 32.—C. surinamensis, Bahia, Brazil, USNM RB62 651: a, basistyle and dististyle; b, mesosome. C. tarsalis, Federal District, Mexico, USNM RB62 706: c, basistyle and dististyle; d, mesosome, NO. 3557 SUBGENUS CULEX—BRAM 109 with a dense tuft of short, pointed spines; basal arm moderate in length, prominently curved; three cercal setae present in a compact group. External process broad on the proximal half, then sharply tapering to a point which reaches slightly beyond the ventral cornu; a prominent additional sclerite on the inner margin of the proximal half. Ventral cornu large, rounded and rugulose. Median process with four to six distinct, sharply pointed teeth. Basal process short, straight, and bluntly rounded. Larva: Antennal tuft located in a constriction near the outer third; antennal shaft spiculate basally. Postclypeal head hair 4 short, single; frontal head hairs 5, 6, and 7 multiple, long, and barbed. Mentum with about 17 teeth; the apical tooth longer and broader than the lateral teeth; subbasal lateral teeth longer than other lateral teeth. Thorax glabrous. Comb with many scales in a patch; each scale rounded apically and fringed with subequal spinules. Siphonal index about 6.0; four siphonal tufts consisting of single setae inserted beyond the pecten. Pecten with about 10 to 14 teeth on the basal third of the siphon; each tooth with one to four coarse barbs on one side. Anal segment completely ringed by the saddle which has several fine spicules distally. MATERIAL EXAMINED.—Five adult males and associated terminalia from Costa Rica, one from Colombia, one from the Dominican Republic, and three from the United States. The lectotype male terminalia was also studied. DistrisuTIon.—Reported from the southwestern United States, Mexico, and Panama. The author has seen specimens in the U.S. National Museum collection from: Santa Anna, Costa Rica; Bara- hona, Santo Domingo, Dominican Republic; and Catera al Colegio, Bogota, Colombia. Culex (Culex) tramazayguesi Duret, 1954 FIGURE 33¢ Culex (Culex) tramazayguesi Duret, 1954, p. 23. SysTEeMATIcs.—Examination of Duret’s (1954) original figures and description of this species suggests the possibility that it is not a member of the subgenus Culex, however, since actual specimen have not been examined, the species is considered a member of the subgenus for purposes of this study. Culex tramazayguesi is readily distinguished from all other members of the subgenus by the characteristic shape and number of the ap- pendicles on the apical lobe of the basistyle. SALIENT CHARACTERS.—Adult female: See table 1. Male terminalia: Basistyle conical, about twice as long as the basal width. Apical lobe of the basistyle prominent, divided into two sec- 221-521-679 110 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 tions: the proximal section is conical and possesses two robust, heavily sclerotized, bluntly rounded rods: the distal section is also conical, but possesses only one heavily sclerotized, bluntly rounded rod: numer- ous setae are also present on the apical lobe. Dististyle normal. Tenth sternite crowned with a dense tuft of short, pomted spines; basal arm short and straight; about eight cercal setae present. Ex- Ficure 33.—C. thriambus, Santa Anna, Costa Rica, USNM RB62 73: a, basistyle and disti- style; b,mesosome. C. tramazayguesi (drawn after Duret, 1954): c, apical lobe of basistyle and dististyle. NO. 3557 SUBGENUS CULEX—BRAM 111 ternal process gradually tapering to a blunt point. Ventral cornu absent. Median process without teeth. Basal process absent. Larva: Unknown. MATERIAL EXAMINED.—None. Known to the author only from the original description. DistriBuTIoN.—Known only from the type locality, Monte Coman, Mendoza, Argentina. Unrecognized Species Culex (Culex) virgultus Theobald, 1901 Culex (Culex) virgultus Theobald, 1901, p. 123. Following the recommendation of Stone (1956 [1957]), Culex virgultus is considered an unrecognized species in the present study. Culex (Culex) guayasi Levi-Castillo, 1953 Culex (Culex) guayast Levi-Castillo, 1953, p. 15. In the original description, Levi-Castillo (1953) described the female and included figures of the male terminalia. I have been unable to utilize the generalized descriptions in keys or otherwise distinguish this species from other members of the subgenus. Since type specimens are not available for study, Culex guayasi is here con- sidered as an unrecognized species. Culex (Culex) quitensis Levi-Castillo, 1953 Culex (Culex) quitensis Levi-Castillo, 1953, p. 190. As mentioned above for C. guayasi, the generalized original de- scription of this species is not sufficient for absolute recognition. Since type specimens are not available for study, C. quitensis is here considered as an unrecognized species. References Apams, C. F. 1903. Dipterological contributions. Bull. Univ. Kansas, vol. 2, no. 2, pp. 21-47. AnpuzB, P. J. 1943a. Estudios de entomologia medica en el estado Merida—Venezuela: La fauna Culicidiana: Descripcion de Culex (Culex) albertoi sp. n. Bol. Ent. Venezolana, vol. 2, pp. 192-195. 1943b. Culex (Culex) beauperthuyi sp. nov. (Diptera: Culicidae). Rev. Venezolana Sanid. Asist. Soc., vol. 8, pp. 459-461. BacuHMANn, A. O., and Casat, O. H. 1962. Notas sobre Culex (Culex) Argentinos (Diptera: Culicidae). An. Inst. Nac. Microbiol., vol. 1, pp. 77-81. 1963. Notas sobre Culex (Culex) Argentinos, 2. Rev. Soc. Ent. Argentina, vol. 25, pp. 39-42. 112 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Bates, M. 1949. 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Culex (Culex) scimitar, a new species of mosquito from the Bahama Islands. Proc. Ent. Soc. Washington, vol. 61, pp. 216-218. BRETHES, J. 1916. Algunas notas sobre mosquitoes Argentinos (su relacion con las enfermedades palucidas, etc. y descripcion de tres especies nuevas), vol. 28, pp. 213-214. 1920. Description d’un nouveau moustique du Perou. Rev. Chilena Hist. Nat., vol. 24, pp. 41-43. CaRPENTER, 8. J., and LaCassz, W. J. 1955. Mosquitoes of North America (north of Mexico), 360 pp. Castro, G. M. 1932. Estudio sobre una especie de Culex, que se cria em buracos de gua- yamu. (Dipt., Culicidae). Rev. Ent. Rio de Janeiro, vol. 2, pp. 97-105. CHRISTOPHERS, S. R., and Barraup, P. J. 1923. Descriptive terminology of male genitalic characters of mosquitoes. Indian Journ. Med. Res., vol. 10, pp. 827-835. Couusss, D. H. 1957. Notes on the culicine mosquitoes of Singapore, 2: The Culex vishnuiz group with descriptions of two new species. Ann. Trop. Med. Parasitol., vol. 51, pp. 87-101. 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Aussereuropaische zweiflugelige insekten, vol. 1, 608 pp. 117 CULEX—BRAM SUBGENUS NO. 3557 Culex (O=character absent; X—character present; ?= presence or absence of TABLE 1.—Salient female characters of New World species of the Culex subgenus character unknown to author) pueq dT M BV UYM PosuUlI STOSO(oIg SSULYICUI OJIN M YIM Id[eg So[Bos IOATIS JO ulo93ed AIM TUNYOUOSE/[ soyeos ail JO soyojed YA BMI J 9e4as IO Seyeos OYA YIM UIMI[eyNIg Bjsooqns pues B4S00 10 SOTeOS OJIYA pota}}BoS YIM SSUIM pueq [eorde oyed 441M InWe7 pueq [eolde ated 44IM BIQhL spueq OUIUM TILM SJUSUIBIS [BSIET, eqtyM Ajos1yUa quourses [esie} pulYy yse'T spueq Teseq oy MOI -IBU YIM SJUeUIZes TeUIMOpg VY spueq [eseq og peoid WLM S}UeUIses [BUIMIOPqY A p9}0orIp Ayepneo yiLM spueq [eurmopqy DOKNDOOHODODOKNHOCOOCOROCOKHOHODODOOONOKOON SSOSSCOMHOSOSOOSCOSSCSOSOCOCOOCOOCOOOC OOOOH OOCOCCoOOX HMOKMOSOKHODDOHOKOOODOOODOCOKOOOOKXOOCOOOCOOOK Moe Ma MAMMA NOM KM MMMM KM NM M AMON AMM MO MMONMOM ROOK KOM MM KM KM MMM OMe MOR On rR OMNM MMA MMM MOM SOCK OOSOOSCSOOCOOCOOOOROOCOOOSOOOKH OOOO OK KOx RAMON MK MMMM NM MMMM MOC OKMNMNOKNHONKOOKNKHMONM KKK MMMM NM KMOKMMMNMNOOKOOKMNKKKONMNMNOKMMNMOM DCOKOODCOKNHOKOCOOKKKOOKKOKOXKOOKNOKNKMNKMMNOM Sl SC OC OCC COSCO SCO SC COO SC COC SCC OSC OOOO OC eS KROKOKAHKMMNMNOMNKMMOKOCOOKKKMMNOKMOKOKOKKOONMNM OKOKODCOODOCOOOKOOKHOCOOCOOOCOODOOOKOOCOOCOO Oa ODOCSOKHOOOCOCOOOKOOCOnMOOCOCOOOCOOCOCOCOCCOCOOR diplophyllum dolosus maracayensis mauesensis erythrothorax maxt carcinoxenus finlayt brevispinosus chidestert interrogator janitor nigripalpus levicastilloi peus bahamensis lygrus abnormalis acharistus airozat apicinus archegus articularis bidens bonneae brethest corniger coronator declarator delys duplicator foliaceus habilitator inflictus lahillei laticlasper mollis VOL. 120 PROCEEDINGS OF THE NATIONAL MUSEUM 118 TABLE 1.—Salient female characters of New World species of the Culex subgenus Culex—Continued pueq OIA & YIIM pesull stosoqoig SBUPYIVUL O}TYM ITM Idyjeg S9TBOS IOATIS JO us0}38d YIM UIM4OUOSET[ sees ey JO soyojzed WIM BIMo[q 9R40s JO SeTBOS O}[YM TILA UUINI]JeynNIg B4sooqns pus BSO0d UO SO[BOS O}IM Poso}}BOs YILM SSULM pueq [eorde syed YILM snuta,T pueq [eorde oped yITM vIqLL spueq OTT. WIM Syuourses [esze, J, ou AToryu9 qJUeUIZeS [BSIv} PUIY 4Se'T spueq [eseq oJIGM MOI -I8U W}LA SJUNUIZeS [BUTMIOpgYy spueq [eseq oyIGM peoiq ITM SjueuIdes [BUTUIOpqy A pe}00r1p Ayjepneo 44th spueq jeurmopqy CONMNMOOCODOOOKNONOCO SCOKMKMOCODCOOOOOHOOOO HOOKMODDOOOOMHOOOO PA OM AMM MM ORM MMA MOM MM SSOCSCSCSCOCOSOOOCOKOHOSO MM OKM KM OOM KM MM PM MM POR KO MM MO OP OO ROK KMODOKOOMNM MMMM NM RODD DODCOOOKOOKKOO MAAN OOHKHMMNMOKNOKMMMOO SCODOKHOOCOOCOKHOOONNM SVS) (NOK) eoNeKorev(eKe)(e(2)(S)\xo®) 5 LZ) 8 & % ‘2 8 ae & eS % Sc 8 = o.s os & 33S 2 ow 2S e'S-. SL § SRE, HS 8-88 SASS ea tS OS Eee Bs BESspsssssesczse ses38 Sie 2.8 8 8 2 See ss Sakis = 8 £2 2d pseu Ges Sk 2S So Ss 2 SkSS SR AaASSESBE SS RAALHHRH HHA HKEASBHE SS NO. 3557 SUBGENUS CULEX—BRAM 119 TABLE 2.—Terminology used in describing structures of the mesosome in Culex (Culex) (*=variation in use of the term within the paper under consideration.) Source Structures Freeborn and External process Ventral cornu Median process Basal process Mesosome Bohart (1951) Carpenter and | Dorsal arm Ventral arm Lateral teeth of not named Phallosome LaCasse * Me phallosome s (1955) Ventral arm not named Lateral teeth of Dorsal arm Phallosome . < phallosome * Lane (1953) Upper horn Upper horn Teeth of Lateral horn of Mesosome . be mesosome mesosome Dyar (1928) Upper arm Upper limb Teeth of Lower arm Mesosome mesosome Martinez Apendice del Primer diente Dientes Apifisis dorsal | Mesosoma Palacios borde interno apical apicales (1950) Roth (1943) Ventral arm not named Teeth of Dorsal arm Mesosome mesosome Bohart and External process Ventral cornu Median process Basal angle Mesosome Ingram (1946) Matheson Upper arm Apical tooth of | Outer plate Lower arm Mesosome (1944) outer plate Natvig (1948) Dorsal sclerite | Ventral sclerite Median sclerite Median sclerite Phallosome Belkin (1962) Inner division Tergal arm Outer division Lateralarmof Phallosome the outer division TABLE 3.—Distributional analysis of the species in the Culex coronator complex Number of Specimens Studied Area c. coronator c. usquatus c. camposi c. ousqua = ¢. Usquatissimus Total I vf 0 0 0 0 7 II 43 3 0 a 0 53 III 11 1 0 12 3 27 IV 2 5 14 1 4 26 Vv 8 13 0 0 2 23 VI 3 3 0) 0 0 6 VII 4 23 0 0 0 27 VIII 26 1 0 0 0 27 IX 0 2 f 0 0 6 x 0 0 24 0 1 25 to So — i=) bo bo J Total 104 51 42 120 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Addendum Culex (Culex) fernandezi Casal, Garcia, and Cavalieri (1966, Physis, vol. 26, pp. 185-192): In the key to species based on male terminalia, CO. fernandezi would key to couplet 49, but differs from C. salinarius in the development of the external process. In the key to species based on the larva, C. fernandezi would key to the vicinity of couplet 16; however, features of the chaetotaxy, particularly the stellate thoracic and abdominal setae, are quite distinctive. This species is known from the Province of Salta, Argentina, and the larvae were collected exclusively from bromeliads. abnormalis, 20 acharistus, 22 aestuans, 90 affinis, 87 aglischrus, 76 aikenit, 90 airozat, 23 alani, 23 alberto, 50 alticola, 24 apicinus, 24 archegus, 28 articularis, 30 aseyehae, 90 autumnalis, 90 azuayus, 84 badger, 62 bahamensis, 32 barbarus, 90 basilicus, 43 beauperthuyt, 96 bickleyt, 33 bidens, 35 bilineatus, 59 biocellatus, 84 bonariensis, 59 bonneae, 36 brehmet, 94 brethest, 38 brevispinosus, 59 campost, 49 caraibeus, 84 carcinoxenus, 40 carmodyae, 84 chidesteri, 42 comitatus, 90 consobrinus, 90 corniger, 43 coronator, 46 coronator mooseri, 46 covagarciai, 53 cubensis, 90 deanei, 42 debilis, 24 declarator, 56 Index delys, 57 dictator, 56 diplophyllum, 57 dipseticus, 90 dolosus, 59 duplicator, 60 eleuthera, 32 éelocutilis, 80 equivocator, 80 eremita, 66 erythrothorax, 62 escomeli, 24 eumimetes, 87 extricator, 68 factor, 84 fatigans, 90 federalis, 62 fernandezi, 120 ferruginosus, 90 jfinlayt, 63 foliaceus, 64 forattini, 56 guayast, 111 habilitator, 66 hassardit, 43 inflictus, 68 inquisitor, 56 interfor, 35 interrogator, 69 janitor, 71 jubilator, 56 kelloggiz, 106 lachrimans, 90 lactator, 43 lahillez, 72 lameniator, 100 lateropunctata, 80 laticlasper, 72 lepostenis, 8O levicastillot, 74 loquaculus, 43 lygrus, 76 maracayensis, 76 mauesensis, 78 maxi, 79 122 PROCEEDINGS OF THE NATIONAL MUSEUM microannulata, 84 microsquamosus, 84 mollis, 80 mortificator, 84 nigripalpus, 84 oswaldot, 86 ousqua, 50 palus, 84 penafieli, 90 petersont, 32 peus, 87 pinarocampa, 88 pipiens, 90 prasinopleurus, 84 proclamator, 56 proximus, 84 pseudojanthinosoma, 91 pungens, 90 quasisecutor, 100 quinquefasciatus, 90 quitensis, 111 raymondii, 90 reflector, 69 regulator, 84 renatot, 92 restuans, 94 revelator, 56 revocator, 90 rigidus, 43 salinarius, 95 saltanensis, 97 scholasticus, 68 scimitar, 98 scutatus, 20 secutor, 100 serotinus, 90 similis, 84 sphinx, 101 spinosus, 103 stenolepis, 104 stigmatosoma, 87 subfuscus, 43 surinamensis, 105 tarsalis, 106 tejerai, 74 thriambus, 107 tisseuilltz, 80 towert, 100 tramazayguesi, 109 usquatissimus, 51 usquatus, 49 vindicator, 56 virgultus, 36, 111 willistont, 106 U.S. GOVERNMENT PRINTING OFFICE: 1967 1 di hia t Bait ad F.! of K , . _ i * i}; ' : i a nf é Bie ae sie ; Le py Mey, Bice: | bari ~~ ' ’ f bale | ¢* Bhs are at ee Te I : ; Beet ed i yale ear ot a besuile 2 aie ‘ ne Sp alan, 54 | : Myra je ‘ a4 rt 7 i ay 6 { , , a P Hahn Cea eit d set | | 1) Fee arian . W' git pi — At eale A ¥ Rew ol) al ee ut Rateett i 4 At ia ie te ev i . en in se belip: a PRE ae 8 ; o on cia Wie . a v5 ce ¥ . ) y ‘ 1 ! ; : 1 ( i m } : / \\ \ : hal , il. 7 i bye init OT P bs * ag wee Bae, } eee Proce ed ries ‘ot the United States National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. a eee a Volume 120 1967 Number 3558 AMBLYCERAN MALLOPHAGA (BITING LICE) FOUND ON THE BUCEROTIDAE (HORNBILLS) ! By Rospert E. Ese. ? Mallophaga of the genera Chapinia and Bucerophagus of the am- blyceran family Menoponidae are found only on hornbills. The purpose of this paper is to redescribe and illustrate the known species in these genera, describe new species encountered, and compare the classification of these lice with that of the hornbills. Menoponidae have been examined from 53 species or subspecies of hornbills (table 13). Presented are descriptions and illustrations of 22 species of Menoponidae of which 17, including 12 new, are species of Chapinia, 3 are species or Bucerophagus, and 2 are new species in a new genus, Bucerocolpocephalum. No previous attempt has been made to examine all the Menoponidae from the hornbills. Clay (1947) included Chapinia and Bucerophagus in her key to the genera of the Menoponidae, but her figures 8 and 9 of the antennae of these genera were transposed accidentally. The genus Chapinia was described by Ewing (1927) for his species C. robusta; later it was described by Bedford (1930) for Menopon bucerotis 1 Modified from a doctoral dissertation submitted to the University of Okla- homa, Norman. 2K. and E. Branch, Dugway Proving Ground, Dugway, Utah. 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Kellogg, 1908, and M. lophocerus Bedford, 1920. Hopkins and Clay (1952) included in the genus Chapinia the additional species, Colpo- cephalum hirtum Rudow, 1866, Menopon acutovulvatum Piaget, 1881, and Allomenopon mjébergi Eichler, 1947, but they considered the generic position of C. hirtum doubtful. In the present study C. hirtum is shown to be a Chapinia, and a neotype is designated; A. mpébergi is shown to be a synonym of M. acutovulvatum. Hopkins (1941) designated a lectotype for Chapinia lophocerus (Bedford), Clay (1949a) designated a lectotype for C. acutovulvata (Piaget), and a lectotype is designated here for C. bucerotis (Kellogg). Piaget (1880) identified a female from hyticeros cassidix (Temminck, 1823) as C. hirtum, but Piaget’s specimen is shown here to be the new species Chapinia lydae. The genus Bucerophagus was described by Bedford (1929) for his species B. africanus and for Colpocephalum productum Burmeister, 1838. For the latter species a neotype was erected by Conci (1950), and a lectotype was designated by Clay (1951a). Hichler (1947) described a new genus for Menopon forcipatum Nitzsch, 1874, but Hopkins and Clay (1952) put MZ. forcipatum in the genus Bucerophagus. A neotype for B. forcipatus (Nitzsch) is designated here from Eichler’s material. Clay (1951la) stated that Bucerophagus africanus and B. productus both infest the two hosts, Bucorvus abys- sinicus (Boddaert, 1783) and B. leadbeateri (Vigors, 1825). No morphological or statistical means were found in the present study to separate the populations of each species on each host so that only the two species, Bucerophagus productus and B. africanus, could be recognized. The phylogenetic arrangement of the hornbills (Peters, 1945) shows scant regard for the geographical regions, and the list winds back and forth between the Ethiopian, Oriental, and Australasian regions (table 13). The mallophagan genera studied here, however, fall into definite species-groups confined to the Ethiopian region or to the Oriental and Australasian regions. It is believed, therefore, that the arrangement of the Mallophaga gives more insight into the origin of the hornbills than study of the host skins. Classification of the hosts is that proposed by Deignan (1963) except for species not discussed by him, for which Peters (1945) has been followed. Skins of the hosts collected in Thailand are in the U.S. National Museum and were identified by Mr. H. G. Deignan. Collections were made possible by assistance from the U.S. Operations Mission to Thailand and the U.S. National Museum. Grateful appreciation is expressed to Dr. Cluff E. Hopla, Oklahoma University, Dr. K. C. Emerson (KCE), Stillwater, Okla., and Dr. Theresa Clay, British Museum (Natural History) (BMNH), for their many suggestions and constant help; to Dr. Clay for the loan NO. 3558 MALLOPHAGA—ELBEL 3 of specimens in the BMNH, in the Meinertzhagen, in the Piaget, and in the G. H. E. Hopkins (GHEH) collections, the latter from the Zoological Museum, Tring, Hertfordshire, England, for obtaining from Dr. Eichler and Dr. von Kéler the loan of specimens in the Zoological Museum, Humboldt University, Berlin, and from Dr. Joio Tendeiro (JT) the loan of specimens in the Centro de Zoologia, Lisboa, Portugal; to Dr. Eric Kjellander for the loan of specimens in the Swedish Museum of Natural History (SMNH); to Dr. Rupert L. Wenzel for the loan of specimens in the Chicago Natural History Museum (CNHM); to Dr. K. C. Emerson and Mr. C. F. W. Muese- beck for the loan of specimens in the U.S. National Museum (USNM), and for mounting mallophagan dried material obtained from that museum. Special thanks are extended to the Chicago Natural His- tory Museum, the Michigan University Museum of Zoology (MMZ), the U.S. National Museum, and Dr. Boonsong Lekagul (BL), Bangkok, Thialand, for permission to examine their collections of hornbill skins for Mallophaga; to Dr. Boonsong Lekagul, Mr. H. G. Deignan, Mr. Kitti Thonglongya of Seato Medical Research Laboratory (SMRL), Bangkok, Thailand, Mr. Wanit Songprakob and Mr. Wichit Suwan Laong, both of Songkhla, Thailand, for help in collecting mallophagan fresh material from Thailand; to Dr. Alfred F. Naylor, formerly of Oklahoma University, for help in statistical comparisons of louse populations on the hosts, Bucerophagus productus and B. africanus; to the Bernice P. Bishop Museum (BPBM), Honolulu, Hawaii, and the Prirodoslovni Musej Slovenije (PMS), Ljubljana, Yugoslavia, for the loan of specimens; and to my wife, Lyda (LE), for help in prepara- tion of the manuscript and in various phases of the study. Explanation of Terms The terminology used in this paper agrees with that of Clay (1947) except as noted below. “Combs of setae” are rows of short, stout setae, each with the alveoli lying close together and approximately in a straight line, on the venter of the third femora and the posterolateral areas of one or more abdominal sternites (figs. 64, 65). “Brushes of setae” are concentrations of setae on the venter of the third femora and the posterolateral areas of one or more abdominal sternites. These may take the form of a few widely spaced setae called “‘small scattered brushes” (figs. 23, 24) or a large number of closely set setae called “large thick brushes” (figs. 68, 69). The setae of the brushes are “normal” if approximately the same length and thickness as the surrounding setae or ‘‘small” if the majority are considerably smaller than the surrounding setae. 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 The “preocular slit” is an emargination with approximately equal and parallel margins in the dorsal-lateral margin of the head immedi- ately anterior to the eye (figs. 23, 24). The “preocular notch” is a similar emargination with triangular or semicircular margins (figs. 25, 26, 64-69). The “metasternal plate’ on the metasternum was first described by Bedford (1920) as a 4-sided plate which was normally expanded anteriorly (figs. 26c, 64c, 66c, 68c, 70). The male and female each possess a complete complement of abdominal segments from the first to the tenth (Cope, 1941). The “pleurites” (“paratergal plates’”’ of some authors) are sclerites which pass around the lateral margin of the abdomen and are sepa- rated from the tergites and sternites by clear divisions (Clay, 1954) (figs. 23, 24). The ‘“postspiracular seta’? on each side of abdominal tergites IIJ-VIII is posterior to the spiracle and is always associated with two small setae, the alveoli of the three setae being contiguous (Clay, 1954). A similar group of three setae is present on abdominal tergite II, although there is no functional spiracle on this tergite. In most Amblycera the postspiracular setae are the most laterad setae of the posterior marginal row (fig. 25p). The “male terminal abdominal tergites’” IX and X are almost completely fused. ‘Abdominal sternite VIII” is fragmented into two parts which have moved laterad. ‘Abdominal sternite IX” strengthens the anterior lip of the genital and anal opening (cloaca), and there is no apparent tenth sternite (Cope, 1941). The “basal plate” (“basal apodeme” of Clay, 1956) of the “male genitalia” supports posteriorly the slender ‘“‘parameres” laterally and the ‘“endomeres” centrally (Ewing, 1927). The “sclerite of the male genital sac” is the sclerite on the walls of the genital sac which is articulated to the basal plate (Clay, 1956) (figs. 18s, 19s). The “lateral horns” of the ‘‘male genitalia” are structures on each side of the large, curved, platelike ‘“endomeres” (Ewing, 1927) (figs. | 1-3, 6, 7). The “female terminal abdominal tergites’’ IX and X are single plates. ‘Abdominal sternite VIII” has a pair of gonopods, lying side by side medially, which are fused at their apices to cover the genital opening (vulva) between segments VIII and IX, and on each lateral side of the gonopods is a fragment of sternite VIII (Cope, 1941). The “internal sclerite of female abdominal sternite VIII’’ appears to be the sclerite that Clay (1961) calls the internal structure of the female genital chamber (figs. 35i, 38i, 391, 491). “Sclerital hooks” are hooklike processes in the female arising on — NO. 3558 MALLOPHAGA—ELBEL 5 either side of the midline of the “ventral sclerite between the vulva and anus” (fig. 27h). The “anal fringe” (‘anal corona” of Ferris, 1923) surrounds the female anus on abdominal segment X (figs. 28, 45, 52). “‘Species-groups” are groups of similar species within a genus. “Fresh material” indicates that Mallophaga were obtained from the host that was collected in the field as contrasted to mallophagan “dried material”? which was obtained from dried museum skins either personally (REE) or by my wife, Lyda. Methods Dried material was obtained from museum skins by lightly fluffing the bird feathers, particularly around the neck and lower belly, over a white surface. Emerson (1954) stated that contamination that occurred on museum skins was well known and that records of Mal- lophaga so obtained should be considered questionable. Mallophaga that were obtained from museum skins were considered here to be stragglers unless they belonged to recognized hornbill genera and unless they were represented by other specimens obtained from additional skins or fresh material of the same host species. Cor- respondingly, about 20 percent of the dried material was considered to be stragglers. The mounting procedure was suggested by Dr. K. C. Emerson (in litt.): Mallophaga were placed in 10 percent potassium hydroxide overnight, transferred to distilled water, and after one hour the body contents were teased out. Specimens were placed in fresh 10 percent potassium hydroxide for 6-12 hours, after which they again were transferred to distilled water. Approxi- materly one-half hour later specimens were put into 40 percent ethyl alcohol. Fifteen minutes later several drops of carbol fuchsin (Ziehl Nielson) were added and allowed to act for one-half hour. Specimens were placed in 70 percent ethyl alcohol for one-half hour, followed by 95 percent ethyl alcohol for 15 minutes. Next, speci- mens were washed in 100 percent ethyl alcohol for a few minutes and placed in Beechwood Creosote for one hour to overnight, after which they were mounted in Gum Damar or other dried resin media. Drawings were prepared from holotypes and allotypes except as noted in the text. All drawings were prepared with the aid of a 300-watt, 35-mm. slide projector as suggested by Dr. K. C. Emerson (in litt.). The monocular microscope with the mounted Mallophaga was turned on its side, the ocular and mirror removed, and the slide projector placed at the lower end of the microscope so that the light projected the image onto a vertical surface, from which the outline was traced on Bristol board or drawing velum. Measurements were obtained by projecting a millimeter scale from a stage micrometer 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 onto the surface. Details of the Mallophaga were added after the microscope was uprighted. Measurements for the tables are in millimeters and were made with the aid of an ocular micrometer. Because of variation in setal number, setae recorded in species descriptions represent the range in numbers of setae from repre- sentative specimens from the material examined. Characters described under genera or species-groups have not been repeated for individual specific descriptions. In each genus or species- group the arrangement of the species is based first on morphological similarity and second on the phylogenetic arrangement of their hosts. Key to Species of Chapinia, Bucerocolpocephalum and Bucerophagus 1. Terminal segment of antenna showing definite signs of division either by transverse line or marginal indentation (figs. 64a, 66a, 68a). . . . . 19 Terminal segment of antenna without signs of division (figs. 23a, 25a). (Genus Chapinia Ewing). . . 2 2. Dorsal-lateral margins of head with a preocular slit (figs. 23, 24). (lophocerus species-group) . . . 3 Dorsal-lateral margins of head with a preocular notch (figs. 25, 26). . . . 7 3. Each lateral margin of abdominal tergites III-VI with a short seta between the spiracle and postspiracular seta (fig.64]l).... cy fy po Rope pontine Abdominal tergites without such short setae (figs. 23, 24). C. robusta Ewing, p. 17. 4. Abdominal sternite II with three median rows of setae. C. bucerotis (Kellogg), p. 15. Abdominal sternite II with one median row of setae on posterior margin . 5 5. Male genitalia with each lateral horn possessing two sharp posterior points; female anal fringe with fewer than 58 setae (figs. 2, 28). C. lophocerus (Bedford), p. 13. Male genitalia with each lateral horn possessing one or two rounded posterior points; female anal fringe with more than 58 setae... . AG: REREG 6. Male genitalia with each lateral horn possessing two rounded posterior points; female anal fringe with more than 70 setae (fig. 1). C. fasciati, new species, p. 12. Male genitalia with each lateral horn possessing one rounded posterior point ; female anal fringe with 60—64 setae (fig. 3) . C. camuri, new species, p. 15. 7. Each lateral margin of abdominal tergites III-VI with a short seta between the spiracle and postspiracular seta (fig.641).......... 8 Abdominal tergites without such short setae... . ee ge ot 8. Male genitalia with endomeres possessing small inner ales eng paired outer rims, each with serrulations on posterior inner margin; female with each lateral projection of ventral sclerite between vulva and anus possessing more than eight thick, posteriorly directed setae (figs. 71, 72sp). C. waniti, new species, p. 22. Male genitalia with endomeres possessing small or large inner plate and paired outer rims without serrulations; female with each lateral projection of ventral sclerite between vulva and anus possessing fewer than eight thick, posteriorly “directed setae’ (ig. /43sp) 2°72. 4%. a) ee ee NO. 10. 11. 12. 13. 14. 15. 16. is 3558 MALLOPHAGA—ELBEL a Male genitalia with endomeres possessing paired outer rims and large shieldlike inner plate with paired lateral flanges and central terminal point; female abdominal sternite VIII with more than 32 setae on posterior margin (figs. 12, 37)... . . . C. malayensis, new species, p. 25. Male genitalia with endomeres possessing paired outer rims and small inner plate that is not shieldlike; female abdominal sternite VIII with fewer than 32 setae on posterior margin... . seins 10 Male genitalia with endomeres OReeaaine Ne on teach tee rim a triangular internal knob wider than long; female terminal abdominal tergite with at most five long setae each side of midline on posterior margin (figs. 8, 33). C. clayae, new species, p. 21. Male genitalia with endomeres possessing on each outer rim a triangular internal knob longer than wide; female terminal abdominal tergite with at least six long setae each side of midline on posterior margin (figs. 9, 35). C. acutovulvata (Piaget), p. 23. Abdominal sternite II with at least two median rows ofsetae. .... . 16 Abdominal sternite II with one median row of setae on posterior margin. (acutovulvata species-group) . . . 12 Male abdominal sternite II with more than 46 total setae; female anal fringe with more than 48 setae... . Mee te 1S Male abdominal sternite II with fewer than 44 torah setae: femmnle anal fringe with fewer than 46 setae... . De ee eat tad mee I Male genitalia with endomeres coneeane orale of ERG cuter rims which are curved inwardly; female abdominal sternite VIII with internal sclerite almost as wide as long (figs. 13, 388i). . . . C. hoplai, new species, p. 26. Male genitalia with endomeres consisting only of paired outer rims which are straight and nearly parallel; female abdominal sternite VIII with internal sclerite much longer than wide (figs. 14, 39i). C. boonsongi, new species, p. 27. Male genitalia with endomeres consisting of paired plates having posterior extension split; female venter of third femora and abdominal sternites IV-VI without brushes (figs. 17, 25) . C. traylori, new species, p. 31. Male genitalia with endomeres either lacking paired plates or, if present, posterior extension not split; female venter of third femora and pos- terolateral margins of abdominal sternites IV-VI each with brushes of normal setae... . Ars. UES Male venter of third ne oma Gabatavole cecalit margins of abdominal sternites IV-VI each with brushes of normal setae; female abdominal sternite VIII without internal sclerite (fig. 40). C. wenzeli, new species, p. 28. Male venter of third femora and abdominal sternites IV-—VI without brushes; female abdominal sternite VIII with internal sclerite having slender posteriorly divergent margins (fig. 43) . . C. blakei, new species, p. 29. Venter of third femora and posterolateral margins of abdominal sternites IV—-VI each with brushes of normal setae... . AAS ar 1 Venter of third femora and abdominal sternites IV-VI pitnout brushes Gigs725226)N i). Mnene . .. . . C.traylori, new species, p. 31. Male genitalia with ore anees ealnveed anteriorly; female terminal ab- dominal tergite with the two median setae on posterior margin as widely spaced as four times the distance between each of the 3-4 long setae each side of midline (figs. 20, 46). . . . . . . C.lydae, new species, p. 32. Male genitalia with parameres slender anteriorly. (hirta species-group) . . . 18 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 18. Sclerite of male genital sac large, nearly twice as long as wide; female un- known (fig. 18s)... ... . . . C, muesebecki, new species, p. 34. Sclerite of male genital sac small, about as wide as long; female terminal abdominal tergite with 5 long setae on posterior margin each side of mid- line, these 10 setae being evenly spaced (figs. 19s, 48). C. hirta (Rudow), p. 35. 19. Venter of third femora and posterolateral margins of fourth abdominal sternite each with three full rows of combs (figs. 64, 65). (Genus Bucerocolpocephalum, new genus). . a Venter of third femora and fourth abdominal sternite without combs. (Genus Bucerophagus Bedford). . . 21 20. Male genitalia with endomeres having a pair of posterior points; female abdominal sternite VIII with internal triangular sclerite (figs. 49, 57). Bucerocolpocephalum emersoni, new species, p. 37. Male genitalia with endomeres lacking a pair of posterior points; female abdominal sternite VIII without internal sclerite (figs. 50, 58). Bucerocolpocephalum deignani, new species, p. 38. 21. Venter of third femora and posterolateral margins of abdominal sternites IV and V, each with brushes of normal setae (figs. 68, 69). . . .. . 22 Venter of third femora and abdominal sternites IV and V without brushes Gigs) 66,067) \<\ coc: ta," . . « Bucerophagus forcipatus (Nitzsch), p. 42. 22. Metasternal plate eens expanded anteriorly (fig. 70). Bucerophagus productus (Burmeister), p. 44. Metasternal plate trapezoidal, expanded anteriorly (fig. 68c). Bucerophagus africanus Bedford, p. 46. Genus Chapinia Ewing Figures 23-26 Chapinia Ewing, 1927, p. 88. [Genotype: Chapinia robusta Ewing, 1927.] Allomenopon Bedford, 1930, p. 153. [Genotype: Menopon bucerotis Kellogg, 1908.] Head triangular, width 1% to 2 times that of length. Forehead much narrower anteriorly. Temples expanded. Antennae 4-jointed, third segment constricted at base, and terminal segment capitate without signs of division. Antennary fossa deep, covered above by expansion of lateral margin of head, posterior margin of which bears eye with double cornea. Dorsal-lateral margin of forehead anterior to eye with preocular slit or shallow notch. Gular region with 3-7 setae varying in length on each lateral margin. Pronotum expanded anteriorly with posterior marginal row of long setae. Metanotum expanded posteriorly with posterior marginal row of long setae and two short setae on each lateral margin. Metanotum separated from mesonotum and from pleurites. Thoracic sternal plates as shown in figures 23b, 25b, and 26c. Metasternal plate trapezoidal, expanded anteriorly, with 6-22 setae. Venter of third femora and posterolateral margins of abdominal sternites IV-VI, each with or without brushes of normal setae. Abdominal segments consist of tergites, sternites, and pleurites, the latter without prolongation of posteroventral angles. NO. 3558 MALLOPHAGA—ELBEL 9 Abdominal tergites each with a posterior marginal row of setae, the most laterad being the postspiracular seta. Each lateral margin of abdominal tergites II-VI with or without a short seta between the spiracle and postspiracular seta. Sternites and pleurites each with a posterior marginal row of long setae and with numerous shorter setae. Male terminal abdominal sternites VIII and IX fused with partial division only from sternite VII (figs. 4, 10, 22, 24, 26). Male genitalia as illustrated for each species, with parameres either expanded an- teriorly or split posteriorly or both. Female terminal abdominal segments as illustrated for each species, with lateral processes arising from ventral sclerite between vulva and anus, with long stout setae but never strong spines. Females larger than males, usually with more abdominal sternal setae, but general shape and chaetotaxy similar to that of males except for terminal abdominal segments. Both Ewing (1927) and Bedford (1930) stated that the ptero- thorax was undivided. As noted by Cope (1941), the sclerotized median button behind the prothorax (fig. 25m) is a vestige of the mesonotum; the supposed mesonotum, the narrow sclerotized band posterior to this button, is a mere extension of the subcoxae. Ewing (1927) stated that the abdomen consisted of 9 segments in the female and 10 segments in the male, but as shown by Cope (1941), the abdomen of both the male and female has 10 segments each. Chapinia resembles most closely Bucerophagus (figs. 66-69) but differs in several characters: The terminal segment of the antenna shows no sign of division in Chapinia, but there are definite signs of division into two parts either by transverse line or marginal indenta- tion in Bucerophagus. The venter of the third femora may have brushes of normal setae in Chapinia and Bucerophagus; similar brushes are present on posterolateral margins of abdominal sternites [V-VI in Chapinia but abdominal sternites 1V and V in Bucerophagus. Each lateral margin of abdominal tergites JI-VI may have a short seta between the spiracle and postspiracular seta in Chapinia, but 1-5 short setae may be present on margins of abdominal tergites II-VIII in Bucerophagus. Male terminal abdominal sternites VIII and IX are fused in Chapinia with partial division only from abdominal sternite VII, but abdominal sternites VIII and IX may be fused in Bucerophagus with a complete division from abdominal sternite VII. Male genitalia have parameres slender or expanded anteriorly in Chapinia but branched anteriorly in Bucerophagus. Lateral processes arising from the ventral sclerite between the female vulva and anus have long stout setae in Chapinia but long stout setae and strong spines in Bucerophagus. The male genitalia and details of the male and female terminal abdominal segments are the best characters for separating species 10 PROCEEDINGS OF THE NATIONAL MUSEUM vou. 120 of Chapinia. Other characters useful in species separation are: The presence or absence of brushes of normal setae on the venter of the third femora and posterolateral margins of abdominal sternites IV-VI; the presence or absence of a short seta on each lateral margin of abdominal tergites III-VI between the spiracle and postspiracular seta; the number of median rows of setae, and the total number of setae on abdominal sternite II. The number and length of setae on the lateral margins of the gular region are too variable to be of much use in separating species. For convenience of classification the species of Chapinia have been arranged into species-groups. Hosts: Species of Chapinia have been found on the genera Tockus, Anorrhinus, Penelopides, MRhyticeros, Anthracocerus, Bycanistes, Ceratogymna, and Buceros of the avian family Bucerotidae. The lophocerus Species-Group Species similar in shape to Chapinia robusta (figs. 23, 24). Dif- fering from other species-groups in the following combination of characters: Dorsal-lateral margins of head with preocular slit; venter of third femora and posterolateral margins of abdominal sternites IV-VI each with brushes of normal setae; each lateral margin of ab- dominal tergites II-VI with or without a short seta between the spiracle and postspiracular seta; females with more abdominal sternal setae than males; abdominal sternite IJ with either one or three median rows of setae; male genitalia with lateral horns on each side of endomeres and with parameres enlarged anteriorly, not split pos- teriorly ; females with sclerital hooks on each side of midline of ventral sclerite between vulva and anus; female abdominal sternite VIII with most of setae similar in size to setae on posterior margin. Hosts: Species of the lophocerus species-group have been found on the genera Tockus, Bycanistes, and Ceratogymna of the avian family Bucerotidae. Species of the lophocerus species-group are all similar in size except that both sexes of Chapinia camuri are smaller than corresponding sexes of other species, and males of C. robusta are larger than other males. The small size of C. camuri might be expected since its host, the 15-inch Tockus camurus, is the smallest known hornbill. The venter of the third femora and posterolateral margins of abdominal sternites ITV-VI each have large thick brushes of normal setae in C. bucerotis but small scattered brushes of normal setae in other species of the lophocerus species-group, although the brushes are slightly thicker in C. robusta. Each lateral margin of abdominal tergites II-VI has a short seta between the spiracle and postspiracular seta in | NO. 3658 MALLOPHAGA—ELBEL 11 all species except in C. robusta. Abdominal sternite II has more total setae in both sexes of C. bucerotis and C. robusta than in corresponding sexes of C. fasciati, C. lophocerus, and C. camuri; this sternite has one median row of setae in all species except C. bucerotis, which has three median rows. The male genitalia have each lateral horn possessing posterior points in all species except C. robusta, which has one sharp median point crossing the broad endomeres; the posterior points are sharp in C. bucerotis and C. lophocerus but rounded in C. fasciatt and C. camuri; C. camurt has only one posterior point instead of two as in the other species; the two points do not reach the slender endomeres in C. lophocerus, but one point crosses the broad endomeres in C. bucerotis (figs. 1-7). In the female the ventral sclerite between the vulva and anus is elevated medially between the sclerital hooks in C. fasciati, C. bucerotis, and C. robusta, is elevated only slightly in C. camuri, and is not elevated in C. lophocerus. The female anal fringe has the most setae in C. fasciati and the fewest in C. lophocerus. Clay (1958) treated populations of the ischnoceran genus Degeeriella as subspecies when the male genitalia were apparently identical or differed only in a minor degree and when there were other minor morphological differences. Because of similarity of the genitalia, Chapinia fasciati and C. camuri could be considered subspecies of C. lophocerus. This would express the similarity of their six host species, which are all members of the Ethiopian genus Tockus. Clay (1958) pointed out that the genitalia, particularly in the Amblycera, might show only minor differences throughout a genus or species-group and great differences in other groups. She therefore concluded that differentiation of the genitalia has taken place at different rates in different groups. Similarly, Johnson (1960) stated that evolution and morphological divergence would not be expected to proceed at the same rate for all free-living species. It would seem that the similarity in the genitalia of C. fasciati, C. lophocerus, and C. camuri would indicate either that evolution has not proceeded as rapidly in these species or that they have not been isolated as long as other species of Chapinia. Clay (1958) stated that if subspecies were populations that would interbreed under natural conditions if they occurred sym- patrically, any morphological differences which might prevent inter- breeding should be considered as specific characters. Johnson (1960) believes that there is little possibility of finding interbreeding popu- lations among lice which are isolated on their hosts. She stated that it would be desirable to treat all stable recognizable forms of Anoplura and Mallophaga as species. Clay (1962) consequently stated that the subspecific category might be useful in some of the ischnoceran genera, but its application in the Amblycera is less satisfactory and 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 should not be used until more is known about the relationships between populations in this superfamily. The members of the lophocerus species-group are arranged according to the phylogeny of their hosts, since this order agrees with the morpho- logical similarities or the parasites. Chapinia fasciati, new species Fiegure 1 Male: Smaller than Chapinia robusta in all measurements; smaller than CO. lophocerus in all measurements except width of metathorax (table 1). Venter of third femora and posterolateral margins of abdominal sternites IV-VI each with small scattered brushes of normal setae. Each lateral margin of abdominal tergites II-VI with a short seta between the spiracle and postspiracular seta. Abdominal sternite II with 32-42 total setae and one median row of setae on posterior margin. ‘Terminal abdominal segments similar to those of C. lophocerus. Genitalia as shown in figure 1, each lateral horn with two rounded posterior points. Female: Larger than Chapinia robusta in all measurements except width of metathorax; smaller than C. lophocerus in all measurements except total length and width of prothorax (table 2). Resembles the male except that abdominal sternite II has 54 total setae. Terminal abdominal segments similar to those of C. lophocerus. Ventral sclerite between vulva and anus elevated medially between sclerital hooks. Anal fringe with 72-86 setae. Discussion: Chapinia fasciati resembles most closely C. lophocerus. The male genitalia have each lateral horn possessing two posterior points which are rounded in C. fasciati but sharp in C. lophocerus. The ventral sclerite between female vulva and anus is elevated medially between the sclerital hooks in C. fasciati but not elevated in C. lophocerus. The female anal fringe has more than 70 setae in C. fasciatt but fewer than 58 setae in C. lophocerus. Material examined: 27 males and 34 females from fresh and dried material collected in the Ethiopian region. Type host: Tockus fasciatus fasciatus (Shaw, 1811). Type material: Holotype male and allotype female from Eden, French Cameroons, Africa, collected by J. Mouchet, BMNH. Para- types: 18 males and 15 females from French Cameroons, Africa, collected by J. Mouchet, BMNH; 1 male from Kasongo, Belgian Congo, Africa, Nov. 13, 1959, collected by P. L. G. Benuit, JT; 3 males and 10 females from CNHM skins from Entebbe, Uganda, Africa, 1895-1916, collected by F. J. Jackson and others, REE; 2 males and 1 female from CNHM skins from Bitya, Cameroons, NO. 3558 MALLOPHAGA—ELBEL 153 Africa, 1924-1927, collected by O. L. Bates, REE; 1 female from CNHM skin from Yokadouma, French Cameroons, Africa, Oct. 19, 1946, collected by A. I. Good, REE; 1 female from CNHM skins from Bwamba, Ruwenzori, Uganda, Africa, 1944-1946, collected by V. Someren, REE; 1 male from CNHM skins from Ebolowa, French Cameroons, Africa, 1952-1953, collected by A. I. Good, REE; 1 male and 3 females from USNM skins from Congo, Africa, 1917, collected by C. R. Aschemeier, REE; from Tockus alboterminatus suahelicus (Neumann, 1905): 1 female from CNHM skins from Sokoke Forest, Kenya, Africa, June 1932, collected by V. Someren, REE; 1 female from USNM skins from Nairobi, Kenya, Africa, 1909, collected by Loring and Mearns, REE. Chapinia lophocerus (Bedford) Fiaures 2, 4, 27, 28 Menopon lophocerus Bedford, 1920, p. 717, pls. 1 (fig. 1), 3 (fig. 1). [Type host: Lophoceros leucomelas= Tockus flavirostris leucomelas (Lichtenstein, 1842).] Chapinia lophocerus (Bedford)—Hopkins and Clay, 1952, p. 67. Bedford did not designate a type from his material which contained a pair of Mallophaga from Lophoceros leucomelas= Tockus flavirostris leucomelas (Lichtenstein, 1842), a pair from LZ. epirhinus=Tockus nasutus caffer (Sundevall, 1851), and two males and one female from L. erythrorhynchos=Tockus e. erythrorhynchus (Temminck, 1823). A lectotype was designated by Hopkins (1941) from the host, Lophoceros leucomelas, since the male from that host agreed best with Bedford’s figure of the male genitalia. Male: Smaller than Chapinia robusta in all measurements (table 1). Venter of third femora and posterolateral margins of abdominal sternites IV-VI each with small scattered brushes of normal setae. Each lateral margin of abdominal tergites II-VI with a short seta between the spiracle and postspiracular seta. Abdominal sternite II with 28-42 total setae and one median row of setae on posterior margin. Terminal abdominal segments as shown in figure 4. Genitalia as shown in figure 2, each lateral horn with two sharp posterior points which do not reach the slender endomeres. Female: Larger than Chapinia robusta in all measurements except width of prothorax and width of metathorax (table 2). Resembles the male except that abdominal sternite II has 34-54 total setae. Terminal abdominal tergite with 22-30 setae, short setae alternating with long, on posterior margin; ventral sclerite between vulva and anus not elevated medially between sclerital hooks (fig. 27). Anal fringe with 42-56 setae (fig. 28). 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Discussion: Bedford (1920) gave the following measurements (inmm). male female length of head 0.25 0.33 width of head 0.53 0.60 width of prothorax 0.38 0.43 width of metathorax 0.55 0.71 width of abdomen 0.85 1.15 total length 1.74 2.36 Except for females being larger in abdominal width and total length, these measurements fall within the ranges of Chapinia lophocerus (tables 1, 2). ©. lophocerus resembles most closely C. fasciatt. The venter of the third femora and posterolateral margins of abdominal sternites [V—-VI each have small scattered brushes of normal setae in C. lophocerus, C. fasciati, and C. camuri but large thick brushes of normal setae in C. bucerotis. Abdominal sternite II of both sexes has fewer than 58 total setae and one median row of setae in C. lophocerus, C. fasciati, and C. camurt but more than 58 total setae and three median rows of setae in C. bucerotis. ‘The male genitalia have each lateral horn possessing posterior points that do not reach the slender endomeres in C. lophocerus, C. fasciati, and C. camuri but one sharp point that crosses the broad endomeres in C. bucerotis; the posterior two points are sharp in C. lophocerus but rounded in C. fasciati, and there is only one rounded posterior point in C. camurt. The ventral sclerite between female vulva and anus is not elevated medially between the sclerital hooks in C. lophocerus, is elevated slightly in C. camurt, but is more elevated in C. fasciati and C. bucerotis. The female anal fringe has fewer than 58 setae in C. lophocerus, at least 72 setae in C. fasciati, and 58-72 setae in C. camuri and C. bucerotis. Material examined: 6 males and 19 females from fresh and dried material collected in the Ethiopian region; lectotype male and syntype female from Transvaal, South Africa, September 1917, collected by G. A. H. Bedford, GHEH; from the type host: 1 male and 4 females from Pretoria Zoo, South Africa, Feb. 10, 1938, collected by G. A. H. Bedford, GHEH; from Tockus n. nasutus (Linnaeus, 1766): 1 male and 1 female from Maroua, North French Cameroons, Africa, 1959, collected by J. Mouchet, BMNH 1960-105; 1 male and 2 females from Manséa, Portuguese Guinea, Africa, Feb. 14, 1951, collected by J. Tendeiro, JT; from Tockus e. erythrorhynchus (Temminck, 1823): 1 male and 4 females from Somaliland, Africa, February 1949, Meinertz- hagen 18708, BMNH; 1 male from USNM skins from Ethiopia, Africa, 1912, collected by Childs Frick, REE; from TYockus f. fla- virostris (Riippell, 1835): 1 female from USNM skins from Ethiopia, Africa, 1912, collected by Childs Frick, REE; from Tockus deckeni (Cabanis, 1869): 6 females from Koka, Ethiopia, Africa, Dec. 13, 1960, collected by Savo Brelih, PMS. NO. 3558 MALLOPHAGA—ELBEL 1 5 Drawings were made of the lectotype male and a female collected in the Pretoria Zoo. Specimens in GHEH. Chapinia camuri, new species FIGURE 3 Male: Smaller than Chapinia robusta in all measurements; smaller than C. lophocerus in all measurements except length of head and width of prothorax (table 1). Venter of third femora and posterolateral margins of abdominal sternites IV-VI each with small scattered brushes of normal setae. Each lateral margin of abdominal tergites II-VI with a short seta between the spiracle and postspiracular seta. Abdominal sternite II with 37 or 38 total setae and one median row of setae on posterior margin. ‘Terminal abdominal segments similar to those of C. lophocerus. Genitalia as shown in figure 3, each lateral horn with one rounded posterior point. Female: Smaller than other Chapinia in all measurements except length of head and width of abdomen (table 2). Resembles the male except that abdominal sternite II has 44-56 total setae. ‘Terminal abdominal segments similar to those of C. lophocerus. Ventral sclerite between vulva and anus slightly elevated medially between sclerital hooks. Anal fringe with 60-64 setae. Discussion: Although smaller in size, Chopinia camuri resembles most closely C. lophocerus. The male genitalia have each lateral horn possessing one rounded posterior point in C. camurt but two sharp pos- terior points in C. lophocerus. The ventral sclerite between female vulva and anus is elevated medially only slightly between the sclerital hooks in OC. camuri and is not elevated in C. lophocerus. The female anal fringe has more than 58 setae in C. camuri but fewer than 58 setae in C. lophocerus. Material examined: 2 males and 2 females from fresh material col- lected in the Ethiopian region. Type host: Tockus camurus camurus Cassin, 1857. Type material: Holotype male and allotype female from Ambam, French Cameroons, Africa, 1955, collected by J. Mouchet, BMNH. Paratypes: 1 male and 1 female with same data. Chapinia bucerotis (Kellogg) Ficures 5, 6, 29, 30 Menopon bucerotis Kellogg, 1908, p. 54, pl. 7 (fig. 12) Bedford, 1920, pl. 3 (fig. 2) (male genitalia). [Type host: Bycanistes cristatus=Bycanistes brevis omissus Peters, 1945.] Chapinia bucerotis (Kellogg).—Hopkins and Clay, 1952, p. 67. Kellogg did not designate a type from his material, which contained 1 male and 1 female syntypes on slides and about 40 syntypes in 16 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 120 alcohol. The slide specimens were remounted and the male is desig- nated hereby as a lectotype; the slide has been so labeled. Approxi- mately one-half of the syntype material formerly in alcohol was mounted. Male: Smaller than Chapinia robusta in all measurements (table 1). Venter of third femora and posterolateral margins of abdominal sternites [V—VI each with large thick brushes of normal setae. Each lateral margin of abdominal tergites II-VI with a short seta between the spiracle and postspiracular seta. Abdominal sternite II with 60-82 setae and three median rows of setae. Terminal abdominal segments as shown in figure 5. Genitalia as shown in figure 6, each lateral horn with two sharp posterior points, one of which crosses the broad endomeres. Female: Approximately the same size as Chapinia robusta (table 2). Resembles the male except that abdominal sternite II has 68-104 total setae. Terminal abdominal tergite with 22-32 setae, short setae alternating with long, on posterior margin; ventral sclerite between vulva and anus elevated medially between sclerital hooks (fig. 29). Anal fringe with 58-72 setae (fig. 30). Discussion: Kellogg (1908) gave the following measurements (inmm). male Jemale length of head 0.33 0.40 width of head 0.65 0.72 width of abdomen 0.80 1.10 total length 2.00 2.80 These measurements are slightly larger than those given here for Chapinia bucerotis (tables 1, 2). C. bucerotis resembles most closely C. lophocerus. The venter of the third femora and posterolateral margins of abdominal sternites IV-VI each have large thick brushes of normal setae in C. bucerotis but small scattered brushes of normal setae in other species of the lophocerus species-group. Each lateral margin of abdominal tergites II-VI has a short seta between the spiracle and postspiracular seta in C. bucerotis but not in C. robusta. Abdominal sternite II of both sexes has more than 58 total setae in C. bucerotis but fewer than 58 total setae in C. lophocerus; this sternite has three median rows of setae in C. bucerotis but one median row of setae on posterior margin in other species of the lophocerus species- group. The male genitalia have each lateral horn possessing two sharp posterior points in C. bucerotis and C. lophocerus, but one point crosses the broad endomeres in C. bucerotis, and the points do not reach the slender endomeres in C. lophocerus; each lateral horn in C. robusta has one sharp median point that crosses the broad endomeres. The ventral sclerite between female vulva and anus is elevated medially between the sclerital hooks in C. bucerotis almost NO. 3658 MALLOPHAGA—ELBEL 7 as much as in C. robusta but is not elevated in C. lophocerus. The female anal fringe has at least 58 setae in C. bucerotis but at most 56 setae in C. lophocerus. Material examined: 34 males, 29 females, and approximately 20 specimens in alcohol from fresh and dried material collected in the Ethiopian region; lectotype male and syntypes, 12 males, 13 females, and about 20 syntypes in alcohol from Kilimanjaro, Tanganyika, Africa, collected by Sjéstedt, SMNH; from Bycanistes bucinator sharpit (Elliot, 1873): 1 female from CNHM skin from Mount Tandan, Mouila, Gabon, Africa, June 9, 1951, collected by H. A. Beatty, REE; 1 female from USNM skins from Congo, Africa, 1917- 1918, collected by C. R. Aschemeier, REE; from Bycanistes bucinator duboisi x sharpii: 1 female from CNHM skin from Yaounde, French Cameroons, Africa, July 12, 1948, collected by A. I. Good, REE; from Bycanistes bucinator duboisi W. Sclater, 1884: 1 male from CNHM« skins from Elat, French Cameroons, Africa, collected by Rev. M. Fraser, REE; from Bycanistes 6. bucinator (Temminck, 1824): 1 male and 1 female from Pietermaritzburg, South Africa, 1917, GHEH; 3 males and 2 females from CNHM skins from Kenya, Africa, 1918-1922, collected by V. Someren, REE; from Bycanistes c. cylindricus (Temminck, 1831): 1 male and 1 female from CNHM skins from Liberia, Africa, February—June 1948, collected by H. A. Beatty, REE; from Bycanistes cylindricus albotibialis (Cabanis and Reichenow, 1877): 2 males and 1 female from Mbalmayo, French Cameroons, Africa, collected by J. Mouchet, BMNH; 1 male from CNHM< skin from French Cameroons, Africa, July 8, 1907, REE; 1 male from CNHM< skin from French Cameroons, Africa, June 25, 1940, collected by A. I. Good, REE; 1 male from CNHM skin from Uganda, Africa, July 15, 1945, collected by V. Someren, REE; from Bycanistes subcylindricus subquadratus Cabanis, 1880: 6 males and 6 females from Uganda, Africa, April 1936, Meinertzhagen 7674, 7708, 7709, BMNH; 1 male from CNHM skin from Kampala, Uganda, Africa, Sept. 2, 1918, collected by V. Someren, REE; 1 male and 2 females from USNM skins from Uganda, Africa, June 1920, collected by H. C. Raven, REE; from Bycanistes b. brevis Friedmann, 1929: 2 males from CNHM skin from Mount Kenya, Kenya, Africa, Novem- ber 1946, collected by V. Someren, REE. Drawings were made of the lectotype male and the syntype female mounted on the same slide. Specimens in SMNH. Chapinia robusta Ewing Fiaurss 7, 23, 24, 31, 32 Chapinia robusta Ewing, 1927, p. 89. [Type host: Ceratogymna atrata (Temminck, 1835).] Chapinia robusta Ewing,—Hopkins and Clay, 1952, p. 68. 221-522—67——_2 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Through the courtesy of Dr. K. C. Emerson, BMNH specimens from the type host, here examined, were determined to be conspecific with the USNM holotype and allotype (USNM 40137, Nytonga, Congo, Africa, Nov. 3, 1917, collected by E. A. Chapin). Male: As illustrated in figure 24. Larger than other species of the lophocerus species-group in all measurements except width of abdomen; approximately the same size as Chapinia traylori (table 1). Venter of third femora and posterolateral margins of abdominal sternites IV-VI each with small scattered brushes of normal setae. Each lateral margin of abdominal tergites II-VI without a short seta between the spiracle and postspiracular seta. Abdominal sternite II with 48-66 total setae and one median row of setae on posterior margin. Terminal abdominal segments as shown in figure 24c. Genitalia as shown in figure 7, each lateral horn with one large sharp point which crosses the broad endomeres. Female: As illustrated in figure 23. Larger than Chapinia camuri in all measurements except length of head; approximately the same size as other species of the lophocerus species-group; slightly smaller than C. traylori in all measurements except length of head (table 2). Resembles the male except that abdominal sternite II has 56-68 total setae. ‘Terminal abdominal tergite with 20-24 setae, short setae alternating with long, on posterior margin; ventral sclerite between vulva and anus elevated medially between sclerital hooks (fig. 31). Anal fringe with 56-62 setae (fig. 32). Discussion: Ewing (1927) gave the following measurements (in mm). male female width of abdomen 0.70 0.95 total length 1.59 2.00 These measurements are not as great as those given here for Chapinia robusta (tables 1, 2). C. robusta resembles most closely C. bucerotis. The venter of the third femora and posterolateral margins of abdom- inal sternites IV-VI each have small scattered brushes of normal setae in C. robusta, slightly more numerous than for other species of the lophocerus species-group, except C. bucerotis, which has large thick brushes of normal setae. Each lateral margin of abdominal tergites II-VI in C. robusta lacks the short seta between the spiracle and postspiracular seta that is present in all other species of the lophocerus species-group. Abdominal sternite II has one median row of setae in C. robusta but three median rows of setae in C. bucerotis. Male geni- talia have each lateral horn possessing one sharp median point that crosses the broad endomeres in C. robusta but two sharp posterior points, one of which crosses the broad endomeres, in C. bucerotis. The ventral sclerite between female vulva and anus is elevated NO. 3558 MALLOPHAGA—ELBEL 19 medially between the sclerital hooks slightly more in C. robusta than in C. bucerotis. Material examined: 4 males and 5 females from fresh and dried material collected in the Ethiopian region; from the type host: 3 males and 3 females from Ambam, French Cameroons, Africa, 1955, collected by J. Mouchet, BMNH; 1 female from CNHM skin from Fougamou, Gabon, Africa, Aug. 4, 1951, collected by H. A. Beatty, REE; from Ceratogymna elata (Temminck, 1831): 1 male and 1 female from Konn, French Cameroons, Africa, Apr. 26, 1947, collected by V. Aellen, BMNH 1954-487. Drawings were made of a male and a female from the type host collected in Ambam, French Cameroons, Africa. Specimens in BMNH. The acutovulvata Species-Group Species similar in shape to Chapinia traylori (figs. 25, 26). Differing from other species-groups in the following combination of characters: Dorsal-lateral margins of head with a preocular notch; venter of third femora and posterolateral margins of abdominal sternites IV—VI each with or without brushes of normal setae; each lateral margin of ab- dominal tergites II-VI with or without a short seta between the spiracle and postspiracular seta; females usually with more abdominal sternal setae than males; abdominal sternite II usually with one median row of setae on posterior margin; male genitalia without lateral horns on each side of endomeres and with parameres enlarged anteriorly, split posteriorly; females without sclerital hooks on each side of midline of ventral sclerite between vulva and anus; female abdominal sternite VIII with most of setae much shorter than those on the posterior margin, Hosts: Species of the acutovulvata species-group have been found on the genera Tockus, Anorrhinus, Penelopides, Rhyticeros, Anthra- coceros, and Buceros of the avian family Bucerotidae. Species of the acutovulvata species-group are all similar in size except that males of Chapinia wenzeli are smaller than other males, and females of C. traylori are larger than other females. The venter of the third femora and posterolateral margins of abdominal sternites IV-—VI each have large thick brushes of normal setae in both sexes of C. waniti, C. acutovulvata, C. malayensis, and C. hoplai; small scattered brushes of normal setae in females of CO. blakei and in both sexes of C. clayae, C. boonsongi, C. wenzeli, and C. lydae; brushes are absent in the male of C. blakei and in both sexes of C. traylort. Each lateral margin of abdominal tergites JI-VI has a short seta between the spiracle and postspiracular seta in C. clayae, C. waniti, C. acutovulvata, 20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 and C. malayensis. Abdominal sternite II has more total setae in both sexes of C. waniti, C. acutovulvata, and C. hoplar than in corre- sponding sexes of CO. malayensis, C. wenzeli, C. blakei, and C. traylori; more total setae in females than in males except in C. lydae; and setae are arranged in one median row except for one or two median rows in C. waniti, two median rows in C. traylori, and three median rows in C. lydae. The male genitalia have endomeres with an inner plate and paired outer rims in C. clayae, C. waniti, C. acutovulvata, C. malayensis, and C. lydae, the paired outer rims possessing internal knobs only in C. clayae and C. acutovulvata and serrulations only in C. waniti; the inner plate possessing paired lateral flanges and central terminal point only in C. malayensis; only paired outer rims in C. hoplai, OC. boonsongi, and C. wenzeli, the posterolateral margin being split in C. wenzeli; and only paired plates in C. blake and C. traylori. The female terminal abdominal tergite has on the posterior margin at most 24 setae except for C. waniti, C. acutovulvata, and C. hoplar with at least 24 setae; of these setae approximately two-thirds are long and one-third are short in C. wanitv, C. acutovulvata, C. malayensis, C. boonsongi, and C. lydae; approximately one-half are long and one- half are short in C. clayae, and C. hoplai; these setae are arranged with at least five long setae on each side of the midline in C. acutovulvata and C. malayensis but at most six setae in all other species of the acutovulvata species-group. In the female the ventral sclerite be- tween the vulva and anus is curved sharply on the anterior margin in C. clayae and C. acutovulvata but is only slightly curved in the other species of the group; on each lateral projection of this sclerite there are 4-6 thick, posteriorly directed setae except in C. waniti which has 10 or 11 and in C. traylori which has 2 or 3 (figs. 43 sp, 44 sp, and 72 sp). The female abdominal sternite VIII has on the posterior margin the most setae in C. malayensis with more than 34 and the fewest in CO. blakei with at most 18. The female anal fringe has at least 44 setae in C. waniti, C. acutovulvata, C. malayensis, C. hoplar, C. boonsongi, and C. lydae but at most 44 setae in C. clayae, C. wenzeli, C. blakei, and C. traylori. Were the species of the acutovulvata species-group arranged according to the phylogeny of their hosts (Peters, 1945), the order would be: C. clayae, C. waniti, C. wenzeli, C. blakei, C. lydae, C. boonsongi, C. malayensis, C. acutovulvata, C. hoplai, and C. traylori, rather than C. clayae, C. waniti, C. acutovulvata, C. malayensis, C. hoplai, C. boonsongi, OC. wenzeli, CO. blakei, C. traylori, and C. lydae, which is based on morphological similarities of the lice. NO. 3558 MALLOPHAGA—ELBEL 21 Chapinia clayae, new species Fiaures 8, 33, 34 Both sexes are smaller than corresponding sexes of Chapinia traylori in all measurements except length of head (tables 1, 2). Male: Venter of third femora and posterolateral margins of abdom- inal sternites [V—VI each with small scattered brushes of normal setae. Each lateral margin of abdominal tergites II-VI with a short seta between the spiracle and postspiracular seta. Abdominal sternite IT with 38-50 total setae and one median row of setae on posterior margin. Terminal abdominal segments similar to those of C. acuto- vulvata. Genitalia as shown in figure 8, endomeres with small inner plate and paired outer rims. Female: Resembles the male except that abdominal sternite II has 66-76 total setae. ‘Terminal abdominal tergite with 12 long and 12 short setae on posterior margin; abdominal sternite VIII with 22-30 setae on posterior margin (fig. 33). Anal fringe with 38-44 setae (fig. 34). Discussion: Although smaller in size, Chapinia clayae resembles most closely C. acutovulvata. The venter of the third femora and posterolateral margins of abdominal sternites IV-VI each have small scattered brushes of normal setae in C. clayae but large thick brushes of normal setae in C. acutovulvata. The male genitalia have endomeres possessing on each outer rim a triangular internal knob that is wider than long in C. clayae but longer than wide in OC. acutovulvata. The female terminal abdominal tergite has on the posterior margin at most 24 setae, of which approximately one-half are long and one-half are short, in C. clayae, but at least 26 setae, of which approximately two-thirds are long and one-third are short, in C. acutovulvata. In addition this margin has on each side of the midline at most four long setae in C. clayae but at least six long setae in C. acutovulvata. The female anal fringe has at most 44 setae in C. clayae but at least 44 setae in C. acutovulvata. Material examined: 19 males and 29 females from fresh and dried material collected in India and Nepal. Type host: Tockus birostris (Scopoli, 1786). Type material: Holotype male, allotype female, and 2 male and 4 female paratypes on same slide from Rajputana, India, March 1937, Meinertzhagen 8855-8856, BMNH. The holotype and allotype are each the second from the right in the rows of males and females as seen under the microscope. Paratypes: 5 males and 12 females with same data except Meinertzhagen 8932; 1 male from Nepal, 22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 December 1935, Meinertzhagen 4859, BMNH; 4 males and 5 fe- males from Nepal, February 1936, Meinertzhagen 4858, BMNH; 1 female from CNHM skins from Kotla, Kangra, East Punjab, India, 1946 and 1948, collected by W. Koelz, REE; 1 male and 1 female from CNHM skins from Bheraghat, Central Provinces, India, Mareh- April 1946, collected by W. Koelz and R. Chand, REE; 1 male from CNHM < skins from Belwani, Kisli, Central Provinces, India, July— August 1946, collected by W. Koelz, REE; 1 female from CNHM skin from Kanha, Central Provinces, India, Aug. 29, 1946, collected by R. Chand, REE; 2 males from CNHM skins from Kalnali, United Provinces, India, February 1947, collected by W. Koelz, REE; 1 male and 1 female from CNHM skins from Nichland, United Provinces, India, February 1947, collected by W. Koelz, REE; 1 female from CNHM skins from Simra, Nepal, Mar. 4, 1947, collected by W. Koelz and R. Chand, REE; 1 female from CNHM skins from Baihar, Balaghat, India, January—February 1949, collected by R. L. Flem- ming, REE; 1 female from USNM skins from India, 1898, 1946-1948, REE; from Tockus g. griseus (Latham, 1790): 1 male from CNHM skins from Nilambus, Madras, India, February—March 1937, collected by W. Koelz, REE. Chapinia clayae is named for Dr. Theresa Clay of the British Museum (Natural History) in appreciation for her continuous assist- ance throughout the study, for the loan of hornbill Menoponidae from the BMNH, and for helping to obtain the loan of hornbill Menoponidae from other museums. Chapinia waniti, new species Figures 71, 72 Male: Larger than Chapinia traylori in all measurements except width of prothorax and width of metathorax (table 1). Venter of third femora and posterolateral margins of abdominal sternites IV-VI each with large thick brushes of normal setae. Each lateral margin of abdominal tergites II-VI with a short seta between the spiracle and postspiracular seta. Abdominal sternite II with 68-78 total setae and one or two median rows of setae. Terminal abdominal segments similar to those of C. acutovulvata. Genitalia as shown in figure 71, endomeres with small inner plate and paired outer rims. Female: Smaller than Chapinia traylori in all measurements except length and width of head (table 2). Resembles the male except that abdominal sternite II has 92-104 total setae. Terminal abdominal tergite with 14-18 long and 10-12 short setae on posterior margin; abdominal sternite VIII with 28-34 setae on posterior margin (fig. 72) ; each lateral projection of the ventral sclerite between vulva and anus NO. 3658 MALLOPHAGA—ELBEL 23 with 10-11 thick, posteriorly directed setae (fig. 72 sp). Anal fringe similar to that of C. acutovulvata, with 56-64 setae. Discussion: Chapinia waniti resembles most closely C. aeutovulvata. Abdominal sternite II of both sexes has more total setae in C. wamniti than in corresponding sexes of C. acutovulvata. The male genitalia have endomeres possessing on each outer rim serrulations on the posterior inner margin in C. waniti but a triangular internal knob in C. acutovulvata. Each lateral projection of the ventral sclerite between female vulva and anus has more than eight thick, posteriorly directed setae in C. waniti but fewer than eight in all other Chapinia. The female anal fringe has at least 56 setae in C. waniti but at most 54 setae in C. acutovulvata. Material examined: 8 males and 7 females from fresh material collected in Thailand. Type host: Anorrhinus galeritus carinatus (Blyth, 1845). Type material: Holotype male and allotype female from Chong, Muang, Trang, Thailand, Mar. 4, 1963, collected by Wichit Suwan Laong, USNM. Paratypes: 6 males and 3 females with same data; 1 male and 1 female from Lamo, Muang, Trang, Thailand, Mar. 3, 1963, collected by Wichit Suwan Laong, USNM; 2 females from Na Wong, Muang, Phatthalung, Thailand, Mar. 6, 1963, collected by Wichit Suwan Laong, USNM. Chapinia waniti is named for Mr. Wanit Songprakob, Songkhla, Thailand, in appreciation for mounting Mallophaga and for directing the activities of the field collector, Wichit Suwan Laong. After my departure from Thailand in April 1963, both boys collected for the Bernice P. Bishop Museum. Chapinia acutovulvata (Piaget) Fiaures 9, 10, 35, 36 Menopon aculovulvatum Piaget, 1881, p. 5, pl. 1 (fig. 4). [Type host: Buceros malabaricus= Anthracoceros a. albirostris (Shaw, 1808).] Menopon acutovulvatum Piaget, 1885, p. 106, pl. 11 (fig. 8). Allomenopon mjébergi Eichler, 1947, pp. 2, 20, figs. 1, 2 (new synonym). [Type host: Anthracoceros converus (Temminck, 1831).] Chapinia mjébergi (Hichler)—Hopkins and Clay, 1952, p. 68. Chapinia acutovulvata (Piaget)—Hopkins and Clay, 1952, p. 67. Dr. Eichler’s specimens of Chapinia mjébergi from Anthracoceros convezus in the Zoological Museum, Humboldt University, Berlin, were loaned through the courtesy of Dr. von Kéler. Comparison of these lice with specimens of Chapinia acutovulvata from the type host discloses no morphological differences between the two series. A lectotype male for Chapinia acutovulvata was designated by Clay (1949a) from the Piaget collection, now in BMNH, BM 777, with 6 syntype females, BM 774 and 776. 24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Both sexes are smaller than corresponding sexes of Chapinia traylori in all measurements except length and width of head (tables D2). ies Venter of third femora and posterolateral margins of abdom- inal sternites 1V-VI each with large thick brushes of normal seta. Each lateral margin of abdominal tergites II-VI with a short setae between the spiracle and postspiracular seta. Abdominal sternite IT with 42—54 total setae and one median row of setae on posterior margin. Terminal abdominal segments as shown in figure 10. Genitalia as shown in figure 9, endomeres with small inner plate and paired outer rims. Female: Resembles the male except that abdominal sternite II has 76-86 total setae. Terminal abdominal tergite with 16-22 long and 10-12 short setae on posterior margin; abdominal sternite VIII with 18-30 setae on posterior margin (fig. 35). Anal fringe with 44-54 setae (fig. 36). Discussion: Although larger in size, Chapinia acutovulvata resembles most closely C. clayae. The venter of the third femora and postero- lateral margins of abdominal sternites IV-VI each have large thick brushes of normal setae in C. acutovulvata but small scattered brushes of normal setae in C. clayae. Abdominal sternite II of both sexes of C. acutovulvata has fewer total setae than in corresponding sexes of C. waniti but has more total setae than in corresponding sexes of C. malayensis. The male genitalia have endomeres possessing on each outer rim a triangular internal knob that is longer than wide in C. acutovulvata but wider than long in C. clayae. ‘These internal knobs are absent in other Chapinia. The female terminal abdominal ter- gite has on the posterior margin at least 26 setae in C. acutovulvata, of which approximately two-thirds are long and one-third are short, but at most 24 setae in C. clayae and C. malayensis, of which approx- imately one-half are long and one-half are short in C. clayae, but approximately two-thirds are long and one-third are short in C. malayensis. In addition this margin has on each side of midline at least six long setae in C. acutovulvata but at most five long setae in C. clayae. ‘The female abdominal sternite VIII has on the posterior margin fewer than 32 setae in C. acutovulvata but more than 34 setae in C. malayensis. Each lateral projection of the ventral sclerite between female vulva and anus has fewer than eight thick, posteriorly directed setae in C. acutovulvata, but more than eight in C. waniti (fig. 72 sp.). The female anal fringe has 44-54 setae in C. acuto- vulvata, at most 44 setae in C. clayae, and at least 56 setae in C. wanite. Material examined: 33 males and 35 females from fresh and dried material collected in the Oriental region; from the type host: 2 females, Piaget, BMNH 1953-21; 2 males and 2 females from Nepal, NO. 3558 MALLOPHAGA—ELBEL 25 December 1935, Meinertzhagen 4872, BMNH; from Anthracoceros albirostris leucogaster (Blyth, 1841): 2 males and 2 females from Myitkyina, Burma, Mar. 26, 1945, collected by the U.S. Typhus Com- mission, BMNH; 3 males, 1 female with same data, USNM; 2 males and 1 female from Stillwell Road, Myitkyina, Burma, Sept. 26, 1945, collected by H. S. Fuller, BMNH 1947-321 (164); 2 males, 2 females with same data, USNM; 2 males from Hin Laem, Tha Khanun, Kanchanaburi, Thailand, Nov. 27, 1952, collected by Robert E. Elbel and H. G. Deignan, USNM; 2 males from Ban Khlua Klang, Prachuap Khiri Khan, Thailand, December 1952, collected by Robert E. Elbel and H. G. Deignan, USNM; 2 males and 1 female from Ban Nam Phu, Phu Khieo, Chaiyaphum, Thailand, Dec. 22, 1952, collected by Robert KE. Elbel, USNM; 1 male and 1 female from Ban Thung Chuak, Salok Bat, Khanu, Kamphaeng Phet, Thailand, June 25, 1953, collected by Robert E. Elbel, USNM; 1 female from Tha Din Daeng, Pa Bon, Pak Pha Yun, Phatthalung, Thailand, July 30, 1962, collected by Wichit Suwan Laong, USNM; 6 males and 6 females from Muang Kluang, Kapoe, Ranong, Thailand, 1962-1963, collected by Wichit Suwan Laong, USNM;6 males and 11 females from Pa Dong Lan, Chumphae, Khon Kaen, Thailand, Dec. 2, 1962, collected by Kitti Thonglongya, SMRL; from Anthracoceros coronatus (Boddaert, 1783): 2 females from CNHM< skins from Kanha, Central Provinces, India, August 1946, collected by Rup Chand, REE; 2 females from CNHM skins from Nawadeh, Bihas, India, Nov. 11, 1947, collected by W. Koelz, REE; 1 male from USNM skins from India and Ceylon, 1874 and 1944, collected by B. H. Swales and S. D. Ripley, REE; from Anthra- coceros convexus (Temminck, 1831): 1 male and 1 female from lot 1584 (TMRN), Zoological Museum, Humboldt University, Berlin; accord- ing to Kichler (1947) the Mallophaga WEC 2268 from this later named host were collected in Sumatra by E. Mjéberg; from » y ‘A uv ‘if Mm fa) a} vi A a fs epi bit ee : a] if year i nal oa fe al Cee pie Vepiaty-4getes Ving) eee ee bia tig ie’ Le ut ; wort sayy car tiv nM : 4 (tay CiaA) Wa healt ‘Sled i sai » Fie 7 ait he ae | ' : 8 ' a 1 ARATE w be aa (ft ty i rl eer \ bi phot an tl ; ; . o: 4 [5 01 ia im By fyibe. ainda ¢ 1) shay erat he Sat t inl f Ww! VPs hare hod apie ie ile (pty tie ‘4 | oe We ne, ea, ; heh fe 1 Mo@e phn ae Pe ; iT Aid et tee ayad we tatu fied) egy Adele Ah 120 eens oP dvi) : ana! en liye aoe met PMLA AS uate . vit ‘4 foment fhab¥ad ; Pe ile genet ; j A mp Magee) prety pul Men ils yt PRN a. - — yegbieeskis oie Maal ie : i ignvte’ Ae Mri Pirie) ie ie ne ee RL i jae Havbalaea lyr) ane in hat ate by ]. FaMngig ht at> (TAGS Pf VRS bd BO a. o 1 kU! 2a eid “hap eye edie WA ae oe f a fi ii? cops Ota heen See a wl ealfagomnty wife vhwadlpahid (he sath ie ai. Otay tiie Len. aren ln Anes Uae ie aie oy «i praia 5 mera me fre ) am! . ans Us 7 v oe i; ee 1 ae 1 >» oe! toe oi Vi Le in Ge a, Proceedings of the United States National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. Volume 120 1967 Number 3561 REVIEW OF SOME SPECIES OF LOXOSTEGE HUBNER AND DESCRIPTIONS OF NEW SPECIES (LEPIDOPTERA, PYRAUSTIDAE: PYRAUSTINAE) By Haun W. Capps! The purpose of this paper is to provide means for reliable identifi- cation of the species of Lozostege Hiibner, to correct invalid synonymy and nomenclature, and to provide names for undescribed species. Some nomenclature, following Hampson’s (1899, p. 207) error in treating Lozostege as a nomen nudum and synonym of Phlyctaenodes Guenée (1864, p. 173), needs correction. Although Hampson (1918, p. 189) corrected the error, the generic placement has not been cor- rected for those new species described in the interim from Mexico, Central and South America because there has been no subsequent revision involving them and because lists of American Lepidoptera published in that period have been restricted to North American species. In addition to the nomenclatural aspect, zoology is also involved. The type-species of Lozostege and Phlyctaenodes are not congeneric. Pyralis aeruginalis Hiibner, the type of Loxostege (Hampson, 1918, p. 189), differs from Pyralis pustulalis Hiibner, the type of Phlyctae- nodes (Hampson, 1899, p. 211), in maculation and genitalia as indi- cated below. 1 Retired, September 1964, Entomology Research Division, Agriculture Re- search Service, U.S. Department of Agriculture, Washington, D.C. il 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Venation.—Loxostege aeruginalis: Forewing (fig. 2) with vein 6 distinctly closer to vein 7 than to middle of cell; veins 8 and 9 long stalked, 10 approximate to 8+9 at cell and parallel for almost half the length of stem; vein 11 subparallel with 10 to costa; terminus of vein la upturned, obsolescent distally or if discernible, forming a loop with 1b. Hindwing (fig. 2a) essentially the same as in P. pustulalis. Phlyctaenodes pustulalis: Forewing (fig. 3) with origin of vein 6 closer to middle of cell than to vein 7; stalking of veins 8 and 9 much shorter than in L. aeruginalis; vein 10 strongly divergent from stalk of 8 and 9 a short distance from cell; vein la acutely divergent from vein 1b from base and terminating on inner margin of wing. Hind- wing (fig. 3a). MALE GENITALIA (see figs. 14 and 18 for terminology) —L. aerugi- nalis: Harpe with a well-developed, hooklike clasper; transtilla with ventrodistal extension conspicuous, narrow and extending to near base of anellus. Uncus with apex rounded distally. Gnathos absent. Phlyctaenodes pustulalis (fig. 5): Harpe unarmed; transtilla without ventrodistal extension. Uncus somewhat bilobed at apex. Gnathos well developed. This study is based, in part, upon more than 1800 dissections of genitalia prepared during the investigation. FEMALE GENITALIA (see fig. 7 for terminology)—L. aeruginalis (fig. 7): Bursa copulatrix with an accessory pouchlike appendage; signum well developed. P. pustulalis (fig. 4): Bursa copulatrix simple, without an accessory pouchlike appendage; signum absent. The Loxostege species helvialis (Walker), mancalis (Lederer), samilalis (Guenée), rantalis (Guenée), occidentalis (Packard), and a species described as Botys obliteralis Walker and heretofore assigned to Loxostege and Phlyctaenodes, are so similar to each other or closely related species in color, maculation, and other characters that exam- ination of the genitalia is often necessary for reliable identification. The delineations were prepared by Mr. A. D. Cushman, scientific illustrator, Entomology Research Division, Agric. Res. Serv., U.S. Department of Agriculture, and are not drawn to scale. The genitalia are in ventral view and those of the males are drawn with one harpe omitted and the aedeagus removed. All photos of adults, except one, are by J. Scott, staff photographer, — U.S. National Museum; figure 146 is used through courtesy of British | Museum (Nat. Hist.). Photos are about three times natural size. NO. 36561 LOXOSTEGE—CAPPS 3 Loxostege Hiibner Loxostege Hiibner, 1827, Verzeichniss bekannter Schmettlinge [sic], p. 352. [Type: Pyralis aeruginalis Hiibner, designation Hampson, 1918, Ann. Mag. Nat. Hist., ser. 9, vol. 2, p. 189.] Characters common to the Loxostege species treated here are dis- cussed below. Apuut.—Antenna simple. Frons usually pointed or conically produced, the modification sometimes limited to the lower margin and more discernible from dorsal view. Midtibia normal and without a hair-pencil, or incrassate and with a hair-pencil, often concealed in a groove. Hind tibia with two pairs of spurs (the relative lengths referred to herein are those of the upper pair of spurs on the hind tibia of the male). Larva.—Chaetotaxy and crochets are typical pyraustid. Pro- thorax with two setae on the prespiracular shield; group VI bisetose. Meso- and metathorax with VI unisetose. Abdominal setments 3-6 with seta IV approximate to seta V, and under spiracle on same pinaculum. Ninth abdominal segment with seta I approximate to seta III and on same pinaculum; setae IV and V absent, only seta VI of this group present. Crochets on abdominal prolegs of tri- ordinal length and arranged in a penellipse. Anal fork absent. Distribution, food plant, and flight data reported in this paper are taken entirely from labels accompanying the specimens examined. Loxostege helvialis (Walker) Ficures 8, 62, 105 Spilodes helvialis Walker, 1859, List of the lepidopterous insects in the British Museum (Nat. Hist.), pt. 18, p. 772. Botys thycealis Walker, 1859, ibid., pt. 19, p. 981. Botys apertalis Walker, 1865, ibid., pt. 34, suppl. 4, p. 1393. Botys citrina Grote, 1867, Trans. American Ent. Soc., vol. 1, p. 23. Loxostege helvialis (Walker) authors.—Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull. 52, p. 382.—Barnes and McDunnough, 1917, Check list of the Lepi- doptera of boreal America, p. 131.—Forbes, 1920, Cornell Univ. Agric. Exp. Sta. Mem. 68, p. 556 [as helvalis.—McDunnough, 1939, Mem. Southern California Acad. Sci., vol. 2, p. 12. Matz.—Alar expanse 18-20 mm. Frons with conical production strong. Antenna somewhat pubescent, cilia dense, very short, length of cilia less than one-half width of shaft. Midtibia slightly incrassate, hair-pencil obsolescent. Outer spur less than one-half as long as inner spur. Labial palpus length less than two times width of eye; first segment white; second segment slightly upturned, luteous 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 dorsally and laterally, white ventrally; third segment short, porrect or slightly downturned, luteous dorsally and laterally, white below. Head, thorax, and abdomen pale ochreous above, whitish below. Forewing: Upper surface pale yellow, somewhat shiny; area along costa, the reniform, discal dot, and two transverse lines luteous; subterminal line absent. ‘Transverse lines smooth, transverse an- terior line bent outwardly, concave or weakly angulate. Posterior transverse line directly posterior from costa to about vein 7, slightly concave outwardly to vein 3, angled inward to vein 2, outward to fold between veins 1b and 2, thence slightly inward to inner margin of wing. Hindwing: Upper surface ground color similar to that of forewing but paler from cell to costa; postmedial line appearing as a continuation of transverse line of forewing and terminating at about vein 2; subterminal line absent. Fringe of fore- and hindwings concolorous with ground color of wings. Under surface of wings paler than upper, with markings evident but weaker. Genitalia (fig. 8): Uncus with apex bluntly pointed. Harpe with simple hairlike scales; the sclerotized hooklike structure short, stout, bifid. Vinculum rather broadly concave midventrally. Aedeagus with an irregular, elongate patch of numerous spinelike cornuti and distally, a strong, single spine and a tapered, subcylindrical structure, the latter with circular grooves; basal keel absent or obsolescent. FEMALE (fig. 105).—Alar expanse 16-20 mm. Color and macula- tion like that of male. Genitalia (fig. 62) with sclerotization of ven- tral margin of ostium weak or absent; ductus bursae broad between ostium and origin of ductus seminalis; accessory pouch small. Larva.—Mature, 22 mm long. Head sordid white, with retic- ulate amber markings; fuscous pigmentation of lateral incision of hind margin inconspicuous or absent. Prothoracic shield sordid white, middorsal markings weak, pale amber; area adjacent to lateral margin darker and extending almost to base of seta Ib. Pinacula bearing setae Ia-Ib and IIJa—IIb of mesothorax separate, but slightly elevated. Pinacula bearing setae IJa-IIb of the meso- and meta- thorax and abdominal segments 1-8 with the sclerotization stronger and the brownish or fuscous pigmentation more extensive than those pinacula above or below. Ninth abdominal segment with pigmented area of pinaculum of seta II small and adjacent to base of the seta; pigmentation of pinaculum bearing setae I and III extensive, covering almost the entire area; pinaculum of seta VI nonpigmented. Typrs.—British Museum (Nat. Hist.): helvialis, female; thycealis, male; apertalis, male and female. American Museum of Natural History: citrina, female, hereby designated lectotype. NO. 3561 LOXOSTEGE—CAPPS 5 TYPE-LOCALITIES.— United States: helvialis and thycealis. North America: apertalis. ‘Phil.’ [Philadelphia, Pa.]; citrina. Foop PLANTS.—Amaranthus, beets. DISTRIBUTION.—UNITED STATES: Massachusetts: Barnstable, Mar- tha’s Vineyard. New York: Long Island (Bellfort, Orient). New Jersey: Lakehurst, Oakland. District of Columbia. Florida: Miami, Sarasota, Siesta Key, Tavernier, Craig. Texas: Brownsville, Black Jack Springs, Mercedes, San Antonio, San Benito, Big Bend Nat. Park, and Montague Co. Oklahoma: Norman. canapa: Ontario: Chatham. SPECIMENS EXAMINED.—72. In FiieHt.— March to October. REeMARKS.—Several species have heretofore been confused with helialis in collections. Six of these species are described as new herein, and some are reliably distinguished from helvialis only by examination of the genitalia. I am indebted to W. H. Tams and E. L. Martin of the British Museum (Nat. Hist.) for comparison of material with the Walker types; and to F. H. Rindge of the American Museum of Natural His- tory for information relative to the cotype of citrina. The male co- type of citrina could not be located in the material at the Philadelphia Academy of Sciences and appears to be lost; therefore, the female cotype at the American Museum of Natural History is designated lectotype of the species. Loxostege pseudohelvialis, new species Figures 9, 63, 106 Mats (fig. 106).—Alar expanse 17-20 mm. Frons and antenna as in helvialis, but male differs otherwise as follows: Midtibia more in- crassate, hair-pencil well developed. Outer spur of hind tibia shorter, about one-third as long as inner. Outer margin of fore- and hindwing darker, concolorous with markings of wings (in helvialis, concolorous with ground color of wings). Genitalia (fig. 9): Similar to those of helvialis but with apex of uncus more sharply pointed; arms of bifid hook slenderer and more angulate distally; vinculum narrower and more attenuate midventrally. Aede- agus similar to that of helvialis but more slender. Frmatr.—Alar expanse 17-21 mm. Similar to male in color and maculation. Outer spur one-half as long as inner spur. Genitalia (fig. 63) similar to those of helvialis but with ventral margin of ostium a narrow, somewhat crinkled sclerotized band; sclerotized portion of ductus bursae between ostium and origin of ductus seminalis shorter, narrower, lateral margins subparallel. 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Typr.—Male, U.S. National Museum, USNM 67595, genitalia slide OBP 290. Typr-LocaLity.—Baboquivari Mts., Pima Co., Ariz. PARATYPES.—UNITED sTaTEs: Arizona: type-locality, 8 o, 4 Q; Brown Canyon, Baboquivari Mts., 7 9; Madera Canyon, Santa Rita Mts., 33 o’, 28 9; Madera Canyon, Pima Co., 1 9; Paradise, Cochise Co., 1 &#; Tempe, 1 @, 1 9; Tucson, 1 o, 1 9; Scottsdale, Maricopa Co., 1 &@; Southwestern Res. Sta., Chiricahua Mts., 2 o, 1 9; Chiri- cahua Mts., 2 co, 1 9; White Mts., 1 9; Huachuca Mts., 2 9; Yavapai Co., 1 9; “Arizona” [no additional locality], 2 o, 3 9. California: San Diego, 7 o', 4 9; La Puerta Valley, 2 o, 3 9. Utah: Beaver, 1 @. New Mexico: Mesilla, 19. Texas: Limpia Canyon, Jeff Davis Co., 1 9. mexico: Sonora: Nainari, 1 o&, 5 9. Paratypes in the U.S. National Museum, Los Angeles County Museum, American Museum of Natural History, the collection of C. P. Kimball, the Canadian National Collection, and the British Museum (Nat. Hist.). Foop puant.—Unknown. IN FLIGHT.—June to September. Remarxs.—The L. pseudohelvialis series contains specimens with labels of both helvialis and citrina. If specimens are in good con- dition, the luteous suffusion along the outer margin of the fore- and hindwing distinguishes specimens of pseudohelvialis from those of helvialis. The more incrassate midtibia and hair-pencil of pseudo- helvialis also distinguish its males from those of helvialis, but exami- nation of the genitalia is necessary for reliable identification of worn females. Loxostege arizonensis, new species Figures 11, 64, 112 Mate (fig. 112).—Alar expanse 20-24 mm. Frons conical. An- tenna ciliate, length of cilia about one-half width of shaft. Midtibia distinctly incrassate, hair-pencil well developed. Outer spur slightly less than one-third as long as inner spur. Similar to pseudohelvialis in color and maculation, but the average size of arizonensis is larger and luteous suffusion along outer margin of fore- and hindwing is more extensive. Genitalia (fig. 11): Uncus pointed. Harpe with hairlike scales simple; padlike production above base of hook narrow, nonscobinate, but with a few simple setae, hooklike structure strong, curved; sacculus normally with a single spine about midway between cluster of spines and base of anellus. Vinculum narrowly attenuate mid- ventrally. Aedeagus with a conspicuous, thin, rounded basal keel and a dense cluster of spinelike cornuti distally. Fremate.—Alar expanse 19-24 mm. Similar to male in color and maculation. Genitalia (fig. 64) with midventral invagination of No. 3561 LOXOSTEGE—CAPPS re ostium divided and rather broad; ductus bursae narrow at origin of ductus seminalis. Typr.—Male, U.S. National Museum, USNM 67596, genitalia slide HWC 7725. Typr-LocaLiry.—Oracle, Ariz. PARATYPES.—UNITED STATES: Arizona: Baboquivari Mts., Pima Co., 9 o&', 4 2; Sabino Canyon, Santa Catalina Mts., Pima Co., 6 o, 8 9; Madera Canyon, Santa Rita Mts., 42 0, 55 9; Santa Rita Mts., 1 &'; Nogales, 4 6, 2 9; Palmerlee, 1 o&; Patagonia, 1 o&; Southwestern Res. Sta., Chiricahua Mts., Cochise Co., 1 &; “SWS” [Southwestern Res. Sta.], 19. mMepxico: Chiapas,1 o. Oaxaca,19. Mesquititlan, 1 9. Guerrero: Iguala, 1 9. Puebla: Tehuacan, 2 9. Sonora: Nainari, 1 o&,4 9. Paratypes in U.S. National Museum, American Museum of Natural History, Los Angeles County Museum, and the Canadian National Collection. Foop pirant.—Unknown. In FLicHT.—July to September in Arizona; May to June in Mexico. ReMARKS.—Labels on some specimens indicate that arizonensis has also been confused with helvialis. Loxostege neohelvialis, new species Figures 12, 66, 111 Mate (fig. 111).—Alar expanse 18-23 mm. With characters of arizonensis and reliably distinguished from it only by examination of genitalia. Genitalia (fig. 12) resemble those of both arizonensis and helvialis; but the aedeagus of neohelvialis has, in addition to dense distal patch of slender spines, a small cluster of strong spinelike cornuti from a narrow, straplike sclerotization which distinguishes the species from all others of the group. Frmate.—Alar expanse, 20-23 mm. Similar to male in color and maculation. Genitalia (fig. 66): configuration of ductus bursae from ostium to origin of ductus seminalis as illustrated; sclerotization of ventral margin of ostium interrupted medially; ductus bursae dis- tinctly sclerotized and longitudinally rugose at origin of ductus seminalis. Typr.—Male, U.S. National Museum, USNM 67597, genitalia slide HWC 7729. Typr-LOcALITY.—Plano, Tex. PARATYPES.—UNITED sTATES: Georgia: Milledgeville, 1 9; Tifton, 1 o&. Florida: Quincy, 2 6, 5 9; Cassadaga, 1 9. Louisiana: 1 o, 19. Mississippi: Bolton, 3 9; Starkville, 1 9; Vicksburg,1 @. Mis- souri: Charleston, 4 &, 2 9. Texas: Brownsville, 1 o&, 1 9; Dallas, 1 9; Plano, 1 9. Arizona: Box Canyon, Santa Rita Mts., 2 o; Madera Canyon, Pima Co., 3 9; Madera Canyon, Santa Rita Mts., 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 2 o, 1 9; Santa Rita Mts., 2 9; Baboquivari Mts., Pima Co., 1 0; Redington, 2 #', 1 9. cuBA: Santiago, 15 o, 10 9. gamarca: 1 9. mExico: Vera Cruz: La Gloria Cardel, 1 9; Coatepec, 19. Yucatan: Chichen Itza, 1 &. Jalisco: Guadalajara, 1 9. Sonora: Nainari, 3 &, 8 9 GUATEMALA: Cayuga, 1 co. costa Rica: Avangarez, 19. PANAMA: Corozal, 3 co’. Paratypes in the U.S. National Mu- seum, Los Angeles County Museum, C. P. Kimball collection, and the Canadian National Collection. Foop puant.—Unknown. In FLIGHT.—June to September. Remarks.—Labels on some specimens indicate that neohelmalis has been confused with both mancalis and helvialis. The lack of a subterminal line on the fore- and hindwings of neohelvialis distinguishes it from mancalis. The sparser cilia of the antenna and incrassate midtibia with a hair-pencil distinguishes the males of neohelvialis from those of helvialis. If the specimens are in good condition, the colora- tion along the outer margin of the fore- and hindwings is darker in neohelvialis than in helvialis; in the former, it is concolorous with the markings of the wings and in the latter, concolorous with the ground color of the wings. Worn females of neohelvialis, however, are reliably distinguished from those of helvialis only by examination of the genitalia. Loxostege nayaritensis, new species FicureEs 10, 65, 117 Mate.—Alar expanse 25 mm. Frons conical. Antenna weakly ciliate, cilia one-half or less than one-half as long as width of shaft. Midtibia incrassate, hair-pencil present. Outer spur about one-third as long as inner one. Resembles both pseudohelvialis and arizonensis in color and maculation; in nayaritensis, markings darker and browner with indentation of transverse posterior line of forewing more directly inward at vein 3 and coloration of reniform extending somewhat be- yond lower angle of cell between veins 3 and 4; hindwing with a similar but smaller patch. Genitalia (fig. 10): Like those of arizonensis but with padlike production broader, more rounded, and setae more numerous; sac- culus without a spine midway between outer cluster of spines and base of anellus; vinculum more broadly concave midventrally. Fremate (fig. 117)—Alar expanse 22 mm. Similar to male in color and maculation except that transverse posterior line of forewing is somewhat straighter from costa to vein 3 and small brownish patch between veins 3 and 4 at lower angle of cell stronger; these differences are not likely to be constant in a larger series. Genitalia (fig. 65) with midventral modification of ostium narrow, not interrupted; ductus bursae conspicuously enlarged at origin of ductus seminalis. NO. 3561 LOXOSTEGE—CAPPS 9 Typr.—Male, American Museum of Natural History, genitalia slide HWC 10,644. Typr-LocaLiry.—Compostela, Nayarit, Mexico. ParatyPE.—Type-locality, 19. In collection of Cornell University. Foop puant.—Unknown. In FLicHt.—August. Remarks.—The rather obscure brownish patch at the lower angle of the cell between veins 3 and 4 of the fore- and hindwings distin- guishes nayaritensis from all other species of the genus that lack a subterminal line. Loxostege polingi, new species Fiaures 14, 67, 109 Mats (fig. 109).—Alar expanse 18-23 mm. Frons conical. An- tenna with cilia short, somewhat pubescent. Midtibia moderately incrassate, hair-pencil present. Outer spur less than one-half as long as inner. Maculation similar to that of arizonensis but with shading along outer margin of fore- and hindwings usually not as sharply defined as in arizonensis. Genitalia (fig. 14): Harpe without digitate setae; pad small, nonscobinate but with a few simple setae; clasper broadly rounded and fimbriate distally. Aedeagus with a distal patch of slender cornuti of irregular length; basal keel very small. Frma.e.—Alar expanse 19-23 mm. Similar to male in color and maculation. Genitalia (fig. 67): configuration of ductus bursae from ostium to origin of ductus seminalis as illustrated; ductus bursae not sclerotized and without longitudinal grooves at origin of ductus seminalis; ductus seminalis somewhat scobinate adjacent to origin. Typr.—Male, U.S. National Museum, USNM 67598, genitalia slide HWC 7828. Typxr-LocaLiry.—Lakeland, Fla. PARATYPES.—UNITED STATES: Florida: Cassadaga 1 9; Gainesville, 2 o, 2 2; Lakeland, 3 9; Lake Placid, 1 9; Lake Placid [Archbold Biol. Sta.], 1 o&, 2 9; Warrington, 1 9; Winter Park, 6 9. Texas: Black Jack Springs, 1 9; Kerrville, 1 =; San Diego, 1 o&; San Antonio, 3 9; ‘Texas’ [no additional locality], 19. Arizona: Madera Canyon, Santa Rita Mts., 1 9. mexico: Guerrero: Iguala, 2 9; Puebla: Tehuacan, 2 9; Zapotitlan, 1 9; Oaxaca: Oaxaca, 1 9; Totolapam, 1 9. Paratypes in the U.S. National Museum, American Museum of Natural History, Los Angeles County Museum, the collection of C. P. Kimball, and the Canadian National Collection. Foop pLant.—Unknown. In Fuicur.—April to September in the United States; June to October in Mexico. 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Remarxs.—The type of polingi bears a label indicating that it had been compared with the cotype of citrina and regarded as conspecific with it, prior to dissection of the genitalia. Other labels also indicate confusion with helvialis and with a more remotely related species, Pionea eupalusalis Walker. The rather broad, fimbriate clasper distinguishes the male genitalia of poling: from those of helmalis and eupalusalis; the configuration of the sclerotization of the ductus bursae between the ostium and collarlike structure at origin of the ductus seminalis is diagnostic for the females of polingi. Loxostege pelotasalis, new species Fiaures 15, 113 Mate (fig. 113).—Alar expanse 18 mm. Frons conical. Antenna somewhat pubescent, cilia very short. Midtibia slightly incrassate, hair-pencil weak. Outer spur one-half as long as inner. Similar to polingi in maculation. Genitalia (fig. 15) similar to those of polingi but with uncus broader, stouter; fimbriate clasper of harpe narrower; aedeagus with a narrow, curved, hooklike cornutus in addition to distal patch of slender spinules. FrmMaLe.—Unknown. Typr—Male, Cornell University, genitalia slide HWC 10,642. Typr-LOcALITy.—Pelotas, Rio Gran do Sul, Brazil. Foop pLrant.—Unknown. In FLIGHT.—May. Remarks.—FEfforts to obtain females of pelotasalis were unsuccess- ful. No other closely related species was noted in the South American material examined; there should be no difficulty in association of the sexes when the females become available. Loxostege boliviensis, new species Fiaures 16, 110 Mats (fig. 110).—Alar expanse 19 mm. Frons conical. Antenna pubescent. Similar to pelotasalis in color but with markings much weaker, barely discernible. Genitalia (fig. 16) resemble those of pelotasalis but with uncus more pointed, the fimbriate clasper of the harpe distinctly narrower and more extenuate basally, and with a conspicuous, strong, hooklike production from near middle of dorsal margin of sacculus. Fremate.—Unknown. Typr.—Male, Canadian National Collection, genitalia slide CNC 3765 MS. TyPE-LOCALITY.—Puerto Suarez, Bolivia. Foop pLaAnt.—Unknown. In FriiacHt.—December. NO. 3561 LOXOSTEGE—CAPPS 11 Remarxs.—Although no females of boliviensis are available, no other closely related species was noted in the material examined from Bolivia. There should be no difficulty in associating the sexes when females become available. Loxostege saltensis, new species Figures 69, 114 Femate (fig. 114).—Alar expanse 23 mm. Frons conical. Similar in color and maculation to pelotasalis, but saltensis is larger and termi- nation of posterior transverse line is farther inward on inner margin. Genitalia (fig. 69): ductus bursae with a rather broad, collarlike expansion between constriction near middle and origin of the ductus seminalis. TypE.—Female, U.S. National Museum, USNM 67599, genitalia slide HWC 10,041. TYPE-LOCALITY.—Salta, Argentina. Foop pLtant.—Unknown. Remarks.—The species is known only from the type; configuration of the ductus bursae between the ostium and the origin of the ductus seminalis is diagnostic for the female. There is no date of collection on the label to indicate time of flight. Loxostege fordi, new species Ficures 18, 84, 107 Mate (fig. 107).—Alar expanse 18-20 mm. Frons_ conical. Antenna with cilia short, length one-half or less than one-half as long as width of shaft. Midtibia incrassate, hair-pencil present. Outer spur minute, about one-fifth as long as inner spur. Color and maculation like that of helvialis. Genitalia (fig. 18): Harpe with a small cluster of digitate setae arising from oblique, ridgelike modification near middle of harpe; distal termination of modification broadly rounded, padlike, and scobinate; sacculus with two groups of spines, the outer composed of a rather dense patch of subequal spines, inner of one to three rather stout spines. Aedeagus without cornutus. Fremate.—Alar expanse 17-20 mm. Similar to male in color and maculation. Outer spur distinctly less than one-half as long as inner spur. Genitalia (fig. 84) with sclerotization below ventral margin of ostium crinkled, median production narrow. Typrz.—Male, U.S. National Museum, USNM 67600, genitalia slide HWC 7731. TYPE-LOCALITY.—Tucson, Ariz. PARATYPES.—UNITED STATES: Arizona: Christmas, | 1; Redington, 3 co’; Yuma, 1 o, 19; Kingman, 1 9; Madera Canyon, Santa Rita Mts., 4 3, 59; Madera Canyon, Pima Co., 1 o’, 5 9; Picacho State Park, 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Pinal Co., 107; Santa Rita Mts., PimaCo.,1@. California: Ibanpah Mts., 3”, 2 9; Providence Mts., San Bernardino Co., 10’, 1 9; Furnace Creek, Death Valley, 1 9; Chino Canyon, Palm Springs, 1c’; Borrego 1°. mexico: Sonora: Carbo [Copete Mine, 30 mi. E.],1 &@. Para- types in U.S. National Museum, American Museum of Natural History, Los Angeles County Museum, and the Canadian National Collection. Foop pLtant.—Unknown. In ruicHT.—April to October. Remarxs.—L. fordi resembles helvialis and polingi in color and maculation, but in well-marked specimens of fordi, a faint subterminal line, absent in both helvialis and polingi, is discernible on the forewing. Differences in the lengths of the outer spurs distinguish the males of these species from each other; in helvialis, the outer is one-half as long as the inner; in polingi, it is about one-third as long as the inner; and in fordi, it is much shorter, only about one-fifth as long as the inner one. ‘The females are reliably distinguished only by examination of the genitalia. Loxostege alpinensis, new species Figures 19, 83, 108 Mate (fig. 108).—Alar expanse 20-23 mm. Frons conical. An- tenna weakly ciliate. Outer spur slightly less than one-half as long as inner spur. Color and maculation as in fordi but with markings often more obscure. Genitalia (fig. 19) resemble those of fordi but with outer group of spines smaller and spinules weaker; inner group in a linear arrange- ment, extending from near base of outer group to middle of sacculus. Frema.e.—Alar expanse 21-23 mm. Color and maculation as in male. Genitalia (fig. 83) somewhat similar to those of fordi but with median production of ventral margin of ostium broader, not crinkled laterad ; origin of ductus seminalis more remote from ostium. Typr.—Male, U.S. National Museum, USNM 67601, genitalia slide HWC 7842. TyYPE-LOCALITY.—Paradise, Ariz. PARATYPES.—UNITED STATES: Arizona: 3 o’, 3 9; White Mts., 15 o, 1 9; Huachuca Mts., 1 9. Texas: Brownsville, 1 o, 1 9; San Benito, 3 o&', 4 2; Brewster Co., 1 co; Montague Co., 1 9. Paratypes in the U.S. National Museum and the Canadian National Collection. Foop pLANtT—Unknown. In FLIGHT.—March to September. ReMARKS.—Specimens of this species are likely to be identified in collections as helvialis and Lozostege obliteralis of authors, not Walker. NO. 3561 LOXOSTEGE—CAPPS 13 Loxostege marculenta (Grote and Robinson) FieureEs 20, 70, 130 Botys marculenta Grote and Robinson, 1867, Trans. American Ent. Soc., vol.1, p. 23, pl. 2, fig. 21—Klots, 1942, Bull. American Mus. Nat. Hist., vol. 79, p. 420 [lectotype designation]. Loxostege marculenta (Grote and Robinson).—Forbes, 1920, Cornell Univ. Agric. Exp. Sta. Mem., no. 68, p. 557. Phlyctaenodes obliteralis (Walker).—Hampson, 1899, Proc. Zool. Soc. London p. 280. Lozxostege obliteralis (Walker) of authors.—Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull. 52, p. 382.— Barnes and McDunnough, 1917, Checklist of the Lepidop- tera of boreal America, p. 13.—McDunnough, 1939, Mem. Southern California Acad. Sci., vol. 2, p. 12. Mate (fig. 130).—Alar expanse 18-24 mm. Frons conical. An- tenna weakly ciliate, cilia about one-half as long as width of shaft. Midtibia slightly thicker than hind tibia, hair-pencil weak. Outer spur about one-fifth as long as inner. Head, thorax, and abdomen pale ochreous above, whitish below. Labial palpus upturned, with third segment short, porrect, concealed by scales; palpus twice as long as width of eye; first segment white, second segment luteous above and laterally, white below, all of third segment luteous. Forewing: Upper surface pale yellow, somewhat subhyaline; area along costa and markings luteous; with three transverse lines, trans- verse anterior and posterior lines narrow, somewhat denticulate, subterminal line smooth, somewhat diffuse and broader than anterior and posterior lines. Anterior transverse line bent obliquely out- ward from costa to vein 1b, thence zig-zag inward and outward to hindwing margin, posterior transverse line bent slightly inward from origin on costa to vein 7, weakly crenulate and broadly concave outwardly to slightly below vein 3, almost directly inward to vein 2, slightly concave outwardly to vein 1b, thence directly posterior to hindwing margin. Subterminal line nearly straight, about midway between outer margin of wing and loop of transverse posterior line at end of cell, subparallel to outer wing margin. Hindwing: Upper surface coloration similar to that of forewing but paler, with two rather diffuse transverse lines appearing as con- tinuations of transverse posterior and subterminal lines of forewing. Postmedial line occasionally weakly crenulate, extending from costa to slightly below vein 2. The subterminal line smooth, parallel to outer margin and distinctly closer to outer wing margin than to postmedial line. Fringe of both fore- and hindwing concolorous with ground color of wing. Under surface of fore- and hindwing whitish with ochreous tinge; markings stronger on forewing but weaker on both wings than above. 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Genitalia (fig. 20): Uncus stout. Harpe with cluster of digitate setae; pad broadly rounded and scobinate distally; sacculus with a long, slender, curved spine and two subequal strongly sclerotized productions, the outer bluntly conical, the inner slender, fingerlike, sometimes sharply pointed distally. Aedeagus stout, without cornutus. Frema.e.—Alar expanse 21-26 mm. Similar to male in color and maculation. Genitalia (fig. 70) with ventral margin of ostium distinctly concave medially. Lectotype.—Female, American Museum of Natural History, Grote and Robinson no. 22995. TyprE LocaLity.— Atlantic District (Penna.).” Foop pLtants.—Ambrosia trifida and Solidago. DISTRIBUTION.—UNITED sTATES: Maine, Connecticut, Massachu- setts, New York, New Jersey, Pennsylvania, Maryland, District of Columbia, Virginia, North Carolina, Alabama, Mississippi, Ohio, Illinois, Missouri, Michigan, Iowa, Texas, and New Mexico. CANADA: Ontario, Quebec. SPECIMENS EXAMINED.—117. In ruicgHt.—May to September. RemARKs.—Labels in collections indicate confusion of marculenta with helvialis, obliteralis, and mancalis. The presence of subterminal lines on the fore- and hindwings distinguishes marculenta from helvialis and others of that group. The markings on the wings of marculenta are somewhat diffuse, often obscure, usually more brownish and with the definition weaker than in mancalis, and the subterminal line of the hindwing is closer to the outer margin in mancalis than in marculenta. ‘The frons is conical in marculenta and round in obliteralis. Loxostege neomarculenta, new species Figures 21, 71, 131 Matr.—Alar expanse 22-24 mm. With characters of marculenta and reliably distinguished from it only by examination of the genitalia. Genitalia (fig. 21): harpe with digitate setae; scobinate area of pad rather narrow; sacculus with a strong subequal bifid spine and a stout, conical production slightly beyond base of spine. Aedeagus without cornutus. Femaue (fig. 131).—Alar expanse 22-23 mm. Similar to male in color and maculation. Genitalia (fig. 71) with two conspicuous, pronglike elements from median area of ventral margin of ostium, margins of prongs occasionally somewhat serrate; ductus bursae distinctly angulate at constriction above origin of ductus seminalis. Typr.—Male, U.S. National Museum, USNM 67602, genitalia slide HWC 7825. NO. 3561 LOXOSTEGE—CAPPS iS TyYpE-LocALIty.— Decatur, IIl. ParatyPEes.—Type-locality, 1 @. Maryland: Plummers Island, 1 o,19. Virginia: Skyland, 19. ‘West Virginia” [no additional locality], 1 9. Foop pLant.—Unknown. In FLIGHT.— May to July. Remarks.—The male genitalia of marculenta and neomarculenta differ chiefly in the armature of the sacculus; the bifurcate spine, shorter outer conical production, and narrower pad distinguish the males of neomarculenta from those of marculenta. The presence of two median prongs arising from the ventral margin of the ostium distinguishes neomarculenta females from those of marculenta. Loxostege pseudobliteralis, new species FIGURES 25, 88, 135 Mate (fig. 135).—Alar expanse 21-24 mm. Frons conical, pro- duction moderate. Antenna weakly ciliate. Midtibia incrassate, hair-pencil distinct. Outer spur minute, about one-sixth as long as inner. Color and maculation as in marculenta and neomarculenta but forewing usually with an indentation of subterminal line evident at vein 2. Genitalia (fig. 25): Harpe with a cluster of digitate setae; sacculus with spine short, somewhat modified, with one or two spinules basad; subconical production at base of spine bearing several slender setae. Aedeagus attenuate distally and with a small patch of coarse, sub- conical granules. FremMaLe.—Alar expanse 22-26 mm. Color and maculation like that of male. Genitalia (fig. 88): ductus bursae rather strongly sclerotized at ostium dorsally and ventrally; ventral margin of ostium strongly convex, somewhat crinkled. Larva.—Mature, 24 mm long. Head sordid white, with reticu- late pale amber markings; a conspicuous small fuscous patch at lateral incision of hind margin. Prothoracic shield sordid white or with a yellowish tinge, narrowly margined with brown or brownish fuscous, the pigmentation slightly interrupted middorsally. Body pinacula of similar brownish pigmentation. Abdominal segments 1-8: Pinacula of seta I and seta II transversely elongate, several times longer than wide; pinacula of seta III subquadrangular longitudinally; pinacula below level of spiracle oval shaped, with pigmentation weaker and less extensive than those dorsad. Ninth abdominal segment: posterior half of pinaculum of paired setae II pigmented; pinaculum bearing setae I-III pigmented; pinaculum of seta VI pigmented. Anal shield with a small, pigmented patch at base of anterior lateral seta. 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Typr.—Male, U.S. National Museum, USNM 67605, genitalia slide OBP 149. Typxr-LocaLiry.—Paradise, Ariz. PARATYPES.—UNITED STATES: Arizona: type-locality, 3 9; No- gales, 19; Baboquivari Mts., Pima Co., 1 o', 3 9; Huachuca Mts., 5 o', 1 9; Santa Rita Mts., Madera Canyon, 10 o’, 6 9; Santa Rita Mts., 1 9; Madera Canyon, Pima Co., 1 9; White Mts., 2 9; “‘Arizona”’ [no additional locality], 1 &@. Texas: Kerrville, 1 &, 1 9; San An- tonio, 1 9; Brewster Co., 2 9. mexico: Morelos: Cuernavaca, 1 9. Paratypes in the U.S. National Museum and the Canadian National Collection. Foop pLAnr.—Morning glory. In FuicHT.—July to September in the United States, May in Mexico. REeMARKS.—Differences in character of the spine and subconical production of the sacculus distinguish the males of pseudobliteralis from those of marculenta and neomarculenta. The shape and character of sclerotization of the ductus bursae at the ostium is diagnostic for the females of pseudobliteralis. Loxostege neobliteralis, new species FicureEs 26, 87, 132 Mate (fig. 132).—Alar expanse 18-24 mm. Antenna, frons, tibial character, color, and maculation essentially the same as in marculenta and neomarculenta but reliably distinguished from them by ex- amination of the genitalia. Genitalia (fig. 26): harpe with digitate setae; a short, sharp, spinelike projection from near middle of oblique modification bearing cluster of digitate setae; pad moderately de- veloped, scobination extensive, rounded or somewhat truncate distally; clasper conspicuously expanded distally and strongly denticulate. Aedeagus without cornutus. Frma.e.—Alar expanse 22-23 mm. Similar to male in color and maculation. Genitalia (fig. 87): ostium subcircular; margin rather strongly sclerotized; ventral margin straight or nearly so. Larva.—Mature, 25 mm long. Similar to larva of pseudobliteralis but with marginal fuscous pigmentation of prothoracic shield inter- rupted on lateral margin; pinacula nonpigmented below level of spiracles on abdominal segments. Ninth abdominal segment with pigmentation of pinaculum bearing paired setae JI interrupted on dorsum; pinaculum of setae I-III and that of seta VI nonpigmented, concolorous with adjacent body area. Anterior lateral seta of anal shield without a fuscous patch at its base. Typr.—Male, U.S. National Museum, USNM 67604, genitalia slide HWC 10,609. NO. 3561 LOXOSTEGE—CAPPS 7s Typr-LocaLiry.—Hubberton, Vt. PARATYPES (16).-—-UNITED STATES: type-locality, 2 #, 1 9; Penn- sylvania: Pittsburgh, 1 o; Oak Station, 2 @. New Jersey: Lake- hurst, 1 9. Maryland: Plummers Island, 1 o&, 2 9. District of Columbia: Washington, 1 o&. West Virginia: 1 9. Mississippi: “Aor. College” [sic] 1 9. Indiana: Lafayette, 1 @. Iowa: Soldier [near] 1 co. CANADA: Quebec: Hummingford, 1 9. Paratypes in the U.S. National Museum and the Canadain National Collection. Foop PpLanr.—Ipomoea. In FuicgHT.—May to September. Remarxs.—The short, spinelike projection from near the middle of the lower margin of the harpe, in combination with the denticulate character of the clasper, is diagnostic for males of neobliteralis. The female genitalia resemble those of pseudobliteralis, but the straight ventral margin of the ostium distinguishes females of neobliteralis from those of pseudobliteralis. Loxostege potosiensis, new species FIGURE 22 Maur.—Alar expanse 22 mm. Frons conical. Antenna ciliate, leneth of cilia slightly less than width of shaft. Midtibia incrassate; hair-pencil present. Outer spur short, about one-fifth as long as inner spur. Color and maculation as in neomarculenta. Genitalia (fig. 22): Harpe with spine from sacculus simple, rather long and slender; distal margin of pad medially angulate, the lower element with fine spinules, slightly bifurcate distally. Aedeagus somewhat attenuate distally, with a patch of short, stout spinules. Fremate.—Unknown. Typr.—Male, U.S. National Museum, USNM 67603, genitalia slide HWC 12,018. Typre-LocaLity.—Tamazunchale, San Luis Potosi, Mexico. Foop pirant.—Unknown. In FiuigHt.—May to August. Remarks.—L. potosiensis is reliably distinguished from pseud- obliteralis and several other species only by examination of the genitalia. The medially angulate distal margin of the pad, in combination with the fine spinulation and distal subequal bifurcation of the ventral element, distinguishes potosiensis males from all others of the group. Loxostege cayugalis, new species FiaureEs 35, 85, 136 Matue.—Alar expanse 21-25 mm. Fronsconical. Antenna ciliate, cilia about as long as width of shaft. Midtibia incrassate; hair-pencil weak. Outer spur one-fifth aslong as inner spur. Similar to pseud- obliteralis in color and maculation. 221-525—67—_2 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Genitalia (fig. 35): Harpe with distal termination of oblique ridge- like modification pointed, curved, hooklike, and nonscobinate; a similar hook at base of cluster of digitate setae; sacculus with two conspicuous spines, one to several small, short spines often associated with outer. Aedeagus with distal patch of subconical granules. Fremate (fig. 136).—Alar expanse 26-28 mm. Color and macula- tion as in male. Genitalia (fig. 85): shape of ostium and character of its ventral margin is diagnostic for females; ostium somewhat variable, nearly round to distinctly oval, ventral margin smooth. Typr.—Male, U.S. National Museum, USNM 67606, genitalia slide HWC 7912. TyYprE-LOCALITY.—Cayuga, Guatemala. PARATYPES.—GUATEMALA: type-locality, 2 o; Quirigua, 1 o’; San Geronimo, 1 9. cosTa RICA: Tuis, 1 o, 1 9; Cachi, 2 9. MEXICO: San Luis Potosi, 1 9; Tamazunchale, 1 9; Morelos: Cuernavaca, 1 9; Chiapas: Soconusco (Finca la Violeta), 38 o, 15 9; Veracruz: Cordoba, 1 9. Paratypes in U.S. National Museum, British Museum (Nat. Hist.), and the Canadian National Collection. Foop pLant.—Unknown. In FuigHT.—April to October. Remarks.—L. cayugalis is likely to be confused in collections with the species described as Pachyzancla cynoalis Druce, which is treated next herein. One of the cayugalis paratypes from Cachi, and those from San Geronimo and Cuernavaca, are from the syntypes of cynoalis. Loxostege cynoalis (Druce), new combination FicureEs 36, 72, 137 Pachyzancla cynoalis Druce, 1895, in Godman and Salvin, Biologia Centrali- Americana, Zoology, Lepidoptera, Heterocera, vol. 2, p. 221 [in part]. Mate (fig. 137).—Alar expanse 23 mm. Frons conical. Antenna ciliate, cilia about as long as width of shaft. Upper spur about one-fourth as long as inner spur. Color and maculation similar to that of cayugalis. Genitalia (fig. 36): Harpe with production from base of digitate setae strongly denticulate; dorsal margin of sacculus with two con- spicuous, stout spines. Anellus stout. Aedeagus with distal patch of small conical granules. Fremate.—Alar expanse 20-21 mm. Similar to male in color and maculation. Genitalia (fig. 72) resemble those of neomarculenta but with contour of ductus bursae undulate between ostium and origin of ductus seminalis. Lecroryrr.—Male, British Museum (Nat. Hist.), genitalia slide HWC 17,454, present designation. Typr-LocaLiry.—Volcan de Chiriqui, Panama. NO. 3561 LOXOSTEGE—CAPPS 19 Foop pLant.—Unknown. DistriputTion.—Guatemala and Panama. SPECIMENS EXAMINED.—3. IN FLIGHT.—May. Remarxs.—Differences in genitalia and length of the upper spurs distinguish the males of cynoalis and cayugalis, but examination of the genitalia is necessary for reliable identification of the females of these two species. Druce did not indicate the number of specimens in the type series of cynoalis but stated that he had specimens from Mexico (Cuer- navaca, Morelos), Guatemala (San Geronimo), Costa Rica (Cachi), and Panama (Chiriqui, Volcan de Chiriqui). Of the five syntypes in the British Museum, the specimen from Volcan de Chiriqui (2000— 3000 ft, Champion), a male, was labelled type by Druce and is the one selected herein as lectotype of cynoalis. The male and female from Cuernavaca and the female from Cachi are not conspecific with the type and represent a new species treated herein. The abdomen is missing from the female from Geronimo; the placement of that specimen is uncertain. Loxostege ecuadoralis, new species FicureEs 37, 138 Mats (fig. 138).—Alar expanse 24-26 mm. Frons weakly conical. Antenna ciliate, cilia about one-half as long as width of shaft. Mid- tibia somewhat incrassate; hair-pencil weak. Outer spur one-fifth as long as inner spur. Similar to cayugalis in color and maculation. Genitalia (fig. 37) resemble those of cayugalis but with dorsal element of extenuation of oblique ridgelike modification of harpe somewhat truncate and with a few slender setae; hook at base of cluster of digitate setae stronger and more pointed than in cayugalis. Frmaute.—Alar expanse 26 mm. Similar to male in color and maculation. Genitalia resemble those of both cayugalis and the next species; distinguished from cayugalis by the much wider nonpigmented, membranous median area dorsad of ostium and the somewhat nar- rower and less constricted ductus bursae between ostium and origin of ductus seminalis. Distinguished from the next species by the smooth, concave, ventral margin of ostium and the narrower and much shorter ductus bursae between ostium and origin of ductus seminalis. Typr.—Male, U.S. National Museum, USNM 67607, genitalia slide HWC 7848. TyPpr-LocaLity.—“Environs de Loja Equateur.” PARATYPES.—ECUADOR: type-locality, 1 o&. PERU: Huanuco, 1 o. BOLIVIA: Bajo Palmar, Chapare Cochamba, 1 9. Paratypes in U.S. National Museum and the Canadian National Collection. 20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Foop pLant.—Unknown. In ruigHT.—November to January. ReMARKS.—Labels on some specimens of ecuadoralis indicate they have been confused in collections with cynoalis and obliteralis. Loxostege neotropicalis, new species Ficuress 24, 86, 133 Mate (fig. 133).—Alar expanse 22-25 mm. Antenna ciliate, cilia about one-half as long as width of shaft. Muidtibia incrassate; hair- pencil present. Outer spur short, one-fifth as long as inner spur. Similar to cayugalis in color and maculation but distinguished from it by genitalic differences. Genitalia (fig. 24): Harpe with distal termination of oblique ridge- like modification padlike, spinulation coarse; width of pad and extent of spinulation variable; sacculus with a single, stout spine arising from dorsal margin. Aedeagus robust, with a distal patch of subconical granules. Frma.e.—Alar expanse 25-28 mm. Similar to male in color and maculation. Genitalia (fig. 86) somewhat like those of cayugalis but ostium of different shape and ventral margin of ostium crinkled. Typr.—Male, U.S. National Museum, USNM 67608, genitalia slide 7871. Typr LOcALITY.—Jalapa, Mexico. PaRaTYPES.—MEXICO: Puebla: Zapotitlan de las Salinas, 3 o&; Guanajuato: Irapuato, 1 9. GuaTeMALA: Volcan Santa Maria, 1 9. COSTA RICA: Juan Vinas, 1 9. VENEZUELA: Rancho Grande [Near Maracay], 1 1,19. Paratypes in U.S. National Museum, American Museum of Natural History, and the Canadian National Collection. Foop pLrant.—Unknown. In riicut.—May to October. Loxostege jaralis (Schaus), new combination Fiaur&s 33, 78, 139 Phlyctaenodes jaralis Schaus, 1920, Proc. Ent. Soc. Washington, vol. 22, p. 220. M atye.—Alar expanse 23mm. Fronsconical. Antenna pubescent. Midtibia incrassate, hair-pencil well developed. Outer spur one-third as long as inner. Similar to pseudobliteralis in color and maculation, but with ground color and markings slightly darker and subterminal line of hindwing closer to outer margin. Genitalia (fig. 33): Sacculus with one to three short, stout spines in outer group; inner group of two to four similar but somewhat larger spines; a larger spine about midway between the two groups. Pad narrow and rather densely spinulate. NO. 3561 LOXOSTEGE—CAPPS yA FrEmMate (fig. 139).—Alar expanse 23-25 mm. Similar to male in color and maculation. Genitalia (fig. 78): configuration of genital plate diagnostic. Typr.—Female, U.S. National Museum, USNM 23829, genitalia slide HWC 6008. TYpE-LOcALITY.—Guadalajara, Jalisco, Mexico. Foop pLtAnt.—Unknown. DISTRIBUTION.—MEXIco: Oaxaca. Puebla: Zapotitlan de las Salinas. San Luis Potosi: Tamazunchale. Jalisco: Guadalajara. UNITED STATES: Arizona: Redington; Baboquivari Mts., Pima Co.; Madera Canyon, Santa Rita Mts. [The U:S. records are new for the species. ] SPECIMENS EXAMINED.—11. In riicHt.—May to September. Remarxks.—The Arizona specimens were confused with helvialis in the U.S. National Museum collection, but the presence of a subter- minal line on the forewing of jaralis distinguishes it from helvialis. The male heretofore has not been associated with females of the species. Loxostege sacculalis Amsel Figures 31, 118 Loxostege sacculalis Amsel, 1956 (1954), Bol. Ent. Venezuela, vol. 10, p. 250. Mate (fig. 118) —Alar expanse 19-22 mm. Frons conical. An- tenna ciliate, cilia slightly shorter than width of shaft. Midtibia incrassate, hair-pencil weak. Outer spur about one-third as long as inner spur. Similar to jaralis in color and maculation. Genitalia (fig. 31): Essentially like those of jaralis but with differ- ences in character of outer termination of dorsal margin of sacculus and contour of ventral margin of oblique ridge of harpe. Typr.—Male, Amsel collection. TYPE-LOcALITY.—Maracay, Venezuela. Foop pLtant.—Unknown. Distrisution.—Known only from type-locality. SPECIMENS EXAMINED.—1. In FLIGHT.—August. Remarks.—No females of sacculalis were available for study; the description of the female genitalia was omitted in Amsel’s treatment of the species. The differences in the male genitalia of jaralis and sacculalis may not be constant; a larger series of males and com- parison of the female genitalia of both species is needed to determine the status of the two names. 22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Loxostege mancalis (Lederer) FIGuRES 27, 89, 119 Botys mancalis Lederer, 1863, Weiner Ent. Monatschr., vol. 7, p. 464, table 9, fig. 4.—Munroe, 1958, Canadian Ent., vol. 90, no. 9, p. 511. Phlyctaenodes mancalis (Lederer).—Hampson, 1899, Proc. Zool. Soc. London, . 208. Paeiee mancalis (Lederer).—Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull. 52, p. 382.—Forbes, 1920, Cornell Agric. Exp. Sta. Mem., no. 68, p. 557.— Barnes and McDunnough, 1917, Checklist of the Lepidoptera of boreal America, p. 131—McDunnough, 1939, Mem. Southern California Acad. Sci. vol. 2, p. 12. Mate (fig. 119).—Alar expanse 20-24 mm. Frons conical. An- tenna ciliate, cilia about one-half as long as width of shaft. Midtibia incrassate, hair-pencil well developed. Outer spur one-third as long as inner. Similar to marculenta in color and maculation, but speci- mens of mancalis are somewhat duller and less ochreous, with defi- nition of markings sharper, subterminal band of forewing usually broader anteriorly, and subterminal line of hindwing closer to outer margin (pale ochreous area between line and outer margin of wing narrower than subterminal line). Genitalia (fig. 27): Harpe without digitate setae; with two con- spicuous subequal hooklike structures; the small one downcurved, arising from near middle of harpe and extending below ventral margin; the large one from sacculus, rather broad basally, distal termination a sharp point; pad moderately broad, rounded, scobinate. Aedeagus somewhat attenuate distally, bluntly pointed and with a patch of small spinules. Frmate.—Alar expanse 22-24 mm. Similar to male in color and maculation. Genitalia (fig. 89): ductus bursae rather strongly sclerotized from ostium to junction with ductus seminalis and some- what incrassate near middle; ostium with lateral production short, spurlike. Larva.—Mature, 28-30 mm long. Head pale amber, reticulation weak. Body color sordid white. Pigmentation of markings on prothoracic shield and body pinacula brown or blackish. Prothoracic shield with pigmentation restricted to lateral margin. Pinacula of seta Ia-Ib and IJa-IIb of meso- and metathorax moderately convex, irregularly ovoid or round in shape, pigmentation strong. On ab- dominal segements 1-8: Pinacula of seta I and seta II conspicuously large, round and flat; pinaculum of seta III reniform and smaller than those of setae I and II; pigmentation of setae I, II, and III strong; pinacula of setae below level of spiracles not pigmented or the pig- mentation obsolescent centrally. Ninth abdominal segment with pinacula not pigmented. No. 3661 LOXOSTEGE—CAPPS 23 Lrectrotyre.—Male, Naturhistorische Museum, Vienna, genitalia slide HWC 17,238. TypxE-Locatity.—North America. PaRALECTOTYPE.—Female, British Museum (Nat. Hist.), genitalia slide HWC 17,327. Foop puLants.—Amaranthus retroflerus, mint, morning glory, tobacco, and Rumez. DIsTRIBUTION.—UNITED STATES: Maryland, District of Columbia, Virginia, North Carolina, South Carolina, Tennessee, Florida, Louisiana, Mississippi, Texas, Oklahoma, Nebraska, Missouri, Illinois, Arizona. MExiIco: Sonora, Jalisco, Morelos, Puebla, Oaxaca. COSTA RICA: Avangarez. SPECIMENS EXAMINED.—169. In FiuigHt.—March to September. Remarks.—In describing mancalis, Lederer indicated that the type series consisted of nine specimens, which were from North America and Brazil “Ex Mus. Caesar, Felder.’”” Only two of these could be located, a male, by Dr. Kasy of the Vienna Museum, and the other, a female, by Mr. Whalley, at the British Museum. I hereby designate the male as lectotype and the female as paralectotype of the species. It is not likely that the syntypes from Brazil are the same species as those from North America, for I have found no specimens of mancalis in the considerable amount of material from Brazil and other South American countries that J have examined. The lack of digitate setae, in combination with the character of the spine arising from near the middle of the ventral margin of the harpe and the hook arising from the sacculus, is diagnostic for the male genitalia of mancalis. The rather smooth, straight, ventral margin and short lateral, spurlike production of the ostium, in com- bination with the incrassate character of the ductus bursae between the ostium and ductus seminalis, distinguish the females of mancalis from all others of the genus. Loxostege ramsdenalis (Schaus), new combination FIGURES 28, 74, 120 Phlyctaenodes ramsdenalis Schaus, 1920, Proc. Ent. Soc. Washington, vol. 22, p. 219. Maue.—Alar expanse 20-23 mm. Frons conical. Midtibia in- crassate, hair-pencil distinct. Similar to mancalis in color and macula- tion but with indentation of transverse posterior line deeper and more acute between veins 1b and 2 of forewing. Genitalia (fig. 28): Somewhat similar to those of mancalis but differing as follows: harpe with a cluster of digitate setae; sclerotized production from near middle of ventral margin bluntly rounded, 24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 spatulate; distal pad narrow, spinules rather stout; sacculus with hook more oblique inwardly and a small spine basad. Frma.e (fig. 120).—Alar expanse 17-20 mm. Color and maculation asin male. Genitalia (fig. 74): ostium with ventral median emargina- tion deep and broadly concave. Typr.—Female, U.S. National Museum, USNM 23827, genitalia slide HWC 6012. TYPE-LOCALITY.—Santiago, Cuba. Foop pLant.— Unknown. DisTRIBUTION.—cUBA: Santiago. CAYMAN ISLANDS: Grand Cayman. mexico: Jalapa and Oaxaca. GuaTEeMALA: Volcan Santa Maria. VENEZUELA: Rancho Grande [near Maracay]. BRAZIL: Pernambuco. BOLIVIA: Corioco. SPECIMENS EXAMINED.—27. In FitigHt.—April to June. Remarks.—As there were no examples of mancalis in the South American material studied, and as ramsdenalis is so similar to mancalis in size, color, and maculation, it seems likely that at least part of the Brazilian syntypes apparently misidentified as mancalis (q.v.) are in fact ramsdenalis. In collections, ramsdenalis has also been confused with Phlyctaenodes conisphora Hampson. Loxostege pergilvalis (Hulst) Ficures 30, 79, 122 Botis pergilvalis Hulst, 1886, Trans. American Ent. Soc., vol. 13, p. 151.—Klots, 1942, Bull. American Mus. Nat. Hist., vol. 79, art. 6, p. 421 [lectotype designation]. Botys pergilvalis Hulst——Hampson, 1899, Proc. Zool. Soc. London, p. 208. [A synonym of Phlyctaenodes coloradensis (Grote and Robinson) in error.] Lozostege pergilvalis (Hulst).—Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull. 52, p. 382.—Barnes and McDunnough, 1916, Contr. Nat. Hist. Lepid., vol. 3, no. 3, p. 191 [type restricted to female of the Henry Edwards collection].— Barnes and McDunnough, 1917, Checklist of the Lepidoptera of boreal America, p. 131—McDunnough, 1939, Mem. Southern California Acad. Sci., vol. 2,p. 12. [As synonym of coloradensis (Grote and Robinson) in error.] Mate (fig. 122).—Alar expanse 20-26 mm. Frons conical. Antenna weakly ciliate. Midtibia weakly incrassate, hair-pencil absent or obsolescent. Outer spur slightly less than one-half as long as inner. Resembles mancalis in color and maculation but differs as follows: Transverse lines narrower and weaker; subterminal line of forewing little, if any, broader anteriorly; hindwing with postmedial line concave outwardly, postmedial and subterminal lines often obsolescent, interrupted, dotlike. Genitalia (fig. 30): Harpe with cluster of digitate setae; pad rounded distally, the spinulation fine, dense; sacculus with two rather large conspicuous spines and a group of small spines adjacent to NO. 3561 LOXOSTEGE—CAPPS 25 base of the large outer spine; dorsal margin of sacculus with median subtriangular production. Frema.tre.—Alar expanse 18-28 mm. Color and maculation as in male. Genitalia (fig. 79): rugose character of postgenital plate diagnostic. LecrotyPpe.—Female, American Museum of Natural History, genitalia slide FHR 4634. Typr-LocaLity.—Arizona. Foop pLant.—Corn? DIsTRIBUTION.—UNITED STATES: Arizona, New Mexico, Texas, Towa, Illinois, and Delaware. caNapa: Ontario. mexico: Federal District, Puebla, and Jalisco. SPECIMENS EXAMINED.—220. In FLigHT.—May to September. Remarks.—Hulst (1886) stated that the type series of pergilvalis consisted of three females from Arizona. Hampson (1899), followed by authors of subsequent check lists of American Lepidoptera, erred in his treatment of pergilvalis as a synonym of coloradensis. The male specimen from Arizona in the Neumoegen collection that bears a pergivalis type label of Hulst (to which reference was made by Barnes and McDunnough (1916)), agrees with coloradensis in color, maculation, and genitalia and is definitely conspecific with colora- densis; however, the female type designated by (Barnes and McDunnough, 1916), and also subsequently designated lectotype by Klots (1942), is distinct from coloradensis and is a valid species. L. pergilvalis and coloradensis differ in both maculation and genitalia. Postmedial and subterminal lines are present on the hindwing of pergilvalis and are absent in coloradensis (fig. 123). The harpe of pergilvalis has a cluster of digitate setae; that of coloradensis (fig. 17) is without digitate setae. Differences in the character of the sclero- tization adjacent to the ostium readily distinguish the female genitalia of pergilvalis from those of coloradensis (fig. 77). Of the rather large series at hand, only one specimen had a plant association; it is labelled ‘‘on corn” and is from Coapa, D. F., Mexico. Whether it was collected as an adult on corn or reared from a larva feeding on corn is not definitely ascertainable. Loxostege coloradensis (Grote and Robinson) Fieures 17, 77, 123 Botis coloradensis Grote and Robinson, 1867, Trans. American Ent. Soc., vol. I ps2os Botys coloradensis Grote and Robinson.—Klots, 1942, Bull. American Mus. Nat. Hist., vol. 79, art. 6, p. 419. Phlyctaenodes coloradensis (Grote and Robinson).—Hampson, 1899, Proc. Zool. Soc. London, p. 208. 26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Lozxostege coloradensis (Grote and Robinson).—Fernald, 1903, 7m Dyar, U.S. Nat. Mus. Bull. 52, p. 382.—Barnes and McDunnough, 1916, Contr. Nat. Hist. Lepid., vol. 3, no. 3, p. 191; 1917, Checklist of the Lepidoptera of boreal America, p. 131—McDunnough, 1939, Mem. Southern California Acad. Sci., vol. 2, p. 12. Maue.—Alar expanse 23-31 mm. Frons conical. Antenna weakly ciliate. Midtibia incrassate, hair-pencil present, buff. Outer spur minute, about one-sixth as long as inner. Color whitish, with a slight ochreous tinge. Markings of forewing similar to those of pergilvalis but somewhat more ochreous and weaker. Hindwing whitish, sub- hyaline, without postmedial and subterminal lines, or the postmedial barely discernible. Genitalia (fig. 17): Harpe without digitate setae; sacculus with two conspicuous subequal spines and an outer cluster of small, subequal spines. Aedeagus with distal patch of short, sharp, spinules. Fema.e (fig. 123).—Alar expanse 27-31 mm. Color and macula- tion asin male. Genitalia (fig. 77): ventral margin of ostium broadly concave, shallow; sclerotization laterad of ostium concave, granulose. Spinulation of bursa copulatrix conspicuous from origin of accessory pouch to junction with ductus bursae. Larva.— Penultimate stage, 18 mm. Head pale amber, reticula- tion weak; a small fuscous patch at lateral incision on hind margin. Body sordid white. Pigmentation of markings of prothoracic shield and body pinacula brownish. Thorax: anterior and lateral margin of prothoracic shield with a brownish patch at bases of setae Ia, Ib, Ic, and IIc; pigmentation discontinuous on dorsum and between setae Ia and Ib; posterior margin concolorous with adjacent body area, without dark markings; prespiracular shield and pinaculum of group VI conspicuously pigmented. Meso- and metathorax with pinacula of setae Ia-Ib, Ila-IIb, IV, V, and VI rather large, pigmentation strong; pinaculum of seta III smaller, pigmentation weaker. Abdominal segments: pinacula of setae I and II of first and eighth segments more strongly pigmented than on second to seventh segments; pinacula of setae III darkly pigmented and of approximately equal size on segments 1-8. Pinacula below level of spiracles not pigmented on segments 1-8. Pinacula of ninth segment not pigmented. Anal shield without conspicuous markings. Luctotypr.—Female, American Museum of Natural History, slide no. 22998 Grote and Robinson, present designation. TypsE-LocaLity.— Colorado. Foop pLant.— Wild sunflower. Distripution.—Arizona, Utah, Colorado, Iowa, Kansas, Missouri, Oklahoma, and Texas. SPECIMENS EXAMINED.— 51. NO. 3561 LOXOSTEGE—CAPPS Di In FuicHt.— April to September. Remarks.—See the discussion of pergilvalis (p. 24). Loxostege cochisensis, new species FIguREs 39, 82, 121 Maur.— Alar expanse 21-26 mm. Frons conical. Antenna weakly ciliate. Midtibia slightly more incrassate than hind tibia; hair- pencil obsolescent or absent. Outer spur one-half as long as inner. Resembles pergilvalis but with ground color usually more ochreous and transverse lines of forewing slightly smoother: postmedial line of hindwing absent, or if evident, smooth, somewhat diffuse, termen concolorous with fringe. Genitalia (fig. 39): Uncus somewhat narrower apically. Harpe with digitate setae; outer half of dorsal margin of sacculus somewhat produced, with numerous, short inwardly directed spines. Aedeagus without cornuti. Fremate (fig. 121) —Alar expanse 19-26 mm. Similar to male in color and maculation. Genitalia (fig. 82): postgenital plate strongly sclerotized, concave, with distinct transverse ridges. Typr.—Male, U.S. National Museum, USNM 67609, genitalia slide OBP 147. Typr-LocaLity.—Palmerlee, Ariz. Paratypes.—Arizona: type-locality, 1 9; [no additional locality], 2 9; Santa Rita Mts. 1 o, 1 9; Southwestern Res. Sta., Chiricahua Mts., Cochise Co., 1 o&, 2 9; Madera Canyon, Santa Rita Mts., 2 0, 3 9; Madera Canyon, Santa Rita Mts., Pima Co., 4 9; Huachuca Mts., 1 9. Texas: The Basin, Chisos Mts., Brewster Co., 2 o, 1 9. Paratypes in the U.S. National Museum, Los Angeles County Mu- seum, and the collection of Mr. C. P. Kimball. Foop pLant.— Unknown. In Fuicut.—June to September. Remarks.—The narrower uncus, in combination with the character of the spinulation on the dorsal margin of the sacculus, is diagnostic for the male genitalia of cochisensis. The character of the postgenital plate (concave with transverse ridges) distinguishes the female geni- talia of cochisensis from all others of the genus. Loxostege jacalensis, new species Fiaures 38, 134 Ma se (fig. 134).—Alar expanse 21 mm. Frons conical. Antenna weakly ciliate, cilia about one-half as long as width of shaft. Midtibia weakly incrassate; hair-pencil absent or obsolescent. Outer spur slightly less than one-half as long as inner. Similar to cochisensis in 28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 color and markings. Genitalia (fig. 38) similar to those of cochisensis but with uncus stouter, fewer spines on dorsal margin of sacculus, spines coarser, anellus broader basally, outer margins of lateral arms parallel or nearly so. Fremaute.— Unknown. Typr.—Male, Canadian National Collection, genitalia slide HWC 17,616. TypxE-LocaLiry.—Jacala, Hidalgo, Mexico. Foop pLaAnt.— Unknown. Remarks.—The species is known only from the type specimen. Loxostege yucatanalis, new species FIGURE 48 Maur.—Alar expanse 20 mm. Frons conical. Antenna weakly ciliate, cilia about one-half as long as width of shaft. Outer spur one-fourth as long as inner. Similar to jacalensis in color and macu- lation, but somewhat paler. Genitalia (fig. 48) resemble those of pseudobliteralis but harpe without digitate setae, pad broader with spinulation finer, aedeagus more robust, cornutus stronger. FremaLe.— Unknown. Typr.—Male, Canadian National Collection, genitalia slide HWC 17,620. Typr-LocaLity.—Chichen Itza, Yucatan, Mexico. Foop pLant.— Unknown. Remarks.—The species is known only from the type-specimen. Loxostege autocratoralis Dyar Fiaures 41, 93, 115 Lozxostege autocratoralis Dyar, 1912, Pomona College Journ. Ent., vol. 4, no. 2, . 747. a eras autocratoralis (Dyar).—Dyar, 1914, Proc. U.S. Nat. Mus., vol. 47, p. 281. Maue.—Alar expanse 20-24 mm. Frons conical. Midtibia incrassate, hair-pencil distinct. Outer spur one-third as long as inner. Resembles nayaritensis in color and maculation but somewhat paler, subterminal line present on forewing. Genitalia (fig. 41): Harpe without digitate setae; dorsal margin of sacculus straight or nearly so, extending almost to middle of harpe. Aedeagus with a rather large patch of spinelike cornuti. Frema.e (fig. 115)—Alar expanse 18-25 mm. Similar to male in color and maculation. Genitalia (fig. 93): postgenital plate strongly sclerotized, concave, scobinate. Sclerotized part of ductus bursae between ostium and origin of the ductus seminalis about as wide as long; ductus bursae rather conspicuously enlarged adjacent to origin of ductus seminalis, with longitudinal grooves. NO. 3561 LOXOSTEGE—CAPPS 29 Typr.—Female, U.S. National Museum, USNM 14432, genitalia slide HWC 6002. TYPE-LOCALITY.—Cuernavaca, Mexico. Foop pLAntT.—Unknown. DisTRIBUTION.—MEXIco: Federal District: Mexico City. Morelos: Cuernavaca. Vera Cruz: Jalapa. San Luis Potosi: Tamazunchale. Michoacan: Zitacuaro. Chiapas: Soconusco. Colima. SPECIMENS EXAMINED.—13. In FLIcHT.—May and June. Remarks.—In collections, specimens of autocratoralis have been confused with those of helvialis and Phlyctaenodes cupreicostalis Dyar, but the presence of a subterminal line on the forewing of autocratoralis readily distinguishes it from those two species. Heretofore, auto- cratoralis has been known only by females. Loxostege corozalis, new species Figures 40, 92, 116 Phlyctaenodes helvialis (Walker).—Dyar, 1914, Proc. U.S. Nat. Mus., vol. 47, p. 281. Mate (fig. 116)—Alar expanse 18-23 mm. Frons conical. Antenna weakly ciliate. Midtibia incrassate; hair-pencil present. Outer spur less than one-half as long as inner. Resembles auto- cratoralis in color and maculation but with subterminal line of fore- wing weaker, often obsolete posteriorly; hindwing usually with an obscure, very narrow ochreous area between subterminal line and termen. Genitalia (fig. 40): Similar to those of autocratoralis but harpe with a cluster of digitate setae; dorsal margin of sacculus shorter, terminating distinctly before middle of harpe; cornuti of aedeagus much weaker, the patch scobinate or obsolescent. FremMaLE.—Alar expense 17-23 mm. Similar to male in color and maculation. Genitalia (fig. 92): somewhat like those of autocratoralis but with sclerotization of postgenital plate weaker; ductus bursae between ostium and origin of ductus seminalis narrower, the sclerotiza- tion longer, about two times as long as wide and without conspicuous longitudinal grooves or enlargement adjacent to origin of ductus bursae. Typr.—Male, U.S. National Museum, USNM 67610, genitalia slide HWC 7864. TyYprE-LocaLiry.—Corozal, Panama Canal Zone. PARATYPES.—PANAMA CANAL ZONE: type-locality, 3 o, 8 2; Barro Colorado Is., Gatun Lake, 307, 2 9; Tabernilla, 1 9. PANAMA: La Chorrera, 3 9. costa rRIcA: Avangarez, 1 co’; Juan Vinas, 2 o, 1 9. GUATEMALA: Cayuga, 1 9. Mexico: Vera Cruz: Jalapa, 1 9; San Luis 30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Potosi: Tamazunchale, 1 9; Chiapas: Santa Anita, 1 o’, 1 2; Soconusco (Finca la Violeta), 2 o, 2 9; Tepechual, 19. Yucatan: Chichen Itza, 21 o, 30 9; Oaxaca: Tuxtepec, 1 o&, 19. Morelos: Cuernavaca, 2 9. Nayarit: El Pautanal, 19. Zacualpan: 19. Paratypes in collections of the U.S. National Museum, American Museum of Natural History, Cornell University, and the Canadian National Collection. Foop pLant.— Unknown. In rLicHT.—April to November. Remarks.—Labels in collections indicate confusion of corozalis with both helvialis and mancalis. The presence of a subterminal line on the forewing of corozalis distinguishes it from helmalis, and the transverse posterior line on the forewing of mancalvs is more denticulate than that of corozalis. Loxostege huachucalis, new species Figures 46, 75, 124 Mats (fig. 124).—Alar expanse 21-24 mm. Frons conical. Anten- na weakly ciliate. Midtibia incrassate, hair-pencil weak. Outer spur one-fourth as long as inner spur. Resembles mancalis in color and maculation but with markings somewhat more ferruginous, lines more denticulate, definition sharper, and postmedial line of hindwing broadly concave. Genitalia (fig. 46): Harpe without digitate setae; sacculus with two conspicuous, subequal spines and an outer cluster of small spines; pad with distal margin concave or somewhat angulate medially, lower element densely spinulate, spinules moderately long and slender. Aedeagus stout, somewhat bifid distally; the attennuate sclerotiza- tions subequal, with the dorsal one denticulate. Frmatre.—Alar expanse 21-24 mm. Similar to male in color and maculation. Genitalia (fig. 75): eighth sternum conspicuously pig- mented and strongly scobinate, midventral configuration as illustrated. Ductus bursae strongly sclerotized from ostium to origin of ductus seminalis and conspicuously reduced in size from midway between ostium and origin of ductus seminalis. Typr.—Male, U.S. National Museum, USNM 67611, genitalia slide HWC 7834. Typr-LocaLiry.—Paradise, Cochise Co., Ariz. Paratyprs.—Arizona: type-locality, 1 o&, 2 9; so. Arizona, 3 o, 3 9; Palmerlee, 6 &, 6 9; Redington, 1 9; Huachuca Mts., 1c’, 4 Q; Madera Canyon, Santa Rita Mts, 3 #,4 9. Paratypes in collections of the U.S. National Museum and the Los Angeles County Museum. Foop puant.—Unknown. In FLIGHT.—June to August. | | NO. 3561 LOXOSTEGE—CAPPS oi Remarks.—L. huachucalis resembles coloradensis much more closely in genitalia than in habitus. In the male genitalia of huachucalis the pad is narrower and more extenuate and the aedeagus is stouter, with distal bifid elements more strongly developed than in coloradensis. The female genitalia of coloradensis are distinguished from those of huachucalis by a difference in the midventral configuration of the eighth sternum, the inconspicuous reduction in width of the ductus bursae from midway between the ostium and the origin of the ductus seminalis, and the stronger spinulation adjacent to junction of the ductus bursae and bursa copulatrix. Loxostege marialis, new species Fiaures 47, 76, 125 Mate (fig. 125).—Alar expanse 24 mm. Frons conical. Antenna weakly ciliate. Midtibia incrassate; hair-pencil distinct, white. Outer spur one-fourth as long as inner. Similar to hauchucalis but with ground color less ochreous, markings somewhat finer, wings slightly more hyaline. Genitalia (fig. 47): Harpe without digitate setae; pad moderately broad, spinulate; sacculus with two conspicuous subequal spines. Aedeagus stout, median constriction conspicuous, bifid distally, the dorsal element strongly sclerotized, narrow, spinulate. Frema.r.—Alar expanse 24-26 mm. Similar to male in color and maculation. Genitalia (fig. 76): ductus bursae of approximately same width from ostium to origin of ductus seminalis; midventral configura- tion of eighth sternum widest at ostium, lateral margins subparallel, pigmentation and granulation ending before ostium. Typr.—Male, U.S. National Museum, USNM 67612, genitalia slide HWC 7903. TypE-LocaLity.—Volcan Santa Maria, Guatemala. ParaTyYPEs.—Guatemala: type-locality, 3 o, 3 9; Duenas, 1 9; Calderas, 1 &. Paratypes in the U.S. National Museum, British Museum (Nat. Hist.), and Canadian National Collection. Foop pLtant.— Unknown. In FLIGHT.—June. Remarks.—The lack of an outer cluster of small spines on the sacculus, the broader pad, and stouter aedeagus of marialis distin- guishes its males from those of huachucalis. The nonpigmented, smooth, membranous, concave area at the base of the midventral con- figuration, in combination with the rather inconspicuous constriction of the ductus bursae midway between the ostium and the origin of the ductus seminalis, is diagnostic of the females of marialis. The paratypes from Calderas and Duenas are from the syntype series of Phlyctaenodes conisphora Hampson. 32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Loxostege purulhalis, new species Fieures 45, 126 Ma tx (fig. 126).—Alar expanse 20-21 mm. Frons conical. Antenna weakly ciliate. Midtibia weakly incrassate, hair-pencil obsolescent. Similar to marialis in color and maculation, but somewhat smaller than that species. Genitalia (fig. 45): Harpe without digitate setae; pad broad, ventral half spinulate; dorsal margin of sacculus with a single spine from near middle and an outer cluster of short, stout spinules. Aedeagus slightly bifid distally, the dorsal element short, attenuate, and with short, stout spinules. FremaLe.— Unknown. Type.—Male, U.S. National Museum, USNM 67613, genitalia slide HWC 7905. Typr-LocaLity.—Purulha, Guatemala. ParRATYPE.—Type-locality, 17; in collection of the U.S. National Museum. Foop piuant.—Unknown. IN FLIGHT.—July. Remarks.—No reliable differences in color and maculation were noted to distinguish purwlhalis from martalis; however, in purulhalis the outer spur is longer and the hair-pencil is weaker than in marialis. The genitalia of purulhalis are more like those of huachucalis than of marialis, but the presence of only one median spine from the sac- culus and the wider pad distinguish the males of purulhalis from those of huachucalis. Loxostege conisphoralis, new species Fiaures 44, 80, 127 Mats (fig. 127).—Alar expanse 22-23 mm. Frons conical. Antenna weakly ciliate. Midtibia incrassate; hair-pencil present, white. Outer spur one-fifth as long as inner spur. Color and maculation similar to that of marialis but hindwing with brownish or fuscous accentuation of veins 3 and 4 adjacent to cell and postmedial line not as broadly concave. Genitalia (fig. 44): Harpe without digitate setae; pad rather elon- gate and of uniform width, densely spinulate distally; sacculus with a conspicuous scalpel-like spine arising from near middle of dorsal margin and a rather large outer cluster of subequal spines. Aedeagus with an inconspicuous patch of minute spinules as cornuti. Fremaue.—Alar expanse 22-26 mm. Similar to male in color and maculation. Genitalia (fig. 80): ductus bursae strongly sclerotized from ostium to origin of ductus seminalis, the sclerotization about as wide as long; median ventral configuration of eighth sternum below NO. 3561 LOXOSTEGE—CAPPS 33 ostium as figured; granulation strong at junction of ductus bursae and bursa copulatrix. Tyrr.—Male, U.S. National Museum, USNM 67614, genitalia slide HWC 7901. Typr-LocaLity.—Chiapas, Mexico. PARATYPES.—MEXIcO: Chiapas: “Santa Anita,” 1 @; La Granja, 1 o'; Soconusco (Finca La Violeta), 33 o&, 5 9. Vera Cruz: Santa Lucrecia, 1 9. Yucatan: Chichen Itza, 1 9. GuaTEMmaLa: Chejel, 1 9; Volcan Santa Maria, 1 co. costa RIcA: Juan Vinas, 19. Para- types in the U.S. National Museum, American Museum of Natural History, and the Canadian National Collection. Foop puant.— Unknown. In FLigHt.—June to November. Remarks.—The scalpel-like spine arising from the dorsal margin of the sacculus and the character of the outer cluster of spinules are diagnostic for the males of conisphoralis, and the midventral configura- tion of the eighth sternum, in combination with the short, broad sclerotization of the ductus bursae between the ostium and origin of the ductus seminalis, distinguish the females of conisphoralis from all others in the genus. Loxostege conisphora (Hampson), new combination FIGuRES 23, 73, 128 Phlyctaenodes conisphora H ampson, 1918, Ann. Mag. Nat. Hist., ser. 8, vol. 11, pe old Mats (fig. 128).—Alar expanse 22 mm. Frons conical. Antenna weakly ciliate. Midtibia incrassate, hair-pencil present, white. Outer spur about one-fifth as long as inner spur. Resembles marialis in color and maculation but wings with brownish irroration and defini- tion of subterminal lines stronger, and veins 3 and 4 of hindwing with fuscous accentuation adjacent to cell. Genitalia (fig. 23): Harpe with digitate setae; medial spine long, slender, bifid distally; dorsal margin of sacculus with outer half broadly convex. Fremae.—Alar expanse 24mm. Color and maculation as in male. Genitalia (fig. 73) similar to those of neomarculenta but with ventral margin of ostium deeply incised medially and lateral margins of the two pronglike projections strongly denticulate. Lecroryre.—Male, British Museum (Nat. Hist.), genitalia slide BM no. 9588, present designation. TypE-LocaLiry.—Calderas, Guatemala. PARALECTOTYPE.—Irazu, Costa Rica, 1, British Museum (Nat. Hist.), genitalia slide HWC 17,451, present designation. DistRIBUTION.—Guatemala, Costa Rica, and Mexico. Feil eet 34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Foop pLant.— Unknown. SPECIMENS EXAMINED.—3. Remarks.—Dissection of the syntypes of conisphora revealed the presence not only of conisphora and marialis, but another undescribed species, the discussion of which follows that of the next species. Loxostege jaliscalis, new species Maue.—Alar expanse 25 mm. Frons conical. Antenna weakly ciliate. Midtibia incrassate; hair-pencil present. Outer spur about one-fifth as long as inner spur. Resembles conisphoralis but differs from it as follows: wings more ochreous, pale straw yellow, and without irroration, markings pale buff; transverse and postmedial lines wider and smoother; subterminal and terminal lines weaker and more diffuse; fringe pale buff; hindwing with accentuation of veins 3 and 4 adjacent to cell absent or obsolescent. Genitalia: Like those of conisphoralis but with uncus somewhat narrower, spine and distal bifurcation stronger, and distal convex production of sacculus much reduced. Frema.n.—aAlar expanse 25mm. Similar to male in color and macu- lation, except accentuation of veins 3 and 4 of hindwing at cell is stronger. Genitalia similar to those of conisphoralis but with scleroti- zation of ductus bursae from ostium to origin of ductus seminalis shorter, conspicuously constricted about midway between ostium and angulate lateral production; and median incision of ventral margin of ostium concave and not as deep as in conisphoralis. Type.—Male, Canadian National Collection, genitalia slide HWC 17,617. TYPE-LOCALITY.—San Luis Potosi, Mexico [46 mi. N.]. ParatyPr.—Tepititlan, Jalisco, Mexico, 1 9. Canadian National Collection. Foop pLant.— Unknown. In FLicHtT.—August and September. Loxostege volcanensis, new species Figures 49, 81, 129 Mate (fig. 129)—Alar expanse 12-22 mm. Frons conical. An- tenna pubescent, cilia very short. Midtibia incrassate, hair-pencil brownish. Outer spur one-sixth as long as inner spur. Resembles conisphoralis in color and maculation but hindwing with postmedial line smoother, not conspicuously bent inward at vein 3, origins of veins 3 and 4 without adjacent brownish accentuation. Genitalia (fig. 49): Uncus conspicuously narrower apically. Harpe without digitate setae; pad narrow, densely spinulate distally; sacculus with large spine, greatly expanded basally, arising from near middle NO. 3561 LOXOSTEGE—CAPPS 35 of dorsal margin. Aedeagus stout, cornutus an elongate, serrate, sclerotized patch. FremMaLe.—Alar expanse 19-21 mm. Similar to male in color and maculation. Genitalia (fig. 81): production of ductus bursae ventrad of the ostium with conspicuous medial emargination. Type.—Male, U.S. National Museum, USNM 67615, genitalia slide HWC 6004. TypxE-LocaLity.—Volcan Santa Maria, Guatemala. PARATYPES—GUATEMALA: type-locality, 2 o@. mexico: Cordoba, 1 9; Orizaba, 1 9; Jalapa, 2 9; Oaxaca, 1 o; Chiapas: La Granja, 1 9; Soconusco (Finca La Violeta), 5 @. costa rica: Juan Vinas, 2 9, Irazu, 1 9. VENEZUELA: Rancho Grande (near Maracay), 1 9. Paratypes in the U.S. National Museum, American Museum of Natu- ral History, Canadian National Collection, and British Museum (Nat. Hist.). Foop pLtant.—Unknown. In FLIGHT.—January to October. Remarks.—The attenuate character of the uncus distinguishes the males of volcanensis from all others of the genus, and the conspic- uous median emargination of the production of the ductus bursae below the ostium is diagnostic for the females of the species. The female from Irazu is from the syntype series of conisphora. Loxostege mellinialis (Druce), new combination Figures 42, 91, 143 Epichronistis mellinialis Druce, 1899, in Godman and Salvin, Biologia Centuali- Americana, Zoology, Lepidoptera, Heterocera, vol. 2, p. 559. Phlyctaenodes phrizalis Dyar, 1914, Proc. U.S. Nat. Mus., vol. 47, no. 2059, p. 395. [New synonymy.] Ma.r.—Alar expanse 28-32 mm. Frons conical. Antenna weakly ciliate. Midtibia incrassate; hair-pencil well developed, white. Outer spur one-third as long as inner spur. Forewing straw-yellow, slightly irrorated with buff. Markings as in female (fig. 42). Hind- wing somewhat paler than forewing, without buff irroration; post- medial line terminating at fold between veins le and 2; brownish patch adjacent to base of vein 2 variable in size and intensity; wedge- like brownish accentuation of veins at termen variable in intensity, occasionally absent or obsolescent. Genitalia (fig. 42): Harpe with digitate setae; a conspicuous denticu- late production arising from dorsal margin of sacculus; pad spinulate ventrally, brushlike. Aedeagus bifid distally, elements subequal, the dorsal one attenuate and spinulate. Frmatx (fig. 143).—Alar expanse 27-31 mm. Similar to male in color and maculation. Genitalia (fig. 91): eighth sternum finely 36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 spinulate, rather smooth in appearance but with a few transverse folds or ridges and a pair of pouchlike receptacles laterad of the ostium, their long axes transverse. Typrs.—British Museum (Nat. Hist.): mellinialis. U.S. National Museum: phrizalis, USNM 16510, genitalia slide HWC 6014. TYPE-LOCALITIES.—Duenas, Guatemala (mellinialis); Zacualpan, Mexico (phrixalis). DISTRIBUTION.—GUATEMALA: Duenas. mexico: Zacualpan. UNI- TED sTATES: Arizona [new record for the United States]. Foop pLaAnt.—Unknown. SPECIMENS EXAMINED.—25. In FLicHTt.—July to September. Remarks.—L. mellinialis and phrixalis were previously known only from the types, and I am indebted to Mr. Whalley for comparison of my material with that of mellinialis in the British Museum (Nat. Hist.). Loxostege venadialis, new species Ficures 43, 90, 145 Maur.—Alar expanse 32 mm. Frons conical. Antenna ciliate, length of cilia equal to width of shaft. Midtibia incrassate; hair- pencil well developed, white. Outer spur slightly less than one-half as long as inner spur. Color and maculation as in mellinialis but with definition of markings weaker, without wedgelike buff or brownish accentuation of veins at termen, and second and third segments of labial palpus paler. Genitalia (fig. 43): Similar to those of mellinialis but with uncus stouter, sacculus wider with dorsal margin somewhat undulate, aedeagus more robust and longer. FreMAte (fig. 145)—Alar expanse 27-31 mm. Similar to male in color and maculation. Genitalia (fig. 90) resemble those of mel- linialis but with eighth sternum more strongly spinulose, with con- spicuous undulate furrows, and with pouchlike receptacles much larger. Typr.—Male, U.S. National Museum, USNM 67616, genitalia slide HWC 6013. TYPE-LOCALITY.—Venadio, Sinaloa, Mexico. ParatyPEs.—Type-locality, 1 o&, 3 2. Foop pLant.— Unknown. In FiticHtT.—No date of collection. Remarks.—The slightly longer cilia and outer spur distinguish the males of venadialis from those of mellinialis. The absence of the NO. 3561 LOXOSTEGE—CAPPS 3 wedgelike accentuation of the veins at the termen in venadialis dis- tinguishes its females from those of well marked specimens of mel- linialis; if specimens are worn, the genitalia must be examined for the reliable separation of those two species. Loxostege labeculalis (Hulst) Ficures 58, 102, 140 Botis labeculalis Hulst, 1886, Trans. American Ent. Soc., vol. 13, p. 152. Lozostege labeculalis (Hulst).—Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull. OZ p. 384.—Barnes and McDunnough, 1917, Checklist of the Lepidoptera of boreal America, p. 132.—McDunnough, 1939, Southern California Acad. Sci. Mem., vol. 2, no. 1, p. 13. Maue.—Alar expanse 16-20 mm. Conical production of frons obsolescent or absent. Antenna weakly ciliate. Midtibia weakly incrassate; hair-pencil weak. Outer spur one-fourth as long as inner spur. Head, thorax, and abdomen ochreous. Ground color of wings pale ochreous, markings dark brown with reddish tinge. Genitalia (fig. 58): Uncus triangulate, apex pointed, slightly blunt. Harpe rather narrow, with digitate setae, clasper short, stout, curved, spinulate distally; sacculus with conspicuous upturned slender spine arising from dorsal margin. Anellus stout. Aedeagus with subequal bifurcate sclerotization and patch of deciduous spines. Large element of bifurcation elongate, somewhat concave, dilated distally; small ele- ment a short, spurlike projection from near base of larger. FEMALE (fig. 140).—Alar expanse 20-23 mm. Color and macula- tion as in male. Genitalia (fig. 102): ostial pouch large, width about equal to depth; ductus bursae extending beyond anterior margin of pouch. Typre.—Female, American Museum of Natural History. TYPE-LOCALITY.—Arizona. Foop purant.— Unknown. DisrrisutTion.—Arizona: Cochise Co.: Palmerlee, Paradise; Hua- chuca Mts., Chiricahua Mts.; Pima Co.: Baboquivari Mts. (Carr Canyon, Pepper Sauce Canyon); Santa Rita Mts., Madera Canyon. Texas: Brewster Co. SPECIMENS EXAMINED.— 39. In Fiicut.—July to September. Remarxs.—The spurlike projection of the aedeagus varies from obsolescent to well developed, and the serration of the margins of the larger element ranges from nearly smooth to distinctly serrate. The ostial pouch is also somewhat variable in shape, particularly adjacent to the origin of the ductus seminalis. 38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Loxostege intinctalis (Dyar), new combination Figure 141 Pachyzancla intinctalis Dyar, 1920, Insec. Inse. Menstr., vol. 8, p. 34. Mats (fig. 141).—Alar expanse 21 mm. Conical production of frons obsolescent or absent. Antenna weakly ciliate. Midtibia incrassate, hair-pencil distinct. Color and maculation as in labeculalis but with brownish accentuation of veins at termen more wedgelike. Genitalia like those of labeculalis but with uncus broader. Frema.e.—Alar expanse 20 mm. Similar to male in color and macu- lation. Genitalia similar to those of labeculalis but larger, with ductus bursae wider and ostial pouch somewhat differently shaped. Type.—Male, U.S. National Museum, USNM 22762, genitalia slide HWC 17,241. TypE-LOcALITY.—Zacualpan, Mexico. Paratypr.—Female, U.S. National Museum, Guadalajara, Mexico, via Schaus collection, genitalia slide HWC 17,242. Foop pLANnt.— Unknown. Remarks.—L. intinctalis is known only from the type and paratype, so the differences distinguishing them from specimens of labeculalis may be individual rather than of specific value. Until this can be deter- mined by examination of additional material of intinctalis, however, it seems best to treat it as a distinct species. Loxostege federalis, new species Fiaures 60, 103, 148 Mats (fig. 148).—Alar expanse 22-24 mm. Frons round. Antenna ciliate, length of cilia about equal to width of shaft. Midtibia weakly incrassate; hair-pencil obsolescent or absent. Outer spur about one- third as long as inner. Ground color of wings straw yellow, markings brownish. Genitalia (fig. 60) similar to those of labeculalis but with uncus longer and more extenuate distally. Fremate.—Alar expanse 23 mm. Similar to male in color and maculation. Genitalia (fig. 103) similar to those of labeculalis but with posterior apophyses longer and a conspicuous constriction of eighth sternum adjacent to ostial opening. Typr.— Male, U.S. National Museum, USNM 67623, genitalia slide HWC 10,005. TypE-LocaLity.— Popocatepetl Park, Mexico, D.F. ParatypEs.— Type locality, 3 o, 2 2; in collection of U.S. National Museum. Foop pLrant.— Unknown. In FLIGHT.—June and July. NO. 3561 LOXOSTEGE—CAPPS 39 Loxostege entephrialis (Schaus), new combination Figures 29, 147 Pyrausta entephrialis Schaus, 1912, Ann. Mag. Nat. Hist., ser. 8, vol. 9, p. 311. Mats (fig. 147).—Alar expanse 25 mm. Frons conical. Antenna ciliate, cilia slightly longer than width of shaft. Midtibia weakly incrassate; hair-pencil obsolescent or absent. Wings: ground color sordid white with an ochreous tinge; irrorated with light brown; markings brownish fuscous. Genitalia (fig. 29): Uncus about as broad as long. Harpe with digitate setae; pad folded, spinulation dense and fine; a conspicuous broad, concave, spurlike projection arising from near middle of harpe and extending below lower margin; sacculus with small distal triangu- late production on dorsal margin. Aedeagus with patch of slender, spinelike cornuti and small finlike basal keel. Fremate.— Unknown. TypE.— Male, U.S. National Museum, USNM 17596, genitalia slide HWC 7952. TyPE-LocALity.—Juan Vinas, Costa Rica. Foop pLiant.— Unknown. In FLIGHT.— October. Remarks.— L. entephrialis is known only from the type. Loxostege cupreicostalis (Dyar), new combination Ficures 60, 104, 142 Phlyctaenodes cupreicostalis Dyar, 1913, Proc. U.S. Nat. Mus., vol. 44, no. 1951, p. 318. Mate (fig. 142).—Alar expanse 21-27 mm. Conical production of frons obsolescent or absent. Antenna somewhat pubescent, cilia very short. Midtibia incrassate, hair-pencil well developed, white. Wings pale yellow, somewhat lustrous and subhyaline, markings brownish; subterminal line absent. Genitalia (fig. 60): Harpe without digitate setae; pad somewhat modified, with distal cluster of rather strong spinules; dorsal margin of sacculus with a short spinelike projection adjacent to emargination, near middle. Aedeagus with a patch of deciduous spinelike cornuti, spinules strongly dilated basally. Fremate.—Alar expanse 25 mm. Similar to male in color and the maculation. Genitalia (fig. 104): the character of sclerotization of the ductus bursae from ostium to origin of the ductus seminalis, in combination with the conspicuously narrow, elongate elements of signum, is diagnostic. LecrotyPpre.— Male, U.S. National Museum, USNM 14449, genitalia slide HWC 6009. 40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 TyprE-LocALiry.— Guadajalara, Jalisco, Mexico. PaRrALEctotyPrE.— Female, U.S. National Museum, genitalia slide HWC 6010. Foop piant.— Unknown. DISTRIBUTION.— MEXICO: Guerrero: Sierra de Guerrero; Iguala. Puebla: Tehuacan. Jalisco: Guadalajara. Chiapas: Soconusco. Sonora: Mainari. VENEZUELA: Aroa. UNITED STATES: Arizona: Baboquivari Mts. [new record for the United States]. SPECIMENS EXAMINED.—9. In FuigHt.— May to October. Remarks.—According to the original description, the species was based on two cotypes, a male and a female. The male, labeled type in Dyar’s script, and the female are hereby designated lectotype and paralectotype, respectively. Although eupreicostalis may eventually require placement elsewhere, it is best placed in Lozostege pending a revision of the subfamily. Loxostege subcostalis (Dyar), new combination Figure 149 Lygropia subcostalis Dyar, 1912, Proc. U.S. Nat. Mus., vol. 42, no. 1885, p. 104. —Klima, 1939, in Junk, Lepidopterorum catalogus, pt. 94, p. 233. Mats (fig. 149).— Alar expanse 15-18 mm. Fronsconical. Antenna weakly ciliate, somewhat pubescent, black with tip white. Midtibia weakly incrassate; hair-pencil obsolescent or absent. Outer spur one-half as long as inner spur. Upper surface of wings brownish yellow, unicolorous; under surface as above, except costa of forewing margined with black. Labial palpus with first and second segments brownish yellow; third segment black. Fore-, mid-, and hindtarsi blackish fuscous. Genitalia similar to those of the species treated next but with the aedeagus of subcostalis lacking the conspicuous, strong, rather long, hooklike distal cornutus. Fremaute.— Unknown. Typpn.— Male, U.S. National Museum, USNM 14208, genitalia slide HWC 7995. TypE-LocaLiry.—Orizaba, Mexico. Foop pLANT.— Unknown. DistrRiBuTION.—MExIco: Vera Cruz: Orizaba and Santa Tosa. GUATEMALA: Chejel. costa rica: Avangarez. UNITED STATES: Texas: Dallas [new record for the United States]. SPECIMENS EXAMINED.—S8. In FitigHt.— May to August. Remarks.—The Texas specimen is labeled ‘“Botis obauratalis NO. 3561 LOXOSTEGE—CAPPS 41 Hulst, Type,” but this appears to be a manuscript name since no record of its description was found in the literature. Dyar erred in referring to the tibiae and tarsi as black; only the tarsi are blackish. The tibiae are brownish yellow. Loxostege sinaloensis, new species Fieure#s 13, 150 Mate (fig. 150).—Alar expanse 15-17 mm. Frons conical. An- tenna weakly ciliate, somewhat pubescent, brownish yellow with tip white. Midtibia weakly incrassate; hair-pencil obsolescent or absent. Outer spur one-half as long as inner spur. Upper surface of wings brownish yellow, unicolorous; under surface same as upper, except costa of forewing narrowly margined with black. Labial palpus with all segments brownish yellow. Entire foretarsus fuscous; first seg- ment of mid- and hind tarsi pale fuscous, remainder of segments brownish yellow. Genitalia (fig. 13): Uncus bluntly pointed. Harpe without digi- tate setae; clasper bifid. Aedeagus with one large, conspicuous, hookhke distal cornutus and a few patches of small, slender spines; basal keel well developed. Frema.te.— Unknown. Typr.—Male, U.S. National Museum, USNM 67617, genitalia slide HWC 7997. TyYPE-LOCALITY.—Venadio, Sinaloa, Mexico. ParatyPEes.—Type locality, 4 o. Foop pLant.— Unknown. Remarks.—L. sinaloensis and subcostalis are closely related; but the differences in coloration of the antenna, third segment of the labial palpus, and tarsi readily distinguish them from each other. The genitalia of both stnaloensis and subcostalis resemble those of helmialis and pseudohelvialis, but the former two species have stouter bifid claspers of the harpe and a different aedeaga! armature. Loxostege chiapasalis, new species Figures 34, 151 Mats (fig. 151).—Alar expanse 22 mm. Frons conical. Antenna ciliate, cilia slightly shorter than width of shaft. Midtibia incrassate; hair-pencil present. Outer spur minute, about one-eighth as long as inner spur. Head, thorax, and abdomen brownish yellow; frons margined with white; thorax and abdomen paler below. Upper and under surfaces of fore- and hindwings brownish yellow, without markings. Labial palpus: first segment with basal half white, re- mainder blackish above, fringe fuscous intermixed with white; second 42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 segment blackish above and laterally, fringe white; third segment blackish. Inner surface of foretibia fuscous, outer surface pale ochreous. Midtibia with a brownish dorsal band, remainder ochre- ous; hind tibia ochreous. All tarsi pale, smoky fuscous. Genitalia (fig. 34): Uncus stout. Harpe with digitate setae; pad bifurcate distally, the elements subequal; a crenulate production anterior to median angle of bifurcation; spinulation restricted to area below crenulate production; sacculus with two widely separated spines. Aedeagus without cornuti. FEMALE.— Unknown. Typp.—Male, US. National Museum, USNM 67618, genitalia slide HWC 17,092. TyPE-LOCALITY.—Chiapas, Mexico. Foop pLant.— Unknown. Remarks.—The species, known only from the type, resembles Psara nigripes Schaus, but chiapasalis is smaller and somewhat more brownish yellow, and the midtibia is brownish fuscous dorsally. The genitalia of chiapasalis are similar to those of cayugalis, differing chiefly in the modification of the pad. Loxostege nigripes (Schaus), new combination FIGURES 32, 68, 152 Psara nigripes Schaus, 1920, Proc. Ent. Soc. Washington, vol. 22, p. 190. Mate (fig. 152).—Alar expanse 28-30 mm. Frons conical. An- tenna ciliate, cilia one-half as long as width of shaft. Midtibia incrassate, hair-pencil weak. Outer spur one-fifth as long as inner spur. Wings above and below orange, without markings. Labial palpus: first segment whitish at base, remainder fuscous; dorsal half of second segment fuscous, remainder whitish; third segment fuscous. Foretibia with some fuscous, more extensive on inner side; mid- and hind tibia orange. All tarsi blackish. Genitalia (fig. 32): Uncus stout. Harpe with digitate setae; pad moderately narrow, spinulation rather coarse; a conspicuous spine arising from sacculus, with distal portion bent horizontally. Aedeagus without cornutus. Fremate.—Alar expanse 27-30 mm. Similar to male in color and maculation. Genitalia (fig. 68): ostium rather broad; ductus bursae eradually tapered from ostium to constriction adjacent to origin of ductus seminalis. Typp.—Male, U.S. National Museum, USNM 23582, genitalia slide HWC 17,090. TypE-LocaLity.— ‘Cayuga, Guatemala’’? Foop pLtant.— Unknown. DisTRIBUTION.—GUATEMALA: Volcan Santa Maria. MExico: Vera NO. 3561 LOXOSTEGE—CAPPS 43 Cruz: Jalapa, Orizaba; Chiapas: Soconusco. SPECIMENS EXAMINED.—14. In FLIGHT.—June to October. ReEeMARKS.—Schaus cited only Cayuga, Guatemala as habitat for the species, but none of the material in the collection is labeled Cayuga. The type and all of the other specimens in the series from Guatemala are labeled Volcan Santa Maria. Loxostege subcuprea (Dognin), new combination Fiaures 61, 174 Lygropia subcuprea Dognin, 1906, Ann. Mag. Ent. Belgique, vol. 50, p. 121.— Klima, 1939, in Junk, Lepidopterorum catalogus, pt. 94, p. 233. Mate (fig. 174).—Alar expanse 20 mm. Frons conical. Antenna somewhat pubescent. Midtibia incrassate; hair-pencil weak. Outer spur slightly less than one-third as long as inner spur. Fore- and hindwings orange, without markings except for fuscous margins. Labial palpus: first segment and most of second segment orange; distal part of second and all of third segment pale fuscous. Genitalia (fig. 61): Uncus pointed. Harpe without digitate setae; clasper strong, hooklike, arising from near middle of harpe; dorsal margin of sacculus with a cluster of coarse, slender spinules near base and a distal group of finer spinules. Aedeagus with a narrow, straplike ventral sclerotization and a distal group of slender spines; basal keel thin, finlike. Frma.Le.— Unknown. Typrr.—Male, U.S. National Museum, USNM 29562, genitalia slide HWC 17,237. Typr-LocaLity.— Metan, Salta, Argentina. Foop pLaAnt.— Unknown. DISTRIBUTION.— ARGENTINA: Type-locality and Tucuman. SPECIMENS EXAMINED.—3. In FuicHt.— February. Remarks.—The species is definitely not congeneric with Lygropia unicoloralis (Guenée), type of the genus Lygropia Lederer. Loxostege clarissalis (Schaus), new combination Figure 176 Nomophila clarissalis Schaus, 1920, Proc. Ent. Soc. Washington, vol. 22, no. 8, p. 217.—Klima, 1939, in Junk, Lepidopterorum catalogus, pt. 94, p. 382. Mats (fig. 176).—Alar expanse 24 mm. Frons conical. Antenna weakly ciliate. Midtibia incrassate; hair-pencil well developed. Outer spur slightly less than one-half as long as inner spur. Head, thorax, and abdomen pale fuscous; patagia pale orange. Forewing ground color fuscous with orange tinge; orange accentuation of veins 44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 terminating before attaining termen. Hindwing orange, margined with fuscous. Genitalia very similar to those of subcuprea but larger and with uncus somewhat longer and more slender. Frema.e.— Unknown. Typr.— Male, U.S. National Museum, USNM 23822, genitalia slide HWC 17,239. TyPE-LOCALITY.— Peru. Foop pLant.— Unknown. DistTRIBUTION.— Peru. SPECIMENS EXAMINED.— 2. Remarks.— The frons, venation, genitalia, and tibial characters of clarissalis exclude it from the genus Nomophila. Loxostege aemulalis (Dognin), new combination Figure 171 Phlyctaenodes aemulalis Dognin, 1905, Ann. Ent. Soc. Belgique, vol. 49, p. 77. Mate (fig. 171).—Alar expanse 23 mm. Frons conical. Antenna weakly ciliate, cilia about one-half as long as width of shaft. Midtibia incrassate; hair-pencil present. Outer spur one-third as long as inner spur. Forewing smoky fuscous, without markings. Hindwing ochreous orange with smoky fuscous border on costa and outer margin. Genitalia essentially like those of clarissalis but with uncus narrower and more attenuate. Frma.Le.— Unknown. Typr.— Male, U.S. National Museum, USNM 29584, genitalia slide HWC 7949. TypE-LocaLiry.— Charuplaya, Bolivia. Foop puant.— Unknown. SPECIMENS EXAMINED.—1 (known only from the type). In FLIGHT.—June. Remarxs.—Although the genitalia of aemulalis and clarissalis are very similar, the lack of orange accentuation of the forewing veins of aemulalis readily distinguishes it from clarissalis. Loxostege naranjalis (Schaus), new combination FiaureE 173 Lygropia naranjalis Schaus, 1920, Proc. Ent. Soc. Washington, vol. 22, no. 8, p. 213.—Klima, 1939, in Junk, Lepidopterorum catalogus, pt. 94, p. 232. Fremate (fig. 173).—Alar expanse 23 mm. Frons conical. Antenna pubescent. Head fuscous, frons margined with orange. Collar, thorax, and patagia orange. Abdomen orange with a black dorsal stripe from second to anal segment. Fore- and hindwings orange, margins black. NO. 3661 LOXOSTEGE—CAPPS 45 Genitalia with ostium broad; ductus bursae long, coiled, with a collarlike sclerotization near origin of ductus seminalis; bursa copula- trix somewhat granulose. Mate.— Unknown. Typr.—Female, U.S. National Museum, USNM 23810, genitalia slide HWC 17,240. TYPE-LOCALITY.—S4o Paulo, Brazil. Foop prant.— Unknown. Remarks.—L. naranjalis is known only from the type, and without the male, its generic placement is somewhat uncertain; however, its affinity with Lygropia is remote and, although the signum lacks the carina of typical Lozostege, its other characters indicate close relation- ship to that genus. Loxostege mojavealis, new species Figures 172, 175 Ferma. (fig. 172).—Alar expanse 22 mm. Frons conical. Antenna simple. Forewing: ground color of upper surface whitish, strongly irrorated with smoky fuscous, markings blackish; definition of trans- verse anterior and posterior lines rather weak, with that of the latter strongest; area between transverse anterior line and base of wing paler than that between transverse anterior and posterior lines; angulation of transverse posterior line sharp and acute on vein 2. Hindwing yellowish orange, postmedial and subterminal lines fuscous, weak but discernible; terminal line black, thin; fringe blackish. Genitalia some- what similar to those of nigripes but with lateral margins of ductus bursae at ostium straplike, ductus bursae with a conspicuous, narrow sclerotization below origin of ductus seminalis. Matr.— Unknown. Typr.—Female, U.S. National Museum, USNM 67619, genitalia slide HWC 8027. TypE-LocaLity.—Mojave Desert, near Phelan, Calif. Foop piuant.— Unknown. In FuicHT.—April. Remarks.—The species is known only from the type. Although mojavealis is somewhat similar to Lozxostege annaphilalis (Grote), of which no females were available for study, it does not seem likely that they are conspecific, since the wings of mojavealis are narrower than those of annaphilalis and the markings different on both the fore- and hindwing. Definition of the markings is sharper on the forewing of mojavealis, with the transverse posterior line much more angulate and acute on vein 2 than in annaphilalis. The hindwing of annaphilalis is without markings except for the terminal dots and fringe. 46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Loxostege fuscivenalis (Schaus), new combination Figure 177 Lygropia fuscivenalis Schaus, 1920, Proc. Ent. Soc. Washington, vol. 22, no. 8, p. 213.—Klima, 1939, in Junk, Lepidopterorum catalogus, pt. 94, p. 233. Mats (fig. 177).—Alar expanse 23 mm. Frons conical. Antenna somewhat pubescent, cilia very short. Outer spur one-third as long as inner spur. Head and collar orange; patagia orange anteriorly, fuscous posteriorly; abdomen fuscous. Forewing dull grayish brown, without markings except for brownish fuscous accentuation of veins. Hindwing brownish fuscous, accentuation of veins black. Labial palpus: first segment orange; upper half and distal portion of second segment fuscous, remainder orange; third segment fuscous. Genitalia similar to those of the species treated next but with vin- culum narrower and anterior production longer. Frma.te.— Unknown. Typr.—Male, U.S. National Museum, USNM 23811, genitalia slide HWC 17,305. TyPE-LOCALITY.—Peru. Foop pLant.— Unknown. DistTRIBUTION.—Peru. SPECIMENS EXAMINED.—2. Remarxs.—L. fuscwenalis is definitely misplaced in Lygropia and its affinity to this group is remote. Although the genitalia of fusci- venalis are somewhat aberrant, all other characters indicate that it is closely related to Loxostege. Loxostege peruensis, new species FiagureE 178 Matz (fig. 178).—Alar expanse 24 mm. Frons conical. Antenna somewhat pubescent, cilia very short. Outer spur one-third as long as inner spur. Head, collar, palpi, and abdomen entirely fuscous. Fore- and hindwings as in fuscivenalis. Genitalia: Uncus short, broad at base, distal termination a sharp point. Harpe without digitate setae; two slender fingerlike projections dorsad of outer end of sacculus, one directed dorsad, the other ventrad ; sacculus with a conspicuous medial production bearing a cluster of strong spines. Aedeagus with two subequal sclerotizations; the long one narrow, straplike, with distal termination a long, slender, curved hook, the other short, bluntly pointed distally. FremaLe.— Unknown. Typr.—Male, U.S. National Museum, USNM 67620, genitalia slide HWC 17,306. TYPE-LOCALITY.—Peru. Foop pLrant.— Unknown. NO. 3561 LOXOSTEGE—CAPPS 47 ReEemMARKS.—The species is known only from the type and is dis- tinguished from fuscivenalis by its fuscous head, collar, and palpi. Because the genitalia of peruensis and fuscivenalis are so similar, the former may eventually prove to be a Mendelian color variant of the latter; but until its status can be definitely determined, it is desirable that a name be available for it. Loxostege rantalis (Guenée) Fieures 51, 94, 153, 154 Nymphula rantalis Guenée, 1854, in Boisduval and Guenée, Histoire naturelle des insectes: Species général des lépidoptéres, vol. 8, p. 405 Hampson, 1899, Proc. Zool. Soc. London, p. 210 [cited as synonym of Phlyctaenodes similalis (Guenée) in error]. Eurycreon rantalis (Méschler).—von Hedemann, 1894, Stettiner Ent. Zeit., vol. 55, p. 286. [Incorrect citation.] Loxostege similalis var. rantalis (Guenée).—Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull. 52, p. 383. Loxostege similalis form rantalis (Guenée).—Barnes and McDunnough, 1917, Checklist of the Lepidoptera of boreal America, p. 132.—McDunnough, 1939, Mem. Southern California Acad. Sci., vol. 2, no. 1, p. 12. Lozxostege similalis of authors.—Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull. 52, p. 383.—Barnes and McDunnough, 1917, Checklist of the Lepidoptera of boreal America, p. 132.—Forbes, 1920, Cornell Univ. Agric. Exp. Sta. Mem. 68, p. 556.—McDunnough, 1939, Southern California Acad. Sci. Mem., vol. 2, no. 1, p. 12.—Schaus, 1940, Scientific survey of Porto Rico and the Virgin Islands, vol. 12, pt. 3, p. 367.—Smith and Franklin, 1954, Kansas Ent. Soc. Journ., vol. 27, no. 1, p. 27.—Okumura [in part], 1961, California Dept. Agric. Spec. Publ., no. 282, p. 48. [Misidentification.] Phlyctaenodes similalis of authors.—Hampson [in part], 1899, Proc. Zool. Soc. London, p. 210. [Misidentification.] Phlyctaenodes similalis of authors.—Wolcott, 1923, Journ. Dept. Agric. Puerto Rico, vol. 7, p. 192; 1936, Journ. Agric. Univ. Puerto Rico, vol. 20, p. 466. [Misidentification.] Botis siriusalis Walker, 1859, List of the lepidopterous insects in the British Museum (Nat. Hist.), pt. 18, p. 563. Botts licealis Walker, 1859, ibid., p. 563. Ebulea murcialis Walker, 1859, ibid., p. 746. Scopula nestusalis Walker, 1859, ibid., p. 784. Scopula diotimealis Walker, 1859, ibid., p. 785. Scopula crinisalis Walker, 1859, ibid., p. 798. Nephopteryx intractella Walker, 1863, ibid., pt. 27, p. 55. Eurycreon crinitalis (Walker).—Lederer, 1863, Weiner Ent. Monat., vol. 7, p. 376 [invalid emendation].—von Hedemann, 1894, Stettiner Ent. Zeit., vol. 55, p. 286. Botys posticata Grote and Robinson, 1867, Trans. American Ent. Soc., vol. 1, p. 22. Botys subfulvalis Herrich-Schaffer, 1871, Corr.-Blatt. Zool. Min. Ges. Regens- burg, vol. 25, p. 20. Eurycreon communis Grote, 1876, Canadian Ent., vol. 8, p. 99. AS PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Eurycreon collucidalis Méschler, 1890, Abh. Senck. Ges., 16, p. 290.— Gundlach, 1891, Ann. Soc. Espafiola Hist. Nat., vol. 20, p. 248. Phlyctaenodes collucidalis (Méschler).— Wolcott, 1923, Journ. Dept. Agric. Puerto Rico, vol. 7, p. 192; 1936, Journ. Agric. Univ. Puerto Rico, vol. 20, p. 466. Pyrausta caffreii Flint and Malloch, 1920, Bull. Illinois Dept. Reg. Educ. (Nat. Hist. Surv.), vol. 13, art. 10, pp. 304. Loxostege caffreii (Flint and Malloch).—Heinrich [in part,] 1921, Ent. News, vol. 32, p. 57. Mate (fig. 153)—Alar expanse 17-22 mm. Frons conical. Antenna pubescent. Midtibia little, if any, incrassate; hair-pencil absent. Outer spur one-half as long as inner spur’ Forewing: eround color dull ochreous or somewhat brownish with markings slightly darker; transverse anterior line rather smooth, irregularly angulated; transverse posterior line denticulate; both lines often weak, diffuse; discocellular patch conspicuous, discal dot weaker. Hindwing often paler than forewing; postmedial line, if evident, diffuse. Genitalia (fig. 51): Uncus narrowing gradually from base to tip. Harpe with cluster of setae arising from ridgelike modification, long, slender, nondigitate; pad somewhat modified, downcurved, hooklike, spinulate; dorsal margin of sacculus with three conspicuous spines; ventral spine of basal pair distinctly more than half as long as dorsal; dorsal spine upright from base for about one-third its length, thence somewhat outwardly oblique and weakly undulate; outer spine approximately parallel to dorsal margin of sacculus and terminating near middle. Aedeagus with a spinulate, irregular, elongate patch. FEMALE (fig. 154).—Alar expanse 17-43 mm. Coloration usually somewhat darker than that of male, more brownish or occasionally somewhat grayish fuscous, markings stronger. Genitalia (fig. 94) with sclerotization of ductus bursae broadly interrupted between ostium and collarlike structure; lateral margins of sclerotized portion of ductus bursae between ostium and collar-like structure parallel or turning but slightly inward adjacent to ostium; an elongate, spinulate sclerotization on dorsum of ductus bursae midway between collar and ostium. ‘Two signa, subequal; the small one connate, spinulate. Larva (fig. 1)—Mature, 24 mm. long. Head sordid white with reticulate pale amber markings and a small fuscous patch at lateral incision of hind margin. Prothoracic shield sordid white with brown- ish or blackish markings; elongate, or a series of smal! spots on dorsum and a narrow, interrupted pigmentation posterior to seta 1b, suffusion present along lateral margin; posterolateral margin extenuated, passing behind and under spiracle and fusing with prespiracular shield. Pinacula brown or blackish. Pinacula above level of spiracles uniformly concolorous, or with but a small area at base of seta paler; those below level of spiracles with a reduction or absence of pigmenta- NO. 3561 LOXOSTEGE—CAPPS 49 tion in central area much larger, often appearing ringlike. Meso- and metathorax with pinacula of setae Ia-Ib and Ila-IIb longitudinally elongate. Abdominal segments 1-8 with pinacula of setae I and IT large, flat, round; pinaculum of seta ITI somewhat elongate or triangu- lar. Abdominal segment 9 with pinaculum bearing paired setae IT, that of setae I-III strongly pigmented; setae VI, VIJ, and VIII also on well-defined pinacula but with central areas paler. Trpes.—U.S. National Museum: rantalis, female, USNM 67625, genitalia slide HWC 17,401. British Museum (Nat Hist.): striusalis, male with head and abdomen missing; lycealis, male, genitalia slide BM 9683; murcialis, female, genitalia slide BM 9680; nestusalis, male with abdomen missing; diotimealis, female, genitalia slide BM 9679, hereby designated lectotype; crinisalis, male, genitalia slide BM 9681, hereby designated lectotype; intractella, male, genitalia slide BM 9678; communis, male, genitalia slide BM 9682, hereby designated lectotype. Museum der Humboldt-Universitat: collucidalis, male, genitalia slide HWC 17,636. American Museum Natural History: posticata, male, lectotype. Illinois Natural History Survey Collection: caffreizi, male. Location unknown: subfulvalis. TyPpE-LocaLit1es.—Probably ‘“‘Amerique Septentrionale, Am. bor.,” or, “Cuba’’: rantalis. United States: siriusalis, licealis, nestusalis, and intractella. New York: communis. Pennsylvania: posticata. Illinois: caffrevi. Santo Domingo: murcialis, diotimealis. Puerto Rico: collucidalis. Unknown: crinisalis, subfulvalis. Foop pLtants.—Alfalfa, Ambrosia trifida, A. artemisifolia retroflexus, Amaranthus sp., Atriplex patula, Batis maritima, beans, clover, corn, cotton, cowpeas, lambsquarters, Partheniwm argentatum, peas, pigweed, radish, ragweed, Rumex spp., smartweed, soybeans, strawberry, sugarbeets, sunflower, Swiss chard, and thistle. DIsTRIBUTION.—UNITED sTATES: Maine, Connecticut, New York, Maryland, District of Columbia, Virginia, North Carolina, South Carolina, Georgia, Florida, Alabama, Louisiana, Arkansas, Texas, Ohio, Indiana, Illinois, Missouri, Iowa, Kansas, Oklahoma, Colorado, New Mexico, Arizona, and California. wrst INpIES: Bermuda, Bahamas, Cuba, Puerto Rico, St. Croix, and St. Domingo. MExico: Vera Cruz, Puebla, Oaxaca, and Jalisco. SPECIMENS EXAMINED.—335. In FuicuT.—April to November. Remarks.—After examining over one hundred genitalia dissections, I believe that the origins cited by Guenée for rantalis (Montevideo) and similalis (Amérique Septentrionale) were confused. The genitalia of the type of rantalis, a female, are unlike any of those of South American specimens but typical of material from North America, Mexico, and the West Indies. Similarly, the genitalia of the type of 221-525—67—_-4 50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 similalis, a male, are unlike those of North American, Mexican, and West Indies material but are like those of South American specimens, some of which were collected at Montevideo, Uruguay. Furthermore, ‘‘Amerique Septentrionale”’ does not appear on the type label of similalis; the only localities on it are ““Am. bor. Cuba.”’ A line appears to have been drawn through ‘‘Cuba’’ indicating cancellation. I am much indebted to Mr. Whalley for examining, dissecting, and furnishing data on the type material in the British Museum essential for verification of synonymy and selection of lectotypes. The types of striusalis and intractella were found to be males, not females as in- dicated by Walker, and Scopula thoonalis Walker proved to be not congeneric with Lozostege. WS. thoonalis is, therefore, removed from the synonymy but, owing to lack of genitalia, it cannot be assigned elsewhere at present. I am also indebted to Dr. H. J. Hannemann of the Humboldt- Universitat Museum for making available the type of collucidalis for dissection and verification of its synonymy. Loxostege occidentalis (Packard) Figures 52, 96, 155, 156 Scopula occidentalis Packard, 1873, Ann. Lyc. Nat. Hist. New York, vol. 10, . 260. urerean occidentalis (Packard).—Hampson, 1899, Proc. Zool. Soc. London, p. 210 [cited as a synonym of Phlyctaenodes similalis (Guenée) in error]. Lozxostege occidentalis (Packard).—Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull. 52, p. 383 (cited as synonym of Lozostege similalis var. rantalis (Guenée) in error).—Barnes and McDunnough, 1917, Checklist of the Lepidoptera of boreal America, p. 132.—MeDunnough, 1939, Southern California Acad. Sci. Mem., vol. 2, no. 1, p. 12 [cited as synonym of Lozostege similalis form rantalis (Guenée) in error]. Maus (fig. 155)—Alar expanse 18-26 mm. Frons_ conical. Antenna pubescent. Midtibia little, if any, thicker than the hind tibia; hair-pencil absent. Typical specimens similar to darker speci- mens of rantalis in color and maculation, those of occidentalis more grayish fuscous and with definition of narrow, sordid white or ochreous shading along inner side of transverse anterior and outer side of transverse posterior lines of forewing more distinct, terminal dots stronger. Examination of the genitalia, however, is necessary to reli- ably distinguish occasional pale brownish specimens of occidentalis from those of rantalis. Genitalia (fig. 52) similar to those of rantalis but with uncus stouter and the two basal spines of sacculus stouter, subequal; ventral spine one-half or less as long as dorsal. FreMALeE (fig. 156).— Alar expanse 20-23 mm. Similar to male in color and maculation. Genitalia (fig. 96) similar to those of rantalis NO. 3561 LOXOSTEGE—CAPPS 51 but ductus bursae distinctly expanded at ostium and lateral margins somewhat angulate. Lxecrotypr.—Female, Museum of Comparative Zoology, MCZ 14264, genitalia slide HWC 17,459. TypE-LocaLity.— California. Foop pLant.— Unknown. Distripution.— California: Soda Springs, Navarro, San Francisco, Claremont, San Diego, Loma Linda, Fair Oaks, Fresno, Yosemite, and Inyo Co. [no additional locality]. SPECIMENS EXAMINED —47. In Fiigur.— April to October. Remarks.— L. occidentalis was based on two males and two females from California received by Packard from the collections of Edwards and Behrens. Of the three specimens (two males and one female) of occidentalis now in the collection of the MCZ, only the female can be definitely associated with the type series; it is labeled “‘Scopula occt- dentalis Packard, California, Edwards” and is hereby designated lectotype of the species. One of the males has no locality label; the other is from California, but there are no indications that either was Edwards’ or Behrens’ material. Differences in the genitalia, both male and female, distinguish occidentalis from rantalis and similalis and clearly indicate that it is a distinct species. The adults of occidentalis have not yet been associated with larvae. Loxostege similalis (Guenée) Fiaures 50, 95, 157, 158 Nymphula similalis Guenée, 1854, in Boisduval and Guenée, Histoire naturelle insects: Species général des Lépidoptéres, vol. 8, p. 403. Phlyctaenodes similalis (Guenée).—Hampson [in part,] 1899, Proc. Zool. Soc. London, p. 210. Tritea ferruginea Warren, 1892, Ann. Mag. Nat. Hist., ser. 6, vol. 9, p.179. [New synonymy.] Phlyctaenodes ferruginea (Warren).—Hampson, 1899, Proc. Zool. Soc. London, . 210. ieuenies garalis Schaus, 1906, Proc. U.S. Nat. Mus., vol. 30, no. 1444, p. 141. [New synonymy.] Mate (fig. 157).—Alar expanse 17-20 mm. Frons conical. An- tenna pubescent. Midtibia little, if any, incrassate; hair-pencil absent. Outer spur about one-half as long as inner spur. Resembles rantalis but duller in appearance, with ochreous tinge lacking; pos- terior transverse line of forewing usually with pale edging along outer margin. Genitalia (fig. 50): Uncus distinctly extenuate; distal third cylin- drical, hairlike scales short, one-half or less as long as proximal ones. o2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Harpe with basal pair of spines on dorsal margin of sacculus of equal length, or ventral spine slightly longer than dorsal. FrmA.e (fig. 158)—Alar expanse 16-20 mm. Usually somewhat darker than male but otherwise similar in color and maculation. Genitalia (fig. 95) similar to those of rantalis but with dorsum of ductus bursae between collarlike structure and ostium without a platelike sclerotization, or if discernible, obsolescent and nonscobinate. Typrs.— U.S. National Museum: similalis, male, USNM 67626, genitalia slide HWC 7754; garalis, male, USNM 9624, genitalia slide HWC 10,032. British Museum (Nat. Hist.): ferruginea, male. TyYPxE-LOCALITIES.— Probably Montevideo, Uruguay: similalis (see discussion under rantalis). Coquimbo, Chile: ferruginea. Sao Paulo, Brazil: garalis. Food plant and larval stage unknown. DIsTRIBUTION.— BRAZIL: Rio Grande do Sul: Pelotas, Guarani; Mato Grosso: Corumba; Minas Gerais: Lassance; Sd40 Paulo: Rio de Janeiro, Campinas; Castro Parana. uruguay: Montevideo. PARA- Guay: Villarica. ARGENTINA: Santa Fe: San Cristobal; Tucuman: Tucuman, Los Vasquez; Cordoba: Pilar; Jujuy: Ledesma; La Rioja: La Rioja. CHILE: Santiago, Cajon Rio Maipo; Malleco: Angol. PERU: no additional locality. Boxtvra: no additional locality. ECUADOR: Loja. SPECIMENS EXAMINED.— 109. In riiagutT.—January to December. Remarks.—In addition to the differences in genitalic structures, the distributions of simdlalis and rantalis are different, the former being restricted to South America and the latter to the United States, Mexico, and the West Indies. Schaus’ Pyralis garalis agrees with similalis in both maculation and genitalia and is definitely conspecific. J am indebted to Mr. Whalley of the British Museum for comparing material with the type of Tritea ferruginea, which also agrees with similalis in maculation and genitalia and is doubtless conspecific. Although similalis replaces rantalis in South America and appears to be common, it has not attracted attention as a pest there; this may be due to some confusion of its larvae with those of Lowxostege bifidalis (Fabricius), which has been reported as a pest of considerable im- portance. Loxostege bifidalis (Fabricius) Fiaures 54, 98, 164, 165 Phalaena bifidalis Fabricius, 1794, Ent. Syst., vol. 3, pt. 2, p. 232. Phlyctaenodes inornatalis Walker, 1865, List of the lepidopterous insects in the British Museum (Nat. Hist.), pt. 34, p. 1456. NO. 3561 LOXOSTEGE—CAPPS 53 Eurycreon evanidalis Berg, 1875, Deutsche Ent. Zeit., vol. 19, p. 134.— Gundlach, 1890, Ann. Soc. Espafiola Hist. Nat., vol. 20, p. 348. Eurycreon obsoletalis Berg, 1875, op. cit., p. 135. Eurycreon evanadalis [sic] Berg—Moschler, 1890, Abh. Senck. Ges., vol. 16, p. 290. Eurycreon bifidalis (Fabricius)—von Hedemann, 1894, Stettiner Ent. Zeit., vol. 55, p. 286; 1896, ibid., vol. 57, p. 3. Phlyctaenodes bifidalis (Fabricius)—Hampson, 1899, Proc. Zool. Soc. London, p. 210.— Wolcott, 1923, Journ. Dept. Agric. Puerto Rico, vol. 7, p. 192. Lozostege bifidalis (Fabricius).—Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull. 52, p. 383.—Barnes and McDunnough, 1917, Checklist of the Lepidoptera of boreal America, p. 131.—McDunnough, 1939, Southern California Acad. Sci. Mem., vol. 2, no. 1, p. 12.—Schaus, 1940, Scientific survey of Porto Rico and the Virgin Islands, vol. 12, pt. 3, p. 368.—Amsel, 1956 (1954), Bol. Ent. Venezolana, vol. 10, p. 249. Loxostege stolidalis Schaus, 1940, Scientific survey of Porto Rico and the Virgin Islands, vol. 12, pt. 3, p. 368. [New synonymy.] Mate (fig. 164).—Alar expanse 20-24 mm. Frons conical. Antenna pubescent. Midtibia but slightly thicker than hind tibia; hair-pencil absent. Outer spur one-half as long as inner spur. Forewing: ground color pale buff, somewhat glossy; markings brownish, obsolescent, diffuse. Hindwing slightly paler than forewing, without markings except for thin terminal line. Fringe of both fore- and hindwings pale buff. Genitalia (fig. 54): Uncus terminating in a sharp point. Harpe with cluster of strong, slender, nondigitate setae from oblique modi- fication; hooklike sclerotization spinulate distally; production of dorsal margin of sacculus truncate. Aedeagus with numerous strong, spinelike cornuti, variable in number and shape. Femate (fig. 165).—Alar expanse 20-23 mm. Similar to male in color and maculation but with the markings stronger and often more fuscous. Genitalia (fig. 98): ostial margins membranous; ductus bursae strongly sclerotized from slightly below ostium to slightly beyond midpoint between ostium and origin of ductus semi- nalis, somewhat constricted near middle, with upper element spinulate, lower element smooth, broad, collarlike; ductus bursae membranous, nonpigmented from collar to origin of ductus seminalis; ductus seminalis strongly sclerotized and curved, hooklike at origin. Larva.—Mature, 24-27 mm. long. Resembles rantalis but differs from it as follows: posterolateral extension of prothoracic shield behind and under spiracle, and fusion with prespiracular shield weaker, usually discernible only in specimens treated with KOH. Pigmenta- tion of pinacula weaker, interrupted. Meso- and metathorax: pig- mentation of pinaculum of setae Ia-Ib weak, restricted to a small patch anterior to base of setae or occasionally with an additional 54 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 posterior patch; pinaculum of setae Ila-IIb with pigmentation stronger and more extensive, restricted to lower portion of pinaculum, crescent shaped; pinacula of setae III, IV-V, and VI nonpigmented or pigmented at margin only. Abdominal segments 1-7: pigmenta- tion of pinaculum of setae I similar to that of setae [A-IB of meso- and metathorax; pigmentation of pinaculum of seta II stronger than that of seta I, constricted at middle; pigmentation of seta II similar to that of setae Ila-IIb of meso- and metathorax; pinacula below level of spiracle with pigmentation restricted to marginal area or absent. Abdominal segment 8: pigmentation of pinacula of setae I and II more extensive than on anterior segments, usually not inter- rupted on margins but with large central area not pigmented; pina- culum of seta III complete, except for small area at base of seta. Abdominal segment 9: pigmentation of pinaculum of paired setae II stronger on anterior and lateral margins, central area and posterior margin with pigmentation obsolescent or absent; pinaculum of setae I-III with pigmentation complete, except for small area at base of setae; pinacula of setae VI, VII and VIII nonpigmented. Anal shield with pigmentation of pattern variable. Typres.—Location uncertain: bifidalis. British Museum (Nat. Hist.): inornatalis. Museum Natural Sciences Argentina: evanidalis, obsoletalis. U.S. National Museum: stolidalis, USNM 34632. TYPE-LOCALITIES.—‘Americae Insulis,” bifidalis. St. Domingo: inornatalis. Argentina, Buenos Aires: evanidalis, obsoletalis. Puerto Rico, Ensenada: stolidalis. Foop pLants.—Cotton, purslane. DistTRIBUTION.—CANADA: Alberta. UNITED sTATEs: Arizona, New Mexico, Texas. mexico: Colima, Oaxaca, Vera Cruz, Morelos. GUATEMALA: Cayuga. PANAMA: La Chorrera. VENEZUELA: Aroa. coLoMBIA: Tolima. PERU: Lima. BRaziu: Rio Gran do Sul. ARGEN- TINA: Santa Fe, Tucuman. paraauay: Villarica. WEST INDIES: St. Croix, Jamaica, Puerto Rico. SPECIMENS EXAMINED.—179. In FLigHT.—February to December. RemARKs.—The inwardly oblique brownish or somewhat fuscous diffuse bandlike marking which extends from the apex of the forewing to the outer angle of the cell, thence fused with the posterior trans- verse line to the inner margin of the wing, distinguishes bifidalis from the other species heretofore treated herein; it is usually weaker in the males, and is often obsolescent but with remnants discernible. The genitalia of stolidalis Schaus were dissected and they are within the range of interspecific variation of bifidalis. NO. 3561 LOXOSTEGE—CAPPS 55 Loxostege brasiliensis, new species Fiaures 55, 99, 166, 167 Mats (fig. 166).—Alar expanse 22 mm. Frons conical. Antenna pubescent. Midtibia slightly more incrassate than hind tibia; hair- pencil obsolescent or absent. Outer spur about one-third as long as inner spur. Resembles bifidalis but somewhat darker, definition of markings of forewing stronger, and fuscous suffusion adjacent to outer margin of hindwing more extensive. Genitalia (fig. 55): Similar to those of bifidalis but with uncus stouter and lateral margins more convex; sacculus without a somewhat extenuate, truncate production from dorsal margin. Frmats (fig. 167).—Alar expanse 22 mm. Similar to male in color and maculation but with markings stronger. Genitalia (fig. 99) similar to those of bifidalis but collarlike structure narrower, with dorsal and ventral margins parallel. Typr.—Male, U.S. National Museum, USNM 67621, genitalia slide HWC 8022. TypE LocaLity.—Campinas, Sao Paulo, Brazil. ParatyPEs.—From type locality 2 9, genitalia slide HWC 8023, in U.S. National Museum. Foop ptant.— Unknown. In FLicHtT.—December. Loxostege piuralis, new species Fiacures 57, 100, 168, 169 Mats (fig. 168).—Alar expanse 21 mm. Frons conical. Antenna pubescent. Midtibia slightly more incrassate than hind tibia; hair- pencil absent or obsolescent. Outer spur about one-third as long as inner spur. Similar to brasiliensis in color and maculation but fore- wing with veins accentuated with fuscous, hindwing with less suffusion adjacent to outer margin. Genitalia (fig. 57): Similar to those of brasiliensis but with uncus somewhat extenuate from near middle to apex; dorsal margin of sacculus more rounded distally; aedeagus with cornuti spinules much weaker. Frema.e (fig. 169).—Alar expanse 16-20 mm. Similar to male in color and maculation. Genitalia (fig. 100) similar to those of brasili- ensis but with sclerotization at origin of ductus seminalis greatly reduced. Typr.—Male, U.S. National Museum, USNM 67622, genitalia slide HWC 8029. TypzE-LocaLity.—Piura, Peru. 56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Paratypes.—From type-locality, 2 9, genitalia slide HWC 8030, in U.S. National Museum. Foop pLtant.— Unknown. In FiuicHt.—February. Loxostege protealis (Warren), new combination Figures 56, 97, 162 Tritaea protealis Warren, 1892, Ann. Mag. Nat. Hist., ser. 6, vol. 9, p. 178. Phlyctaenodes protealis (Warren).—Hampson, 1899, Proc. Zool. Soc. London, 1 alle Maue.—Alar expanse 21 mm. Frons conical. Midtibia not incras- sate; hair-pencil absent. Outer spur about one-half as long as inner spur. Resembles eneanalis in color and maculation but upper surface of forewing somewhat duller; area in cell between discal dot and reni- form ochreous; terminal dots black, conspicuous; hindwing with post- medial line diffuse but evident. Genitalia (fig. 56): Similar to those of the species described as Pyrausta eneanalis Schaus and treated next, but distinguished from them by the stronger ampulla, longer spine from dorsal margin of sacculus, and character of the cornuti of the aedeagus. Fema ce (fig. 162)—Alar expanse 20-21 mm. Similar to male in color and maculation but with ochreous patch between discal dot and reniform stronger. Genitalia (fig. 97) similar to those of eneanalis but distinguished from them by difference in character of sclerotiza- tion adjacent to ostium, in length of ductus bursae from ostium to origin of ductus seminalis, and degree of sclerotization at origin of ductus seminalis. Lecrotypr.—Male, British Museum (Nat. Hist.). TypE-LocALity.—San Lorenzo Is., Callao, Peru. Foop pLaAnt.— Unknown. DistripuTIoN.—PERU: Trujillo and Angasmarca. SPECIMENS EXAMINED.—#. In ruicHt.—No dates of collection on labels. Remarks.—The species was based on two males and one female, all from the same locality. The male, with genitalia dissected, is hereby designated lectotype of protealis. Loxostege eneanalis (Schaus), new combination Figures 53, 101, 160, 161 Pyrausta eneanalis Schaus, 1923, Zoologica, Sci. Contr. New York Zool. Soc., vol. 5, p. 45. Mate (fig. 160).—Alar expanse 18 mm. Frons conical. Antenna pubescent. Midtibia slightly thicker than hind tibia; hair-pencil obsolescent or absent. Outer spur about one-half as long as inner spur. Resembles darker specimens of occidentalis in color and maculation. NO. 3561 LOXOSTEGE—CAPPS 57 Genitalia (fig. 53): Similar to those of occidentalis, harpe with a single spine arising from dorsal margin of sacculus; an obscure ampulla dorsad of cluster of long, slender setae; spinelike cornuti of aedeagus stronger. FrMa.e (fig. 161).—Alar expanse 17-20 mm. Similar to male in color and maculation. Genitalia (fig. 101): the conspicuous dilation of ductus seminalis in combination with character of sclerotization ad- jacent to ostium, ductus bursae between ostium and origin of ductus seminalis, and ductus seminalis at origin, are diagnostic for females. Typr.— Male, U.S. National Museum, USNM 26524, genitalia slide HWC 17,693. TypE-LocaLity.—Conway Bay, Indefatigable Island, Galapagos Archipelago. Foop pLtant.— Unknown. Disrripution.—Indefatigable and South Seymour Islands, Gala- pagos Archipelago. SPECIMENS EXAMINED.—4. In FLIGHT.— April. RemMArRKs.— Examination of the type revealed that it is a male, not a female as stated by Schaus. Ostrinia obliteralis (Walker), new combination Ficure 146 Botys obliteralis Walker, 1865, List of the lepidopterous insects in the British Museum (Nat. Hist.), pt. 34, p. 1392. Loxostege obliteralis (Walker).—Barnes and MceDunnough, 1917, Checklist of the Lepidoptera of boreal America, p. 131. Pyrausta ainsliet Heinrich, 1919, Journ. Agric. Res., vol. 18, p. 175. [New synonymy.] Typrs.—British Museum (Nat. Hist.): obliteralis, female; U.S. National Museum: ainsliei, male, USNM 22544. Typxr-LocaLitiss.— North America: obliteralis; Knoxville, Tennes- see: ainslier. Remarks.—F. H. Benjamin was the first to note discrepancies in the sex of the type of obliteralis and the application of the name. In his unpublished notes, compiled about forty years ago during his examination of types of American species in the British Museum, he noted that the type of obliteralis was a female and not a male (as stated in the original description), and that the species to which the name was applied in collections in the United States was probably a Pyrausta instead of a Loxostege. It was not until recently, however, that Walker’s obliteralis was definitely determined to be conspecific with ainslier by the comparison of females and genitalia of that species with the type of obliteralis. 58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Hence, ainsliei falls as a synonym of obliteralis. I am indebted to Mr. Whalley of the British Museum and to Dr. Duckworth of the U.S. National Museum for comparison of the material and verifica- tion of the suspected synonymy. Ostrinia penitalis (Grote), new combination Botis penitalis Grote, 1876, Canadian Ent., vol. 8, p. 98. Pyrausta penitalis (Grote).—Fernald, 1903, in Dyar, U.S. Nat. Mus. Bull 52, p. 391. Marion’s (1957) paper transferring nubilalis (Htibner), commonly referred to as the European corn borer, from the genus Pyrausta Schrank to Ostrinia Hiibner was restricted to species occurring in Europe. Because of that fact, treatment of its contemporary Amer- ican congeners, Pyrausta ainsliei and Pyrausta penitalis (Grote), was omitted. P. ainslier is placed in synonymy under obliteralis herein, but penitalis is not a Loxostege or Phlyctaenodes species and has never been assigned to either of them; in a strict sense it is not an essential part of this study. Due to its association with ainsliet and nubilalis, however, there is considerable economic literature involved. P. penitalis is also hereby assigned to Ostrinia for its correct placement and uniformity of nomenclature in references to obliteralis, ainsliei, and penitalis. Phlyctaenodes parvipicta (Barnes and McDunnough), new combination Figure 170 Loxostege parvipicta Barnes and McDunnough, 1918, Contr. Nat. Hist. Lepid. North America, vol. 4, p. 161—McDunnough, 1939, Southern California Acad. Sci. Mem., vol. 2, no. 1, p. 13. Mats (fig. 170).— Alar expanse 13 mm. Frons conical. Antenna ciliate, cilia longer than width of shaft. Midtibia not incrassate; hair-pencil absent. Upper surface of forewing ochreous, markings pinkish. Hindwing white, without markings. Genitalia like those of pustulalis but with uncus and vinculum narrower, gnathos stouter, and aedeagus longer and more slender. Frema.e.—Alar expanse 14mm. Similar to male in color and macu- lation. Genitalia like those of pustulalis but with anterior and posterior apophyses shorter and collarlike structure of ductus bursae more strongly sclerotized. Lrectotypr.— Male, U.S. National Museum, USNM 67627, geni- talia slide HWC 7815. Typxr-LocaLtiry.—Olancha, Inyo Co., Calif. Foop pLant.— Unknown. NO. 3561 LOXOSTEGE—CAPPS 59 DistrisuTion.— Arizona: Mojave Co. California: type-locality. Nevada: Clarke Co. SPECIMENS EXAMINED.— 17. In FiuicHT.— March to June. Remarks.—Of the American species studied, only parvipicta has the characters of Phlyctaenodes, to which it is hereby transferred; it is not closely related to the vibicalis group as indicated in the original description. The species was based on three males and two females. Two of the specimens were labeled as type male and type female and the remainder paratypes. The specimen labeled ‘“‘Lozostege parvipicta B. & McD., male type” is hereby designated lectotype of the species. References GUENEE, M. A. 1864. Deltoides et Pyralites. Vol 8 in Boisduval and Guenée, Histoire naturelle des insectes: Species général des lépidoptéres, pp. 173, 405. Hampson, G. F. 1899. A revision of moths of the subfamily Pyraustinae and Pyralidae. Proc. Zool. Soc. London, pp. 207-212. 1918. Descriptions of new pyralide of the subfamily Pyraustinae. Ann. Mag. Nat. Hist., ser. 9, vol. 2, p. 189. Hisner, J. [Huebner] 1816-27. Verzeichniss bekannter Schmettlinge [sic], p. 352. Marion, H. 1957. Complément a la classification et nomenclature de Pyraustinae d’Europe. Entomologiste, vol. 13, p. 129. Ficures 1-7.—Loxostege rantalis (Guenée): 1, larval setal arrangement of prothorax, meso- thorax, and abdominal segments 1, 3-6, 8, 9, using Dyar and Heinrich system of termi- nology (TI=prothorax, TI]=mesothorax, Al=Ist abdominal segment, A3-6=3rd to 6th abdominal segments, A8=8th abdominal segment, A9=9th abdominal segment). Loxostege aeruginalis (Hubner): 2, forewing; 2a, hindwing. Phlyctaenodes pustulalis (Hubner): 3, forewing; 3a, hindwing; 4, female genitalia; 5, male genitalia; 5a, aedeagus Loxostege aeruginalis (Hubner): 6, male genitalia; 6a, aedeagus; 7, female genitalia (Ap= accessory pouch, Bc=bursa copulatrix, Db=ductus bursae, Ds=ductus seminalis, Os=ostium, Sn=signum). NO. 3561 LOXOSTEGE—CAPPS 61 Ficures 8-13.—Loxostege, male genitalia: 8, 8a, helvialis (Walker); 9, 9a, pseudohelvialis, new species; 10, 10a, nayaritensis, new species; 11, lla, arizonensis, new species; 12, 12a, neohelvialis, new species; 13, 13a, sinaloensis, new species. 62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Ficures 14-19.—-Loxostege, male genitalia: 14, 14a, polingi, new species; 15, 15a, pelotasalis, new species; 16, 16a, boliviensis, new species; 17, 17a, coloradensis (Grote and Robinson); 18, 18a, fordi, new species; 19, 19a, alpinensis, new species. (Ae=aedeagus, An=anellus, Cl=clasper, Dms=dorsal margin of sacculus, Gn=gnathos, Hp=harpe, Ms=modified setae, Obr=oblique ridge, Pd=pad, Ts=transtilla, Un=uncus, Vn=vinculum.) No. 3661 LOXOSTEGE—CAPPS 63 Ficures 20-25.— Loxostege, male genitalia: 20, 20a, marculenta (Grote and Robinson); 21, 2la, neomarculenta, new species; 22, 22a, potosiensis, new species; 23, 23a, conisphora (Hampson); 24, 24a, neotropicalis, new species; 25, 25a, pseudobliteralis, new species. 64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Ficures 26-31.—Loxostege, male genitalia: 26, 26a, neobliteralis, new species; 27, 27a, mancalis (Lederer); 28, 28a, ramsdenalis (Schaus); 29, 29a, entephrialis (Schaus); 30, 30a, pergilvalis (Hulst); 31, 31a, sacculalis Amsel. NO. 3561 LOXOSTEGE—CAPPS 65 Ficures 32-37.—Loxostege, male genitalia: 32, 32a, nigripes (Schaus); 33, 33a, jaralis (Schaus); 34, 34a, chiapasalis, new species; 35, 35a, cayugalis, new species; 36, 36a, cynoalis (Druce); 37, 37a, ecuadoralis, new species. 221-525— 67 5 66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Ficures 38-43.—Loxostege, male genitalia: 38, 38a, jacalensis, new species; 39, 39a, cochisen- Sis, new species; 40, 40a, corozalis, new species; 41, 41a, autocratoralis (Dyar); 42, 42a, mellintalis (Druce); 43, 43a, venadialis, new species. NO. 3561 LOXOSTEGE—CAPPS 67 Ficures 44-49.—Loxostege, male genitalia: 44, 44a, conisphoralis, new species; 45, 45a, purulhalis, new species; 46, 46a, huachucalis, new species; 47, 47a, marialis, new species; 48, 48a, yucatanalis, new species; 49, 49a, volcanensis, new species. 221-525—67——6 68 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Ficures 50-55.—Loxostege, Male genitalia: 50, 50a, similalis (Guenée); 51, 51a, rantalis (Guenée); 52, 52a, occidentalis (Packard); 53, 53a, eneanalis (Schaus); 54, 54a, bifidalis (Fabricius); 55, 55a, brasiliensis, new species. NO. 3561 LOXOSTEGE—CAPPS 69 Ficures 56-61.—Loxostege, male genitalia: 56, 56a, protealis (Warren); 57, 57a, piuralis, new species; 58, 58a, labeculalis (Hulst); 59, 59a, federalis, new species; 60, 60a, cupre- tcostalis (Dyar); 61, 61a, subcuprea (Dognin). 70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Ficures 62-69.—Loxostege, female genitalia: 62, helvialis (Walker); 63, pseudohelvialts, new species; 64, arizonensis, new species; 65, nayaritensis, new species; 66, neohelvialis, new species; 67, polingi, new species; 68, nigripes (Schaus); 69, saltensis, new species. NO. 3561 LOXOSTEGE—CAPPS 71 Ficures 70-75.—Loxostege, female genitalia: 70, marculenta (Grote & Robinson); 71, neomarculenta, new species; 72, cynoalis (Druce); 73, conisphora (Hampson) 74, rams- denalis (Schaus); 75 huachucalis, new species. a2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 : | Ficures 76-81.—Loxostege, female genitalia: 76, marialis, new species; 77, coloradensis (Grote & Robinson); 78, jaralis (Schaus); 79, pergilvalis (Hulst); 80, conisphoralis, new species; 81, volcanensis, new species. NO. 3561 LOXOSTEGE——CAPPS Ficures 82-89.—Loxostege, female genitalia: 82, cochisensis, new species; 83, alpinensis, new species; 84, fordi, new species; 85, cayugalis, new species; 86, neotropicalis, new species; 87, neobliteralis, new species; 88, pseudobliteralis, new species; 89, mancalis (Lederer). 74. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Ficures 90-96.—Loxostege, female genitalia: 90, venadialis, new species; 91, mellinialis (Druce); 92, corozalis, new species; 93, autocratoralis (Dyar); 94, rantalis (Guenée); 95, similalis (Guenée); 96, occidentalis (Packard). NO. 2561 LOXOSTEGE—CAPPS 15 Ficures 97-104.—Loxostege, female genitalia: 97, protealis (Warren); 98, bifidalis (Fab- ricius;) 99, brasiliensis, new species; 100, piuralis, new species; 101, eneanalis (Schaus); 102, labeculalis (Hulst); 103, federalis, new species; 104, cupreicostalis (Dyar). PLATES (Figures 105-178) Figures 105—116.—Loxostege, right wings: 105, helvialis (Walker), 9 ; 106, pseudohelvialis, new species, o' paratype; 107, fordi, new species, co paratype; 108, alpinensis, new species, o type; 109, poling, new species, & type; 110, boliviensis, new species, o& type; 111, nechelvialis, new species, o' paratype; 112, arizonensis, new species, o' paratype; 113, pelotasalis, new species, o type; 114, saltensis, new species, type; 115, autocratoralis t (Dyar), 2 type; 116, corozalis, new species, co type. Ficures 117-128.—Loxostege, right wings: 117, nayaritensis, new species, 9 paratyp 118, sacculalis Amsel, @; 119, mancalis (Lederer), @ lectotype; 120, ramsdenalis (Schaus), 2 type; 121, cochisensis, new species, Q paratype; 122, pergilvalis (Hulst), o; 1235 coloradensis (Grote and Robinson), 2 ; 124, huachucalis, new species, o' type; 125, marialis, new species, o' type; 126, purulhalis, new species, o type; 127, conisphoralis, new species, ype; > ) I ype; ) o paratype; 128, conisphora (Hampson), & paralectotype. Figures 129-140.— Loxostege, right wings: 129, volcanensis, new species, o& type; marculenta (Grote and Robinson), o; 131, neomarculenta, new species, 9 paratype; neobliteralis, new species, o' paratype; 133, neotropicalis, new species, o’ type; jacalensis, new species, o type; 135, pseudobliteralis, new species, co’ paratyy cayugalis, new species, 9 paratype; 137, cynoalis (Druce), &@ lectotype; 138, ecwadoralis, new species, co’ type; 139, jaralis (Schaus), 2 type; 140, labeculalis (Hulst), 9. Ficures 141-148.— Loxostege and Ostrinia, right wings: 141, L. intinctalis (Dyar), & type; 142, L. cupreicostalis (Dyar), & type; 143, L. mellinialis (Druce), 2; 144, L. phrixalis (Dyar), 2 type; 145, L. venadialis, new species, 2 paratype; 146, O. obliteralis (Walker), Q type; 147, L. entephrialis (Schaus), & type; 148, L. federalis, new species, o type. Figures 149-160.— Loxostege, right wings: 149, subcostalis (Dyar). & type; 150, sinaloensts, new species, co’ paratype; 151, chiapasalis, new species, & type; 152, nigripes (Schaus), o type; 153, rantalis (Guenée), &; 154, rantalis (Guenée), 9 type; 155, occidentalis (Packard), co; 156, occidentalis (Packard), 9; 157, similalis (Guenée), co type; 158, similalis (Guenée), 9 ; 159, garalis (Schaus), & type; 160, eneanalis (Schaus), & type. Ficures 161-172.—Loxostege and Phlyctaenodes, right wings: 161, L. eneanalis (Schaus), 2; 162, L. protealis (Warren), 9 ; 163, L. stolidalis (Schaus), & type; 164, L. bifidalis (Fabricius), o; 165, L. bifidalis (Fabricius), 2 ; 166, L. brasiliensis, new species, o type; 167, L. brasiliensis, new species, 9 paratype; 168, L. pruralis, new species, o& type; 169, L. piuralis, new species, 2 paraty pe; 170, P. parvipicta (Barnes and McDunnough), @ lectotype; 171, L. aemulalis (Dognin), #7 type; 172, L. mojgavealis, new species, 9 type. Ficures 173-178._—Loxostege, right wings: 173, naranjalis (Schaus), 2 type; 174, subcuprea (Dognin), & type; 175, mojavealis, new species, 9 type; 176, clarissalis (Schaus), o type; 177, fuscivenalis (Schaus) o type; 178, peruensis, new species, o type. US GOVERNMENT PRINTING OFFICE: 1967 Proceedings of the United States National Museum SMITHSONIAN INSTITUTION « WASHINGTON, D.C. Volume 120 1966 Number 3562 NOTES ON FLIES CAPTURED IN TREETOPS IN MALAYA (DIPTERA: EMPIDIDAE, NERITDAE, PLATYSTOMATIDAE, SEPSIDAE, MUSCIDAE) By GrorcE C. STEYSKAL ! An unusually interesting lot of Diptera captured on treetop plat- forms in the Selangor Gombak Forest Reserve, Malaya, by Dr. H. Klhott McClure, of the U.S. Armed Forces Institute of Pathology, was presented to this author for determination. The following notes are the result. All specimens are deposited in the collections of the United States National Museum (USNM). The following abbreviations, referring to leg parts, chaetotaxy, and wing veins, have been used throughout: a=anterior, acr= acrostichal, ad=anterodorsal, av=anteroventral, cr=coxa or coxae (cz, =forecoxa, etc.), d=dorsal, de=dorsocentral, f=femur or femora (fi=forefemur, etc.), fo=fronto-orbital, h=humeral, mspl=meso- pleural, ntpl=notopleural, p=posterior, pa=postalar, pd=postero- dorsal, ppl=propleural, pra=prealar, prs=presutural, prsc=pres- cutellar, pv=posteroventral, sa=supra-alar, sc=scutellar, sl= sublateral, stpl=sternopleural, t=tibia or tibiae (t;,=foretibia, etc.), ta=anterior crossvein, tp=posterior crossvein, v=ventral, vti=inner vertical. 1 Entomology Research Division, Agriculture Research Service, U.S. Depart- ment of Agriculture, Washington, D.C. 1 Ds PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Family Empididae Hybos melanauges, new species FIGURES la-e Mauz.—Length of body 3 mm; wing 3.25 mm. Color black, only basitarsi brownish. Sides of thorax, posterior part of mesonotum, and coxae, with thin white pruinosity; middle of mesonotum shining with very little microsetation; bristles black; hairs of coxae, dorsal side of f, and f3, and abdomen white. Figure 1.—Hybos melanauges, new species: a, antenna; b, right middle tibia and tarsus, anterior view; c, wing; d, postabdomen, right side; ¢, postabdomen, left side. Hybos meracrus, new species: f, wing; g, tip of abdomen; h, right middle tibia and tarsus, an- terior view (x=position of bristle on posterior side); i, antenna. Head with eyes contiguous above, ommatidia of upper half approx- mately twice as large as lower ones; antenna as in figure 1a, arista NO. 3562 MALAYAN FLIES—STEYSKAL 3 bare; proboscis slightly shorter than height of head; palpi a little shorter than proboscis, very slender, with only one rather small apical bristle. Thoracic chaetotaxy: 2 ntpl, 1 pa, 1 de (also a few small anterior dorsocentral hairs), 1 acer, 1 sc; pa, de, acr in transverse line some distance anterad of scutellum. Wing (fig. 1c) hyaline, with poorly developed brown stigma; halter pale yellowish. Legs slender; f; somewhat clavate, at apical third 0.15 as wide as long. Bristles of legs as follows: t; with one short median d and one long preapical ad; tf, as in figure 1b; f; with complete row of 16-17 staggered av, mostly approximately as long as femoral diameter, in apical half with 3-4 pv; ts with one long median ad, followed a short distance apicad by a smaller d, one ad at apical 1/7, and one each ad and pd preapicals; ¢; with basiventral basitarsal bristle and each tarsal segment of all tarsi with more or less distinct preapical ad and pd; basitarsus,; with a secondary basiventral and 2 median pv; basitarsus; slightly enlarged. Abdomen with sparse whitish hairs of moderate length; postabdo- men (fig. 1d—e) scarcely greater in diameter than preabdomen, dorsal valves each with 2 apical processes and rows of black bristly hairs. Hototyrre.—Male, Malaya, Selangor Gombak Forest Reserve, tree platform, February 24, 1962 (H.E. McClure), USNM 67631. This species runs in Frey’s key (1953, pp. 57-71) to H.. nigronitidus Brunetti, described from northern India and recorded by Frey from northeast Burma; it differs therefrom by its lack of “stiff hairs’? on the humerus, narrowly brown knees (except possibly on middle legs), pale hairs on the under side of the tarsi, and ‘wollig weissliche Behaarung”’ [wooly whitish hairs]. Hybos meracrus, new species Figures 1f-7 FErmMaLe.—Length of body 3 mm; wing 3.25 mm. Head, thorax, and abdomen shining black; legs, including coxae, yellowish, all knees narrowly piceous; ¢, dark drown dorsally; /; blackish above in apical 0.15; apical tarsal segments brown. Head with eyes contiguous above, ommatidia of upper half twice as large as lower ones; antenna (fig. 17) black, arista short plumose; proboscis brown, slightly shorter than height of head; palpi nearly as long as proboscis, slender, black, bearing a few small bristles at and near apex. Thorax with pale gray pruinosity on sides and in humeral grooves; mesonotum with scanty grayish pruinosity laterally; mesonotum posteriorly and scutellum and metanotum with somewhat denser 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 yellowish microsetation; middle of mesonotum with very scant and minute microsetation; strong blackish bristles as follows: 1 népl, 1 sa, 1 dc, 1 acr (prescutellar) ; all other thoracic setae hairlike, yellowish. Wings (fig. 1f) pale brownish with deep brown stigma; venation as shown; halter pale yellowish. Legs slender, f; somewhat clavate, at apical fourth 0.12 as wide as long; coxae and most of legs with pale yellowish hairs; f and ¢ with bristles as follows: ¢, with one long median d and one long preapical ad, also with nearly erect fine ventral hairs nearly as long as tibial diameter; ¢, (fig. 1h) including one p of moderate length at position indicated by x; {3 with complete row of 9 av, longest (median) some- what longer than greatest diameter of femur; also 2 ad, one slightly apicad of middle and one at apical %; ¢; with one d shortly apicad of middle and one each a and pd preapicals; tarsi with moderately long hairs, one basiventral bristle on basitarsus, and basitarsus,, each segment of all tarsi with more or less distinct preapical ad and pd; basitarsus, slightly enlarged. Abdomen with sclerites black, lightly grayish pruinose and with sparse whitish hairs of moderate length; terminal segments (fig. 19) with bristles black. Ho.iotype.—Female, Malaya, Selangor Gombak Forest Reserve, tree platform, February 24, 1962 (H. E. McClure), USNM 67632. This species runs in Frey’s key (1953) to H. apicalis Frey, but differs by having j; with less black apically and the upper ommatidia en- larged, or to H. annulipes Frey, from which it differs also in its dif- ferently colored f/f; and the differently bristled ¢. Both of those species are from Burma. If one considers the 2nd wing vein suffi- ciently bowed apically to place this species in Frey’s subgenus Pseu- dosyneches, it would run to H. palawanus Frey, differing therefrom by its hairy arista and by the presence of only 2 bristles on f;. Family Neriidae The following key is offered in the hope that it will be found an improvement over the one by Aczél (1954, p. 511). Key to Asiatic Genera of Subfamily Neriinae 1 (4). Antennal socket largely unpolished, matt or at most with slightly greasy appearance; only 1 ntpl. 2 (3). Third antennal segment short oval with acute tip blending directly into apical, densely white-felted arista; tips of t; and all f of # distinctly clavate: lange species © salasc. aces hack oie Rhoptrum Enderlein Third antennal segment oval to long oval, with rounded tip; tip of t; and all f of @ ordinary, never clavate or swollen; vti usually weakly de- veloped, short and hairlike; smaller species . . Telostylinus Enderlein 4 (1). Antennal socket polished, strongly shining. co — i) 4 NO. 3562 MALAYAN FLIES—STEYSKAL 5 5 (8). Two strong equal fo; anterior nipl strong; 4th wing vein strongly bowed downward before ta, Ist basal cell at widest part therefore nearly twice as wide as length of ta; occiput and prothorax weakly developed. 6 (7). Stump veins on 8rd and 4th veins; anterior de strong; f; of @ with small spinules below... . .. . . . Stypocladius Enderlein 7 (6). Stump veins lacking; antenies a: ae small; f; of o& with very heavy spines on small tubercles. . . . .. . . Nipponerius Coquillett 8 (5). Only one weak fo and sometimes anterior thereto another rudimentary one; anterior ntpl usually very weak; 4th wing vein only gently bowed basad of ta, Ist basal cell therefore only slightly wider than length of ta. 9 (10). Occiput and prothorax greatly lengthened, occiput nearly or quite aslongaseye.... . . . . Gymnonerius Hendel 10 (9). Occiput and prothorax little levethened! occiput much shorter than eye. Paranerius Bigot Neither Aczél nor I have been able to place Protonerius Meijere (1924, p. 29) in our keys. Subfamily Telostylinae This subfamily is distinguished from the Neriinae by the lack of a distinct dorsal expansion of the facial plate at the bases of the antennae. The genus Telostylus Bigot, 1859 (type: T. binotatus Bigot, 1859) is distinguished from the two other known genera by having only 1 nétpl and 1 (prescutellar) de, a long and pointed 3rd antennal segment, and by the basitarsus, of the o sometimes being swollen. Nobody so far has had the temerity to offer a key to Telostylus. The following one is offered as a preliminary contribution since it is constructed largely from descriptions—although TJ. philippinensis Cresson has been placed from examination of the type specimen in the USNM, and 7. niger Bezzi, from material in the USNM collected by Baker and apparently part of the original lot. Aczél’s statement (1955, p. 32) that 7. benotatus Bigot is known only from Borneo is erroneous, since the original description cites Celebes only, and Aczél’s redescription from Borneo specimens must therefore be suspect. Specimens of 7. binotatus that agree well with Bigot’s description and the concept here presented were taken in the Selangor Gombak Reserve by McClure on October 21, 1961 and October 27, 1962. Key to Species of Telostylus Bigot 1 (4). Mesofrons between the eyes dull black all the way across; mesonotum, scutellum, and pleura largely or wholly black; halter yellow. 2 (3). Legs yellow with preapical dark bands; 3rd antennal segment largely yellowish; dull black supra-alar spot present; wing length 3.75 mm; basitarsus, of ¢’ not broadened (Philippine Islands) .. T. niger Bezzi 3 (2). Legs black; 3rd antennal segment black; mesonotum wholly shining black; wing length 5.5 mm; ? @ basitarsus, broadened (type ?, Sumatra andeninailand) ieee. tet eens see eee he apicatus Edwards 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 4 (1). Mesofrons between the eyes more or less yellowish or reddish medially; halter yellow (except in 7’. remipes). 5 (6). Mesonotum and pleura wholly blackish; f; yellow brown, f; missing in type; 3rd antennal segment black, except at base; wing length 5 mm Gihailandyen 2. vs aes . . T. inversus Hennig 6 (5). Mesonotum variegated He Bromma or pieckish stripes or spots or largely yellowish; pleura with considerable yellowish color. 7 (8). Wing span 5.5 lines (=11.5 mm); body length 3.5 mm; halter black; f yellowish with black tips; ¢@ basitarsus,; broadened (Macassar). T. remipes (Walker)? 8 (7). Wings 4-4.5 mm. long; legs yellowish, f with black tips and sometimes dark preapical bands. 9 (12). Mesonotum yellowish without median marks, only intrahumeral and supra-alar black marks; f; with blackish tips; 3rd antennal segment with extreme tip blackish; scutellum yellow; abdomen yellowish or reddish. 10 (11). Pleura with large blackish mark above cz,;; mesonotum with large intrahumeral spot and posterior thereto a small presutural spot (Philippine Islands). ...... .. . TT. philippinensis Cresson 11 (10). Pleura with 3 blackish marks (Taiwan) . . T. decemnotatus Hendel 12 (9). Mesonotum with median dark marking and often other stripes and spots. 13 (14). Mesonotum with median stripe only; f, blackish in apical third, f; of type missing (Ceylon). .... . . .. T. latibrachium Enderlein 14 (13). Mesonotum with dark markings besides median stripe. 15 (16). Black supra-alar spot lacking; f,; dark brownish, base and preapical band yellow; mesonotum yellowish with pair of presutural black spots; wing length 4 mm (Celebes; Malaya; ?Borneo) .. T. binotatus Bigot 16 (15). Black supra-alar spot present. 17 (18). Pleura yellowish in anterior half, dark brown posteriorly; abdomen blackish brown; scutellum wholly black, lateral pair of bristles very small; f; with apicodorsal blackish mark (islands off Sumatra). T. babiensis Meijere 18 (17). Pleura wholly yellowish or with 1-3 black spots; scutellum more or less reddish medially; 3rd antennal segment black. 19 (20). Pleura with 3 distinct black spots; f without preapical dark marks; ord antennal segment black in apical half; abdomen with blackish central stripe (Philippine Islands). . . T. maccus Osten-Sacken 20 (19). Pleura wholly yellowish or with one indistinct blackish mark between ca, and cx; at least fg with preapical dark band; abdomen blackish brown (Krakatoa; Java; so. India)... T. trilineatus Meijere Family Platystomatidae Plagiostenopterina Hendel Subgenus Meringomeria Enderlein Meringomeria Enderlein, 1924, Mitteil. Zool. Mus. Berlin, vol. 11, p. 108.— Bezzi, 1928, Dipt. Fiji Is. (Brit. Mus.), p. 98.—Hennig, 1941, Arb. Morphol. Taxon. 2 These are the only differentiating characters I can glean from Walker’s short description as Coenurgia remipes (1859, p. 164). NO. 3562 MALAYAN FLIES—STEYSKAL Pi Entom. Berlin-Dahlem, vol. 8, p. 21.—Frey, 1964, Notulae Ent., vol. 44, pod. Stenopterella Malloch, 1931, Proc. U.S. Nat. Mus., vol. 78, no. 15, pp. 13, 16. Enderlein designated Plagiostenopterina trivittata (Walker) as type of Meringomeria, although he also included P. aenea (Wiedemann), which is the type of Plagiostenopterina. Malloch cited P. trivittata also as the type of Stenopterella, distinguishing it from other segregates of Plagiostenopterina in a key and included therewith also P. marginata (Wulp) and a new species, P. trivittigera Malloch. Hennig and Frey have indicated the synonymy of Stenopterella. The nature of the “spiracles” of the 4th and 5th tergites of the female, well within the sclerotic margins, would seem doubtful, but a macerated specimen of P. trivittata does indeed show large ramose tracheae connected with them. I prefer to maintain Meringomeria as a subgenus and offer the following key to include all of the species that can be brought into it at this time, although information on the position of spiracles of the ultimate preabdominal segments is not available for P. newrostigma Bezzi, P. pallidipes Frey, and P. olivacea Hendel. Key to Species of Plagiostenopterina Subgenus Meringomeria 1 (2). Wing with cloud about tp and isolated spot anterad thereof dark brown; costal cell hyaline. . . .... 2.2... .. P. neurostigma Bezzi 2 (1). Wing without such markings. 3 (8). Mesonotum lightly clayey-yellow pruinose without shining longi- tudinal vittae. 4 (5). Cx and f dark brown; 9: spiracles of 5th tergite far mesad of lateral margins, but somewhat closer thereto than to each other and near posterior margin. . . 1 es oy OP. marginata’: (Wulp) 5 (4). Femora nearly wholly gelicws 6 (7). Mesonotum with distinct blackish de lines interrupted at suture; abdomen wholly greenish. . . . ... =... P. olivacea Hendel 7 (6). Mesonotum without such lines, with only faint traces of brownish dc lines; abdomen with yellowish transverse bands; 9: spiracles of 5th tergite close to lateral margins. . . . P. leytensis, new species 8 (3). Mesonotum with pair of longitudinal shining stripes. 9 (10). Pruinosity of mesonotum whitish, rather light; cr and f blackish; 9: spiracles of 5th tergite distant from lateral margins, twice as far from each other as from posterior margin. . . P. planidorsum (Walker) 10 (9). Pruinosity of mesonotum yellowish, heavy; at least f, largely yellowish. 11 (14). Costal cell hyaline. 12 (13). Base of abdomen wholly greenish; costal wing margin from stigma to end of 4th vein with continuous brown seam; 9: spiracles of 5th PETOITEL es) hal ots . .. .. P. pallidipes Frey 13 (12). Base of abdomen seiewiatns wings swith apical spot isolated from dark brown stigma by hyaline space; ?: spiracles of 5th tergite only their diameter from each other and from posterior margin. P. hebes Hendel 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 14 (11). Entire costal margin of wing from root to end of 4th vein with brown seam; abdomen wholly greenish. 15 (16). Shining mesonotal vittae gradually tapered posteriorly; abdomen with dense, yellow pruinose anterior tergal margins; 9: spiracles of 5th tergite near anterior margin, only twice their diameter from each other, surface of tergite dull. . .... . P. trivittata (Walker) 16 (15). Shining mesonotal vittae tapered rather suddenly at posterior end; pruinose tergal margins of abdomen faint; 9: spiracles of 5th tergite near posterior corners, widely separated, surface of tergite shining. P. trivittigera Malloch Plagiostenopterina (Meringomeria) hebes Hendel Plagiostenopterina hebes Hendel, 1914, Abhandl. Zool.-Bot. Ges. Wien, vol. 8, p. 70. The type and only described specimen is a male from Singapore. McClure captured a female on the Selangor Gombak tree platform, on May 7, 1961, which agrees very well with Hendel’s description. Spiracles of 5th tergite as in preceding key, those of 4th tergite near posterior corners; front dull blackish with very narrow shining median longitudinal stripe and many black hairs; abdomen shining, lightly transversely carinulate, and with black hairs on all dark parts, whitish hairs at the yellowish base. Plagiostenopterina (Meringomeria) leytensis, new species Frmaue.—Length of wing 6.5 mm. Head brownish yellow; antennal grooves and pair of ill-defined occipital spots piceous; 3rd antennal segment, palpus, and front brown. Hairs of front whitish to yellowish. Medifrons wholly dull. Thorax largely brownish; middle of mesonotum, mesopleuron, and sternopleuron metallic greenish; nearly entire thorax covered with moderately dense whitish tomentum or coarse pruinescence, that on mesonotum uniform and not showing shining stripes, but with a trace of brownish dorsocentral lines; all hairs whitish to yellowish, except a few black ones on anterodorsal corner of mesopleuron and adjacent humerus; all bristles black. Legs with all cx and f yellowish, only f; with small black antero- apical spot; ¢ black, only basitarsus; brown. Bristles of cz, d and v rows of f;, and 6-8 bristles and several hairs on dorsal third of f, black; all other vestiture of f (hairs) whitish. Wings light yellowish hyaline, entire costal margin from root to tip of 4th vein with dark brown seam, broadest near tip of wing. Basal 4/5 of 2nd basal cell and basal half, except narrow strip along anterior edge, of anal cell bare. Squamae and their fringes whitish. Halter blackish. Abdomen metallic greenish black; compound 1st dorsal segment with transverse pruinose yellowish band in 8rd quarter; basal 2/5 NO. 3562 MALAYAN FLIES—STEYSKAL g of next 2 segments and diffuse basal part of last (5th) preabdominal segment yellowish; entire 5th tergite pruinose; yellowish parts and entire 5th tergite covered with whitish pile; apical part of basal tergite and dark apical parts of 3rd and 4th tergites covered with moderately dense and long, recumbent rufous hairs; spiracles of 4th tergite narrowly within apical corners of tergite; those of 5th tergite narrowly within lateral margins of tergite opposite base of ovipositor; basal segment of ovipositor piceous, with many erect whitish hairs. Houotryrr.—Female, Tacloban, Leyte, Philippine Is. (Baker), USNM 67978. This species, as shown in the preceding key, is apparently most closely related to Plagiostenopterina marginata (Wulp) and to P. olivacea Hendel, should the latter be found definitely to belong to Meringomeria. Plagiostenopterina (Meringomeria) planidorsum (Walker) Charax planidorsum Walker, 1860, Trans. Ent. Soc. London (new series), vol. 5, p. 325.—Hendel, 1914, Genera Insectorum, fase. 157, p. 50; 1914, Abhandl. Zool.-Bot. Ges. Wien, vol. 8, p. 53. Plagiostenopterina (M.) planidorsum (Walker).—Frey, 1930, Notulae Ent., vol. LOM peo One pair, Selangor Gombak Forest Reserve, tree platform, 9 Nov. 10, 1962, @ Jan. 19, 1963. Walker described a female from Burma, Hendel expressed an opinion that the species might be related to P. trivittata (Walker), and Frey made the combination and recorded the species from Palawan. The Malayan specimens agree well with Walker’s description and are indeed close to P. trivittata (see key, p. 7). Plagiostenopterina (Meringomeria) trivittigera Malloch Plagiostenopterina (Stenopterella) trivittigera Malloch, 1981, Proc. U.S. Nat. Mus., Vol S,.10. 15, p.. 17. Selangor Gombak Forest Reserve, tree platform, 2 7,19, May 7, 1961; 2 9, May 13, 1961; 19, Sept. 29, 1962. These are apparently the second recorded captures. The specimens agree well with the type from Singapore in the USNM. Family Sepsidae Dicranosepsis coryphea, new species Figures 2a-f Maure.—Length of body 3.4 mm; wing 2.5 mm. General color metallic purplish black; propleura reddish brown; labellae and most of legs yellowish; f, and f,; becoming dark brown apically; base and apex of ¢; brown; apical swollen part of & black; 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 apical 2 or 3 segments of all tarsi brown; basimesal spot on 3rd antennal segment reddish brown. Head (fig. 2e): front shining, at antennae 0.66 as broad as at level of anterior ocellus; cheeks very narrow, without outstanding posterior bristle; 3 moderately developed vibrissae. Thorax with mesonotum very lightly brownish pruinose; with the following white pruinose areas: anterior part of humeral grooves, Ficure 2.—Dicranosepsis coryphea, new species: a, wing; b, middle leg, posterior view; c, forefemur and tibia, anterior view; d, foreleg, less tarsus, posterior view; ¢, head, profile view; f, male abdomen, apex. lateral stripe on notopleuron, stripe above cz, on propleuron and for only a short distance on sternopleuron, posterior half of sternopleuron, entire hypopleuron (lightly). Chaetotaxy: 1 h, 2 ntpl, 1 sa, 2 de, 2 sc, 1 mspl. Wing pale brownish hyaline, at root and in humeral and larger part of costal cell dark brown; venation as in figure 2a; halter yellowish; squama and its long cilia yellowish. NO. 3562 MALAYAN FLIES—STEYSKAL i Legs (figs. 2b-d): ex, C-shaped, with heavy pilose ventral projection (fig. 2d); f; with strong triangular median projection and without basal hairs or bristles (figs. 2c, d); f: and f; with bristles; f, moderately swollen (fig. 26); ¢; with ventral bristle near tip, latter shining black, dilated into 2 roundish tubercles and with 3 curved, preapically com- pressed and broadened bristles (fig. 2b). Abdomen smooth and shining; segment 8 strongly constricted basally; segment 2 without strong bristles; segment 3 and posterior segments with well-developed bristles, most of which are approximately half as long as apical epandrial pair (fig. 2f). Postabdomen black, processes of epandrium each ending in pair of slender, mesally directed prongs. Houotyrr.—Male, Malaya, Selangor Gombak Forest Reserve, tree platform, Oct. 20, 1962 (H.E. McClure), USNM 67579. This species is evidently close to Dicranosepsis bicolor (Wiedemann), a taxon including a number of entities that Hennig has considered to be members of a “‘Formengruppe.”’ The peculiar modification of t, in D. coryphea has not been mentioned in connection with any of the other forms or related species. The genus Dicranosepsis Duda has been placed by Hennig as a synonym or possible subgenus of the neotropical genus Palaeosepsis Duda, but the relationships and rank of these taxa need further study and at present I prefer to consider them as distinct genera. Family Muscidae Dichaetomyia prolixa (Walker), new combination Aricia proliza Walker, 1864, Journ. Proc. Linn. Soc. London, vol. 7, p. 218. Spilogaster proliza (Walker).—Stein, 1901, Zeits. Hym. Dipt., vol. 4, p. 205. Mydaea proliza (Walker).—Stein, 1918, Ann. Mus. Nat. Hung., vol. 16, p. 185.— Séguy, 1937, Genera insectorum, fasc. 205, p. 289 (in list of “species dubiae vel incerti generis’). One female, Selangor Gombak Forest Reserve, tree platform, Dec. 23, 1962, agrees well with Walker’s description. The species is rather characteristically colored, mesonotum reddish with broad central blackish stripe stopping at yellowish scutellum, abdomen largely yellowish with posterior margins of tergites 3 and 4 rather broadly blackish, and tergite 5 with a pair of well-developed, oval, black lateral spots, legs with all f and ¢, and ¢ yellowish, and all tarsi blackish; 4 postsutural dc bristles. Although clearly a Dichaetomyia, the species is not to be found in Malloch’s keys (1925a, p. 324; 1925b, p. 323) nor in his key of 1929 (p. 401), where it runs to D. rufa (Stein), a species with a wholly reddish mesonotum. 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Genus Fraserella, new name Fraseria Malloch, 1932, Ann. Mag. Nat. Hist., vol. 10, no. 10, p. 309 [type by original designation: F. fulvomaculata Malloch]—Séguy, 1937, Genera insec- torum, fase. 205, p. 3878. [Preoccupied by Fraseria Bonaparte, 1854 (Aves).] Both Malloch and Séguy have referred this genus to the Muscinae, although it is not very conformable therein. The thoracic squama, being in close contact with the base of the scutellum and with straight- ish apical margin, would seem to indicate such a reference. In Séguy’s key (1937, p. 370), the type species traces easily by its hairy eyes; the two additional species I am including at this time, however, have eyes that are scarcely hairy at all and will therefore run to Mesembrina or Muscina. Malloch, in his unpublished key to world genera of Muscidae (now on file in USNM), compares Fraseria to Mesembrina and distinguishes the former by its bare pteropleuron. Malloch, however, also brings out Fraseria in two other places, comparing it in the first instance with Bryantina and Balioglutum (distinguished from those genera by its wholly hairy arista, lack of any seta on wing veins except costa, and 4 postsutural dc) and in the other instance with Gymnodia (distinguished by plumose arista and strong presutural acr). Fraserella resembles Muscina in many respects, but the flattish abdomen and quite differently bristled ¢;, including a distinct calcar, of Muscina separate it from Fraserella. The new species described below is not very similar to F. fulvomaculata, but it fits no better with any other genus than Fraserella and hardly seems distinct enough to require a new genus. I am indebted to Roger W. Crosskey, Commonwealth Institute of Entomology, London, for notes on the type of F. fulvomaculata and for pointing out that Phaonia corbettt Malloch should also be referred to Fraserella. The development of the latter species was described by J. S. Sewill (1931, pp. 233-235) as taking place in the inflorescence of nipa palms and it is possible that the new species has a similar habitat. Fraserella altivolans, new species Figures 3, 4 Ma.e.—Length of wing 8.25-8.5 mm. Color generally black; 1st, 2nd, and base of 3rd antennal segments reddish; oral margin between vibrissae reddish; infraocular bare area brownish; all bristles and hairs black, except very short whitish marginal cilia of thoracic squama. Head proportions (fig. 3c): front at narrowest part 0.115 of total width of head; medifacies strongly narrowed at insertion of vibrissae; arista as in figure 3d, long plumose; anterior ommatidia approximately twice as great in diameter as posterior ones; bristles and hairs similar NO. 3562 MALAYAN FLIES—STEYSKAL 13 to those of female (fig. 3b); occipital ocular margin bare, whitish pruinose. Thorax with changeable pruinosity, when viewed from rear with central stripe, stripes in lines of de bristles, and broad lateral areas, grayish. Chaetotaxy: 4 h in row; 2 ntpl; 1 sl; 1 prs; 1 small pra and 3 strong sa, 3 pa; 2 small posterior ia; 2+-4 dc; 3 strong presu- tural acr and several much smaller and finer pairs of postsutural acr ending in row of 6 strong prsc; 6 marginal sc; 1 ppl and 1 strong stigmatal, each accompanied by a group of long strong bristly hairs, 1+2 stpl, the posterior 2 close together; complete row of about 18 Figure 3.—Fraserella altivolans, new species: a, female head, profile; b, male wing; c, diagrammatic male head, anterior and lateral views; d, male arista. posterior mspl. All thoracic hairs long and coarse, only the follow- ing areas bare: prosternum, center of propleuron, oblique stripe from center of mesopleuron to just posterad of cx,, pteropleuron, hypo- pleuron, supra-alar and postalar declivities (except a few small hairs just below pa), suprasquamal ridge, lower surface of scutellum, me- tanotum. Sides of scutellum with many coarse hairs. Legs simple, only f, quite slender in apical fourth; claws and pul- villi moderate in size. Chaetotaxy: jf, with 2 complete rows of pd, 1 complete row of long pv; t, with short median p and many short ad; f. with 5 pv shorter than femoral diameter, about as many av only half as long as pv, and 3 closely adjacent preapical pd; & with 2 p in apical half, 12d and 4 v preapicals; f,; with complete comb of densely placed disticlinate bristly ad hairs about as long as femoral 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 diameter, complete row of about 20 av (about 5 of which definitely bristles, others hairlike), a series of pv bristly hairs in basal half similar to av series; tf; with 5 median av somewhat shorter than tibial diameter, some bristly pd hairs (a slightly larger one at apical third might be called calcar), distinct apical d and av, other apicals no larger than surrounding bristly setae. T; is distincly compressed on apical half of posterior surface and is furnished with a narrow but deep sulcus in pd middle half. Cz; bare at apex above. Wing (fig. 3a) hyaline; veins brown to black, only swollen base of costa as far as humeral crossvein yellowish; tegula and basicostal scale black; alar squama with blackish margin; thoracic squama closely adjacent to base of scutellum, with nearly straight apical margin, white and with very short whitish marginal cilia; costal vein at swollen base with rather long bristles, otherwise with very short depressed spinules; all other veins bare; halter with dark brown knob. Abdomen rotund, swollen, covered dorsally with yellow pruinosity in somewhat changeable pattern, giving it a finely peppered appear- ance, except on basal 2/3 of 1st apparent tergite. Whole surface of abdomen covered with fairly dense decumbent hairs, marginal seg- mental hairs somewhat longer and more bristle-like; 1st ventral segment hairy. Postabdomen as in figures 4a-c; 5th sternite with broad, V-shaped emargination and 2 digitate apical protuberances (fig. 46). FrEmMaLe.—Wing length 7.9-8.6 mm. Similar to male, except as follows. Head (fig. 36): front slightly broader anteriorly, 0.38 of total width of head; anterior ommatidia 1.25 times as large as posterior ones. Legs with f; bearing complete rows of long pd and pv and small group of medial a; ¢, with 1-2 pd in apical half, nearly complete row of 9-10 rather short ad, long apical ad, pd, and pv; t; lacking the regular palisade of ad of male, but with a number of irregular bristles, of which 2 near middle stand out; t; also with pd bristles a little coarser than in male, but calcar no more distinct; posterior compression and sulcus as In male. Abdomen in general appearance much as in male; postabdomen as in figure 4d (ovipositor), 8th sternite consisting of 2 elongate cuneiform sclerites; spermathecae not seen. Hototypse.—Male, Malaya, Selangor Gombak Forest Reserve, tree platform, Dec. 9, 1961 (H. E. McClure); allotype, same data, Dec. 17, 1961; paratypes, same data, 1 o, Nov. 10, 1962, bait trap 2000’; 1 o', May 20, 1961; 1 9, Mar. 3, 1961; 1 9, Apr. 1, 1961; 19, June 3, 1961; 1 9, June 10, 1961; 1 9, Oct. 21, 1961; 2 9, Sept. 1, 1962; USNM 67824. NO. 3562 MALAYAN FLIES—STEYSKAL 15 The relationships of this species may be gathered from the discus- sion preceding the description and from the following key. Giff x UA fis # SANS LN ee } ams \ f rate Ficure 4.—Fraserella altivolans, new species: a, male postabdomen, lateral and posterior views; b, last preabdominal sternite; c, inner copulatory apparatus; d, female postabdomen (ovipositor). (Views: A=anterior, D=dorsal L=lateral, V=ventral, z=ejaculatory apodeme.) 16 PROCEEDINGS OF THE NATIONAL MUSEUM Vou. 120 Key to Species of Fraserella 1 (2). Eyes hairy; legs fulvous or orange yellow; 4th wing vein strongly bent forward apically; 4 sc. F. fulvomaculata (Malloch), new combination 2 (1). Eyes practically bare; legs black; 4th wing vein very weakly bent forward apically. 3 (4). 3 sc (plus bristle-like posterior discal hairs); antenna black; abdomen gray pruinose. ... . F. corbetti (Malloch),? new combination 4 (3). 5-6 sc; antenna basally reddish; abdomen yellowish-pruinose. F. altivolans, new species References Aczféi, M. 1954. Nertidae von Neuguinea (Diptera). Treubia, vol. 22, pp. 505-531, pl. it. 1955. Nerdiiae von Indonesien (Dipt. Acalyptratae). Treubia, vol. 23, pp. 19-40. Bicot, J. 1859. Dipterorum aliquot nova genera. Rev. Mag. Zool., vol. 7, pp. 305- 315, pl. 11. Frey, R. 1953. Studien tiber ostasiatische Dipteren, 2. Notulae Ent., vol. 33, pp. 57-71. Mautocg, J. R. 1925a. Notes on Australian Diptera with descriptions of thirteen new spe- cies. Australian Zool. vol. 3, pp. 322-338. 1925b. The Anthomyiid genus Dichaetomyia Malloch (Diptera) in the Phil- ippines. Philippine Journ. Sci., vol. 26, pp. 312-332. 1929. Fauna Buruana: Diptera, Fam. Muscidae. Treubia, vol. 7, pp. 390-408. 1931. Exotic Muscaridae (Diptera), 31. Ann. Mag. Nat. Hist., ser. 10, vol. 7, pp. 185-200. MEIJERE, J. C. H. DE 1924. Studien iiber siidostasiatische Dipteren, 15: Dritter Beitrag zur Kenntnis der sumatranischen Dipteren. Tijdschr. Ent., vol. 67, suppl., pp. 1-64. Sievy, E. 1937. Diptera Fam. Muscidae. Genera insectorum, fase. 205, pp. 1-603, pls. 1-9. SEwILu, J. W. 1931. Life cycle of a fly pest (Anthomyiid), on Nipa fruticans. Malayan Agric. Journ., vol. 19, pp. 233-235. WALKER, F. 1859. Catalogue of Dipterous insects collected at Makassar in Celebes, by Mr. A. R. Wallace, with descriptions of new species. Journ. Proc. Linn. Soc. London, vol. 4, pp. 90-172. 3 Described as Phaonia corbetti Malloch (1931, p. 187). Proceedings of the United States National Museum SMITHSONIAN INSTITUTION +« WASHINGTON, D.C. Volume 120 1967 Number 356 EUPHILOMEDES AROSTRATA, A NEW MYODOCOPID OSTRACOD FROM MALDIVE ISLANDS, INDIAN OCEAN By Louis S. KornickER Associate Curator, Division of Crustacea While participating in the U.S. Program in Biology, International Indian Ocean Expedition, aboard the Stanford University Research Vessel Te Vega, the author obtained marine Ostracoda from the waters along the southwest coast of Ceylon and from among atolls forming the Maldive Islands. ‘This paper describes a new species collected in the Maldives. Field collecting was sponsored by the National Science Foundation through the Woods Hole Oceanographic Institution. I wish to thank Miss Caroline Bartlett and William Risley for preparing the illus- trations. Through the courtesy of Dr. Gerhard Pretzmann of the Natural History Museum in Berlin, I was able to examine for com- parative purposes specimens of Philomedes polae Graf from the Gulf of Suez. Subfamily Philomedinae G. W. Miiller, 1912 Genus Euphilomedes Poulsen, 1962 Philomedes (part), Miller, 1912. Philomedes (part) Skogsberg, 1920. Euphilomedes Poulsen, 1962. Type-species: Huphilomedes nodosa Poulsen, 1962, by present designation. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 GENERIC CHARACTERS.—Only a few additions need to be added to Poulsen’s diagnosis of the genus (1962, pp. 359-361) to include the new species described herein. Shell hingement of the genus was not discussed by Poulsen. Fu- philomedes arostrata bears a tripartite hinge with strongly sclerotized posterior elements. Whether this character is present in all members | of the genus is not known. Muscle scars consist of numerous oval or polygonal individual muscle scars located near, or anterior to, the: middle of each valve. Second antenna: Basal spines are not always present on exopodite. They are not present on EH. arostrata and were also reported to be absent on Huphilomedes oblonga (Juday, 1907, p. 145) and EHuphilo- medes multichelata (Kornicker, 1959, p. 230). Sixth limb: The end joint of FE. arostrata is unusual because it is not produced posteriorly. Frontal organ: Not noticeably widened at middle and with some surface spines near tip on appendage of male of FE. arostrata. Surface spines also were reported on Huphilomedes polae (Graf, 1931, p. 37, fie): OccuRRENCE.—Species of Huwphilomedes have been reported from subtropical, tropical, and warm temperate regions (Poulsen, 1962, p. 359). The new species is the second reported from the Indian Ocean. Euphilomedes debilis (Brady, 1902) was previously reported from Trincomali on the northeast coast of Ceylon facing the Bay of Bengal. Euphilomedes arostrata, new species Figures 1-10 Hototyrr.—Gravid female described in this paper, USNM 112658 (figs. 1-4). ParatypE.—Male described in this paper, USNM 112659 (figs. 5-10). TyprE LocaLity.—A male and female were collected about 100 meters apart between Imma and Hura Islands, Male Atoll, Maldive Islands on March 20, 1964. Specimens were in sand obtained from upper surface of bottom in a patch of marine grass in one-third to one-half meters of water. DiaGnosis.—Euphilomedes with four primary claws and seven to nine (possibly more) secondary claws. Second joint of exopodite of | sixth limb not produced posteriorly. Hinge of shell terminating posteriorly in sclerotized angular process on each valve. Shell sur- | face with irregular polygons and without lateral or posterior processes. — DESCRIPTION OF FEMALE.—Shell (figs. 1, 2): Oval in lateral view with greatest height near middle (fig. 14); prominent rostrum; broad rostral incisure, with compressed posterior edge (fig. 1c). Anterior NO. 3563 EUPHILOMEDES AROSTRATA—KORNICKER 3 margin of rostrum and anteroventral margin of shell with scalloped outline formed by crescent-like marginal denticulations. Shell sur- face with irregular polygons and normal pore canals; polygons con- taining pore canals smaller than average; about 23 polygons larger than average, hyaline, slightly in front of valve middle, representing locations of muscle attachments; hyaline, large, oval spot, with polygon border lies dorsal to hyaline polygons (fig. 2F). Anterior two-thirds of dorsal margin of valves hinged by transparent ligament following arcuate curvature of shell outline; posterior one-third with tripartite straight hinge. Hinge not affecting arcuate shell outline, but clearly visible in dorsal view and in lateral view when light is passed through closed shell. Posterior elements on each valve an angular sclerotized process (figs. 24—E); left valve process containing a small forward- facing recess which apparently receives backward-projecting process of right valve (figs. 2B, c). Thin flap behind posterior process of left valve projecting medially, apparently resting on smooth, slightly depressed surface behind posterior process of right valve when shell is closed; upper surface of flap containing polygons similar to those on lateral surface of valves. Anterior hinge element less prominent than posterior; no tooth-and-socket arrangement observed. Medial hinge element of each valve consisting of straight bar without teeth. Sur- face of each valve above medial hinge element bearing two rows of minute tubercles or crenulations. Left valve broadly overlapping right along anterodorsal margin. Thin sclerotized septum projecting downward from dorsal margin of each valve, starting near middle in front of anterior hinge element and terminating just behind posterior hinge element. Septum clearly visible in medial view at anterior and posterior hinge elements where it is set back from contact plane of valves; septum striated at anterior hinge elements. Several septa seem to be present in vicinity of anterior hinge element, especially on left valve. Inner lamella (inner lamella, as used here, is peripheral part of chitin lining of epidermis) broad with vestibule, terminating at each end of dorsal ligament. Line of concrescence lies about halfway between inner and outer margin of inner lamella, closer to inner margin in anteroventral section, closer to outer margin in postero- dorsal section. Marginal zone with simple, fairly closely spaced true and false radial pore canals. Selvage narrow, with wide, corrugated, lamellar prolongation having serrate outer edge and terminal spines (figs. lr, 2c). List midway between selvage and inner margin of inner lamella; about ten striations between list and inner margin of anterior part of inner lamella (figs. 1G, H). Hairs, flaring distally and with concave, cuplike ends abundant, protruding from normal pore canals (figs. 24, H); short, tapered hairs 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 sparsely distributed over shell surface; long, tapered hairs, as well as flared hairs on lateral surface of shell near outer margins, appear to protrude from false radial pore canals. About 11 long, bifurcating hairs bearing secondary hairs form row on inner lamella behind rostrum (figs. 1p, £); a small tapered hair, immediately below rostral incisure, is followed by a space and then a bare hair on left valve, and three bare and two brushlike hairs on right valve; these are followed by a space and then 16 or 17 brushlike hairs, all having bases on or near list (figs. 1a, H); short, bare hairs are distributed between list and inner margin of posterior and posteroventral parts of inner lamella (figs. 2a, B). Dimensions.—Length 1.37 mm., height 0.92 mm. First antenna (fig. 34): First joint elongate with clusters of fine hairs on broad surface. Second joint about three-fourths length of first, with numerous fine hairs; annulated bristle near middle of dorsal margin; slightly shorter bristle near anteroventral corner; short bristle near middle of distal margin. Third joint about one-half length of second; anterodorsal margin with short, bare bristle and long plumose bristle; anteroventral margin with bare bristle about length of third joint; short spines on ventral margin. Fourth joint about twice length of third; anterodorsal margin with two long, slender bristles about length of joint, both bristles with short, fine hairs; anteroventral corner with about four bristles, two long, two short; ventral margin with short spines. Fifth joint slightly longer than third; anterodorsal corner bears slender bristle with short hairs. End joint (or joints) fused, with numerous bristles (ventral bristle may be attached to end of fifth joint). Second antenna (figs. 3B—E): Exopodite: first joint about twice leneth of following joints combined ; second to ninth joints trapezoidal, each joint smaller than preceding, all without basal spines; mediodistal margin of second to eighth joints with comb of slender spines; second to ninth joints with long bristles; bristles on second and third joints without natatory hairs, bristles on fourth to ninth joints with natatory hairs; bristles on second and third joints with short marginal spines, some spines also observed on bristles of fifth to ninth joints. End joint with five or six bristles; two long with natatory hairs, three or four short without hairs. Endopodite: two jointed; first jot with long bristle wreathed by long hairs and five short, bare bristles; second joint bears long bristle with two wreaths of long hairs, minute spine at distal end of joint. Mandible (figs. 3F, H, 1): Coxale endite bifurcate, with rows of slender spines. Basale: ventral margin with seven bristles; two long, distal bristles with wreaths of long hairs; five short, proximal bristles with short hairs. Dorsal margin with three bristles: two at distal corner; NO. 3563 EUPHILOMEDES AROSTRATA—KORNICKER 5 one with long hairs, proximal to middle. Medial surface with six short bristles and spines proximal to middle. Exopodite reaches about middle of first endopodite joint; tip with point; two terminal bristles of about equal length, both with short hairs, but slightly longer hairs on proximal bristle. Endopodite: distoventral corner of first joint with one short, bare bristle and three long bristles with wreaths of long hairs. Second joint: dorsal margin with ten bristles; two long and two short proximal to middle; six at about middle, one of latter much shorter than rest. Ventral margin with five bristles: two bristles distal to middle; three bristles at distal corner, clawlike, with sharp secondary spines. End joint with two claws of about equal length and about three bare bristles. Medial surfaces of basale and second endopodite joint provided with hairs. Maxilla (figs. 3a, 3, K): Dorsal margins of precoxale and coxale with thin, transparent epipodial appendages fringed with long, fine hairs. Coxale with short, bare bristle on distal, dorsal margin. Basale: distal dorsal margin with long, stout bristle with wreaths of long hairs; distal ventral margin with shorter bristle with hairs; narrow, elongate lobe with terminal bristle with base on distal medial margin near posteroventral corner, terminal bristle reaching end of endopodite; base of exopodite on distal lateral margin. Exopodite bears three bristles: proximal short, bare; two terminal bristles extend past end of endopodite, one with long, other with short hairs distally. Endopodite: first joint elongate, with clusters of stiff hairs along anterior margin and medial surface; distal anterior margin bears long bristle with wreaths of long hairs; distal lateral margin bears three (?) bristles with short hairs. End joint with numerous bristles, all with secondary hairs or spines, some clawlike. Precoxale bears three endites: first with about nine bristles; second about seven; third about nine plus one short proximal bristle. Fifth limb (figs. 4a—c): First exopodite joint: middle, distal, anterior margin with two bristles, both bear long hairs; distal, medial corner with short, bare bristle; triangular process in front of main tooth extends almost to tip of tooth. Main tooth composed of four con- stituent teeth: distal tooth, large, bifurcate; following tooth slender, with four secondary teeth; next tooth slender, with about nine sec- ondary teeth; fourth tooth small, without secondary teeth. Bristle with hairs, proximal to fourth tooth. Second exopodite joint has large curved tooth with small marginal teeth on inner curvature, small spine near distal lateral corner. Sixth limb (fig. 4p): Protopodite: first endite with long bristle bear- ing long, stiff hairs and two short bristles with fine hairs; second endite with one proximal and three distal bristles; third endite with eight distal bristles, all except shortest bristle bear stiff hairs, fourth endite 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 with one medial and eight distal bristles; epipodial appendage repre- sented by two short bristles. Second joint of exopodite twice as broad as long; distal margin bears 17 bristles, 4 with long, thin hairs, remainder with one or two wreaths of stiff hairs; medial surface with fine hairs; unlike others in genus, joint not produced posteriorly. Seventh limb (fig. 4&): Cleaning bristles: five in distal group, three dorsally, two ventrally; two (?) proximal bristles; each bristle with three or four bells; distal surface with short, closely spaced hairs. Terminal comb: simple distal tooth with three or four teeth on either side; these teeth have secondary teeth at base. Terminal pegs: two, one or both with short, marginal spines. Furea (figs. 4r, @): Each lamella has 13 claws: primary claws number 1, 2, 7, 12; secondary claws 8, 4, 5, 6, 8,9, 10, 11,13. Primary claws 1 and 2 separated from lamella, 7 and 12 joined to lamella; secondary claws all separated from lamella. Primary claws decrease in length proximally on lamella; claw 12 unusually broad at base. Secondary claws 3, 4, 5, 6 about same length, shorter than claw 7, longer than claw 8; claws 8, 9, 10, 11 decrease in size proximally. Primary claw 12 shorter than secondary claw 11, longer than claw 13. Lamella near basis of claws 1 and 2 bears cluster of long hairs. Claw number 1, right lamella: Posterolateral margin has row of about 24 teeth arranged in sets of two to four; teeth at distal part of claw generally shorter than teeth at midclaw. Central portion of medial surface bears two large teeth followed by about eight smaller teeth. Claw number 1, left lamella: posteromedial margin has row of about 24 teeth similar to row on posterolateral margin of claw number 1 of right lamella; similar row of about 29 teeth on postero- lateral margin. Primary claws 2 and 7 have two rows of teeth, claw 12 has single row. Secondary claws: broad base; spines along anterior and posterior margins become progressively finer and longer on distal half. Eyes: Medial eye with brown pigment in preserved material. Lateral eyes absent. Frontal organ (fig. 44): Two jointed, with suture proximal to middle; construction with faint suture, near proximal end; tip rounded, with- out hairs. Upper lip (figs. 41-x): Small, helmet shaped, with small, unpaired conical process on anterior part; lip and process bear hairs. Irregular protuberances occur between upper lip and frontal organ. Eggs: Specimen contained about 10 oval eggs. DESCRIPTION OF MALE.—Shell (figs. 5-7): Oval in lateral view with greatest beight near middle, more elongate than shell of female (figs. 5A-E). Prominent rostrum and broad rostral incisure similar to female. Rostral anterior margin and shell anteroventral margin NO. 3563 EUPHILOMEDES AROSTRATA—KORNICKER th have scalloped outline formed by crescent-like marginal denticulations. Shell surface with irregular polygons and normal pore canals (fig. 6p). Polygons having pore canals are smaller than average. Distribution of pore canals differs in males and females; males have fewer large canals through which large hairs pass and more small canals containing minute hairs. Large hyaline polygons interpreted as areas of muscle attachment on the female valve, less distinct on male valve. Oval hyaline area with polygon border distinct, located dorsal to hyaline polygons. Hinge elements of male identical to those of female (fig. 7). Inner lamella broad with vestibule as in female, but narrower along ventral margin. Radial canals not observed in male. Line of con- crescence not identified with certainty on male, but may lie just within inner margin of selvage. I{ so, marginal zone of male is much narrower than that of female. Selvage of male similar to that of female. List as well as striations on anteroventral part of inner lamella not observed on male. Distribution of hairs differs on male and female: Hairs that flare distally and have concave tips less abundant than on female, restricted to anterior one-third of valve surface and to ventral margin (fig. 6c); minute tapered hairs apparently more abundant, more or less evenly distributed over valve surface. Long tapered hairs more abundant on posterior one-third, especially along posterior margin; also present along ventral and anterodorsal margins interspersed with flaring hairs. Single tapered hair located behind posterior hinge elements of left and right valves on both male and female. Normal pore canals from which hairs protrude located in polygons smaller than surrounding polygons. Polygons containing flared hairs generally bordered by five polygons; polygons with long tapered hairs by six polygons; polygons containing minute tapered hairs by only four polygons. Pore canals containing minute tapered hairs apparently smaller in diameter than pore canals with long tapered or flared hairs. Ten long hairs on left valve and nine on right form row on inner lamella behind rostrum; not bifurcate like hairs similarly located on female shell (figs. 5r, G). Small bare hair located below rostral sinus in same position as on female. Row of 19 bare hairs on right valve and 21 on left on anterior part of ventral inner lamella; not brushlike as are similarly located hairs on female (figs. 5a, 3). About 32 short, tapered hairs, more or less in two rows, on inner lamella extending from its posteroventral part to posterior hinge element (fig. 5x). Dimensions.—Length 1.00 mm., height 0.60 mm. First antenna (fig. 8a): First joint without bristles; second joint with transverse rows of hairs along ventral margin and three bare bristles; third joint with one bare bristle on ventral margin and two bristles on dorsal margin; of these the shorter is bare, the longer has 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 wreaths of long hairs. Fourth joint weakly separated from third; dorsal] margin with two long bristles, each with wreath of long hairs; distoventral corner with four long, bare bristles. Fifth joint small, inserted ventrally between fourth and sixth joints; sensory bristle with numerous thin filaments on broad base, two or three more distal short filaments and about four short terminal filaments. Sixth joint with one bare bristle distally. End joints bearing five slender bristles and two long, stout c- and f-bristles, each with about 12 short filaments. Broad surfaces of third joint and distal part of second joint with hairs. Second antenna (figs. 8B—G): Exopodite: first joint elongate with small mediodistal spine; second joint about one-fourth length of first; third joint about twice length of second; distal margins of second to eighth joints with comb of short spines. First joint without bristle; bristle of second joint with marginal spines, reaching end of third joint; on left appendage only, second bristle provided with marginal spines on proximal part, natatory hairs on distal one-half (second bristle at this location atypical for family, and probably not present on all males of species). Bristles on joints 3-8 with natatory hairs, without marginal spines. End joint with one short, and three long, stout bristles, all with natatory hairs, and two short, slender bristles, the longer of these having hairs. Endopodite: 3-jointed; first joint with five short, bare bristles and one long, stout bristle having short hairs on distal one-half and wreath of long hairs near middle; second joint elongate with two long, slender bristles on margin near middle; third joint elongate, arcuate, with broadest part proximal to middle; two short, annulated bristles along convex margin, one near distal, other near proximal end; tip of third joint with short, curved nonannulate bristle and four or five serrate transverse ridges. Mandible (figs. 94, B): Coxale endite not observed. Basale: ventral margin with five short, slender, bare bristles and two long, stout bristles with wreaths of long hair; one of the slender bristles farther in from margin than others. Dorsal margin with two bristles at distal corner and one near middle, all bare. Medial surface with about five short bristles near proximoventral corner. Exopodite reaches middle of first endopodite joint; with two subequal bare terminal bristles, proximal bristle with few short hairs; tip of joint with blunt process provided with hairs. Endopodite: distoventral corner of first joint with one short, bare bristle and three long bristles with wreaths of long hairs. Second joint: Dorsal margin with proximal group of two and distal group of six bristles; one short bristle located on medial side between proximal and distal groups. Ventral margin with two bristles proximal to middle and three NO. 3563 EUPHILOMEDES AROSTRATA—KORNICKER g bristles near distal corner. End joint with two claws of about equal length, one short claw, and three bare bristles. Medial surfaces of basale and second endopodite joint provided with hairs. Maxilla (fig. 9c): About two-thirds size of female maxilla. Bristles seem more weakly developed than on female. Dorsal margin of precoxale and coxale with epipodial appendage fringed with long hair. Precoxale with several short bristles along distal margin. Long, stout bristle on dorsodistal margin of female not observed on male. Endopodite first joint elongate, with hairs along anterior margin. End joint with numerous bristles. Precoxale bears three endites provided with numerous bristles. Fifth limb (figs. 9p, @, H): Endites and exopodite joints provided with numerous bristles; first and second exopodite joint with elongate, triangular process; outer lobe of third exopodite joint with two plumose bristles. Sixth limb (fig. 98): Protopodite: first endite with long, bare distal bristle and three short, medial bristles provided with hairs; second endite with one proximal and three distal bristles; third endite with two stout, plumose bristles and about five short, bare bristles; fourth endite with plumose proximal bristle and eight distal bristles; epipodial appendage represented by three short, bare bristles, bristles are followed by small bifurcated process. Second joint of exopodite not produced posteriorly; distal margin bears 15 bristles, all with hairs; medial surface, anterior and posterior margins with fine hairs. Seventh limb (figs. 9F, H, 1): Cleaning bristles: four in distal group, two dorsally, two ventrally; three proximal bristles; each bristle with one to four bells; surface hairs not observed on bristles. Terminal comb contains about nine teeth; some have short, marginal spines. Terminal pegs: two, one or both with short, marginal spines. Diameter of seventh limb about one-half that of female limb. Furca (figs. 10a-£): Each lamella has 11 claws: primary claws 1, 2, 6, 10; secondary claws 38, 4, 5, 7, 8, 9, 11. Medial and lateral row of teeth on all primary claws. Claw number 1 contains 50 to 60 teeth on each side arranged in sets containing 4 to 7 teeth arranged on echelon. Secondary claws have spines along anterior and pos- terior edges. Medial side of lamella near bases of primary claws 1, 2, 6 bears clusters of hair. Copulatory limb (figs. 10, Fr): Long, slender, divided into three lobes having bristles; one lobe has large curved tooth. Eyes (figs. 10c, n): Medial eye with brown pigment in preserved material. Large lateral eyes, each with about 19 ommatophores visible in lateral view; suture divides ommatophores into two parts. 221-527—_67——2 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Frontal organ: 2-jointed with minute spines or hairs at tip. Upper lip: Small, helmet shaped. Irregular protuberances occur between upper lip and frontal organ. Comparisons.—Other species of Huphilomedes having a furca with more than one secondary claw placed between two distally located primary claws and a third primary claw are Euphilomedes multi- chelata (Kornicker) (1959, p. 230), Huphilomedes oblonga (Juday) (1907, p. 145), Huphilomedes polae (Graf) (1931, p. 37). Hach fureal lamella of EH. arostrata has four primary claws, but each lamella of both E. multichelata and E. oblonga has five. Polygons forming the shells of £. arostrata do not have the shingle-like texture of shells of E. multichelata and E. oblonga. The seventh limbs of the male of L. arostrata have three proximal bristles compared to only one on #. multichelata and E. oblonga. Euphilomedes polae is closely related to E. arostrata. The two bristles on the second joint of the endopodite of the male of E. arostrata are longer and are located more proximally on the joint than the bristles on E. polae; the third joint of the endopodite of E. arostrata has one bristle at the distal end, whereas E. polae has two. Graf designated two specimens, both females, captured in plankton of the Red Sea, as Philomedes species. Only the furca was described and illustrated. ‘The distribution of primary and secondary claws on that appendage suggests close relationship with E. arostrata, but lack of knowledge of other appendages prevents further comparison. Specimens of Huphilomedes polae and Philomedes species were found in the same sample, so it is possible that Philomedes species is the female of E. polae. The second exopodite joint of the sixth limb of E. arostrata differs from species of Huphilomedes, Philomedes, Scleroconcha, and Para- philomedes in which that appendage has been described in not being produced posteriorly. Sixth limbs of species of Huphilomedes having a distribution of furcal claws similar to 2. arostrata have not been described. Literature Cited Brapy, G.S. 1902. On new or imperfectly-known Ostracoda, chiefly from a collection in the Zoological Museum, Copenhagen. Trans. Zool. Soc. London, vol. 16, pt. 4, pp. 179-210, pls. 21-25. Graf, H. 1931. Expedition S.M.S. “Pola” in das Rote Meer: Die Cypridinidiae des Roten Meeres. Denkschr. Akad. Wiss. Wein. Math. Naturw. Klasse, vol. 102, pp. 32-46, figs. 1-10. Jupay, C. 1907. Ostracoda of the San Diego region, 2: Littoral forms. Univ. California Publ. Zool., vol. 3, no. 9, pp. 135-150, pls. 18-20. Kornicker, L. 8. 1959. Ecology and taxonomy of recent marine ostracodes in the Bimini area, Great Bahama Bank. Publ. Inst. Marine Science, Univ. Texas, vol. 5 (1958), pp. 194-399, 89 figs. MU.uer, G. W. 1912. Ostracoda. In Das Tierreich. Preuss. Akad. Wiss., pt. 31, 4384 pp., 92 figs. PoutseEn, E. M. 1962. Ostracoda-Myodocopa, Part 1 (Cypridiniformes-Cypridinidae), 414 pp. [Dana-Report No. 57.] SKoasBera, T. 1920. Studies on marine ostracods, part 1 (Cypridinids, Halocyprids, and Polycopids). Zool. Bidr. Uppsala, suppl., vol. 1, 784 pp., 153 figs. i PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 — mh Ficure 1.—Euphilomedes arostrata, new species, female: a, left lateral outline of complete specimen; B, lateral view left valve showing muscle spots and marginal hairs; c, lateral view left valve showing depressed area below antennal sinus; p, medial view rostrum right valve; E, medial view anterior left valve; Fr, detail posteroventral selvage right valve; c, medial view anteroventral part right valve margin; H, medial view anteroventral part left valve margin. (Same scale, in mm.: a, B;c, D, G, H; E, F. Key:i. m.= inner margin of lamella; |. c.=line of concresence; 1.=list; e=edge of valve; s=selvage.) NO. 3563 EUPHILOMEDES AROSTRATA—KORNICKER 1133 Cf CF SRO Og | SD aa ae | ae CARS COS Ficure 2.—Euphilomedes arostrata, new species, female: a, medial view left valve posterior hinge element; B, ventral view posterior hinge elements of partly open connected valves (posterior end of valves to left); c, dorsal view posterior hinge element of partly open con- nected valves (posterior end of valves to left); p, medial right valve posterior hinge ele- ment; E, lateral view left valve posterior hinge element; F, lateral view left valve muscle scars (anterior to left); Gc, medial view part of valve margin with calcium carbonate re- moved by dilute acid; u, lateral view anteroventral part of left valve showing distribution of surface hairs. (Same scale, in mm.: A-F, H; G.) 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Ficure 3.—Euphilomedes arostrata, new species, female: A, lateral view left Ist antenna with frontal organ and medial eye; B, medial view right second antenna; c, 2nd antenna endop- odite; p, section of bristle on 3rd joint second antenna, outer edge on bottom; E, medial view end joints right second antenna; Fr, medial view left mandible exopodite; G, medial view distal end right maxilla; 4, lateral view right mandible; 1, medial view left mandible (marginal bristle of basal not shown); J, lateral view right maxilla (all bristles not shown); K, lateral view left maxilla exopodite, anterior to left. (Same scale, in mm.: A, B; c-G, I, K; H, J. Key: pco=precoxale; CO=coxale; Ba.=basale; Ex.=exopodite; E.=endop- odite; I-III =endite joints.) NO. 3563 EUPHILOMEDES AROSTRATA—KORNICKER 1S) Ficure 4.—Euphilomedes arostrata, new species, female: a, posterior view 4th joint and two lateral bristles of 3rd joint in 5th limb; B, posterior view Ist joint of left 5th limb (looking through part of 2nd joint); c, posterior view 2nd joint of right 5th limb (all bristles not shown); p, medial view left 6th limb; £, distal end 7th limb; F, lateral view right furca; c, detail of where claws 2-13 join lamella of right furca; u, frontal organ and median eye; 1, profile from frontal organ to upper lip; J, profile of upper lip; x, detail of projecting proc- ess on upper lip. (Same scale, in mm.: a-c, E, G, K; D, F, H, J;1. Key: I-IV=endite joints.) 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 J (oe Ficure 5.—Euphilomedes arostrata, new species, male: a, right lateral view of complete specimen showing right lateral eye; B, medial view right valve; c, dorsal view right valve; p, medial view left valve; £, dorsal view of partly open complete specimen; showing lateral eyes; F, anterior part left valve; c, anterior part right valve; H, posterior part right valve; 1, dorsal view posterior part complete specimen; Jj, anterior ventral part left valve. (Same scale, in mm.: A-E, I, F; G, H, J. Key: i. m.=inner margin of lamella.) NO. 3563 EUPHILOMEDES AROSTRATA—KORNICKER lye Ficure 6.—Euphilomedes arostrata, new species, male: a, dorsal view anterior part left valve; B, dorsal view anterior part right valve; c, lateral view left valve with area bearing flared hairs stipled; small oval outlines lucid spot; p, medial view anterior dorsal part right valve showing distribution of hairs and polygons; £, lateral view anterior part left valve showing distribution of hairs; Fr, lateral view posterior part left valve showing distribution of hairs. (Same scale, in mm.: A, B, D, E; C} F.) 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 005 D Ficure 7.—Euphilomedes arostrata, new species, male: a, medial view dorsal hinge left valve with details of anterior and posterior elements; B, medial view dorsal hinge right valve with details of anterior and posterior elements; c, dorsal view posterior hinge element left valve; p, dorsal view anterior hinge element left valve; £, dorsal view posterior hinge element right valve; F, dorsal view anterior hinge element right valve. (Scale in mm.: A; B; C-F.) NO. 3563 EUPHILOMEDES AROSTRATA—KORNICKER RR Ss ie att DY y — a WH ot “al CNH Ficure 8.—Euphilomedes arostrata, new species, male: a, medial view right first antenna; B, lateral view joints 3-7 left second antenna exopodite showing sclerotized areas; c, lateral view joints 1-9 right second antenna exopodite; p, detail showing double bristle on distal corner of joint 2 left 2nd antenna; £, detail showing bristles on joints 8-9 right 2nd antenna; r, 2nd antenna endopodite; c, detail of tip of 2nd antenna endopodite. (Same scale, in mm.: A, C, F; B, D, E, G.) 9 20 PROCEEDINGS OF THE NATIONAL MUSEUM vou. SF Ly mms if i a eh Ficure 9.—Euphilomedes arostrata, new species, male: a, lateral view left mandible; 3, lateral view exopodite and first endopodite joint left mandible; c, maxilla (all bristles not shown); p, distal part 5th limb (all bristles not shown); £, medial view left 6th limb; F, tip of 7th limb; Gc, detail of bristle of 5th limb epipodial appendage; u, 7th limb and epipodial appendage of fifth limb; 1, left (above) and right (below) 7th limbs. scales immmi. A," C, EG) 15 B,D. iks He E= endopodite; I-[V=endite joints.) (Same Key: Pco= precoxale; Co=coxale; Ba=basale; 120 NO. 3563 EUPHILOMEDES AROSTRATA—KORNICKER Al Ficure 10.—Euphilomedes arostrata, new species, male: a, ventral view furca and sclerotized connectives; B, furca and sclerotized connectives viewed from slightly above lateral; c, lateral view left lamella of furca; p, part of distal tooth on left and right lamella of furca showing distribution of secondary teeth and hairs at base; £, lateral view of right lamella of furca and copulatory limb; Fr, distal ends of copulatory limbs; c, lateral view frontal organ and median eye; H, lateral eye. (Same scale, in mm.: A, B, EF; C, G, Hj D, F.) U.S. GOVERNMENT PRINTING OFFICE: 1967 i ; ay f ira i a Mf i iy Pe ‘ \ 7 t oe y vii a Peg re et ie beh ulead ’ ade mele fetes i: Proceedings of the United States National Museum SMITHSONIAN INSTITUTION +« WASHINGTON, D.C. Volume 120 1967 Number 3564 A COMPARISON OF AUSTRALASIAN AND AMERICAN SPECIMENS OF HEMISQUILLA ENSIGERA (OWEN, 1832) (CRUSTACEA: STOMATOPODA) By WituiAM STEPHENSON! Drs. Raymond B. Manning, John C. Yaldwyn, and the present author each decided independently and simultaneously that the status of the Australian specimens of Hemisquilla merited reconsideration. Drs. Manning and Yaldwyn graciously consented to this author’s undertaking the main investigation while at the Allan Hancock Foundation. Here, California material was available mostly from the Hancock Foundation collections. Australian material has been obtained from various museums in Australia (particularly the Aus- tralian Museum, Sydney). Chilean material was limited to the small, complete collection of the Smithsonian Institution; unfor- tunately no recent collections from Chile have been obtained. Dr. John C. Yaldwyn has kindly loaned me the only known New Zealand specimen. Institutions referred to by abbreviations are: Allan Hancock Foun- dation (AHF); Australian Museum, Sydney (AM); Dominion Mu- seum, Wellington (DM); Scripps Institution of Oceanography (SIO) ; United States Nationa) Museum, Smithsonian Institution (USNM). _ Measurements throughout are in millimeters. Carapace lengths were measured with dial calipers, and other dimensions with the 1 Department of Zoology, University of Queensland, St. Lucia, Brisbane, Australia. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 calibrated eyepiece of a stereomicroscope. Both are accurate to ca. 0.1 mm. Two considerations are involved: first, whether the material from Australia and New Zealand (i.e. Australasia) is identical with the American, and secondly, what names to apply to the different popu- lations. The author is deeply grateful to Drs. Raymond B. Manning and John C. Yaldwyn for helpful suggestions, to Miss May Rees for assistance in computations, and to Mr. I. F. Horton for advice on statistical matters. Thanks are also due to the directors and cura- tors of Crustacea of the above-mentioned institutions for allowing me to examine their Hemisquilla collections. I also wish to thank the directors and curators of Crustacea of institutions not mentioned: Johns Hopkins Marine Laboratory, the Queensland Museum, Bris- bane, and the Tasmanian Museum, Hobart. Names of Chilean Material At the generic level the following names have been employed: Gonodactylus Latreille, 1825 (e.g., by H. M. Edwards, 1837; Nicolet, 1849; Miers, 1880; and Bigelow, 1894); Pseudosquilla Dana, 1852 (e.g., by Rathbun, 1910; Kemp, 1913) and Hemisauilla Hansen, 1895 (e.g., by Schmitt, 1940; and Manning, 1963b). Most workers have used the specific epithets styliferus or stylifera, following H. M. Edwards’ (1837) description of Gonodactylus styliferus. Rathbun (1910) showed that G. styliferus H. M. Edwards is a homonym of G. styliferus (Lamarck, 1818), now Pseudosquilla ciliata (Fabricius, 1787). She substituted the specific epithet bigelowi, and Gonodactylus styliferus H. M. Edwards became Pseudosquilla bigelowi Rathbun. This name has been used by a number of American workers for Cali- fornian material (e.g., Hilton, 1915a, 1915b; Buchsbaum and Milne, 1960). Meanwhile Australian records have been under P. stylifera (H. M. Edwards) by Whitelegge (1900) and Kemp (1913); and under H. stylifera (H. M. Edwards) (following Schmitt, 1940) by Stephenson (1953, 1954) and Stephenson and McNeill (1955). Manning (1963b) has shown that Gonodactylus ensiger Owen, 1832, from Chile is clearly the Hemisquilla that H. M. Edwards described as G. styliferus. Manning stressed the color similarities between Owen’s description and preserved Chilean material, which leaves no doubt that Hemisqulla ensigera (Owen, 1832) has priority as the name for the Chilean form. He is also using this name for Australian material (Manning, 1967). NO. 3564 HEMISQUILLA ENSIGERA—STEPHENSON 34 Distribution Randall’s (1839) Hawaiian record, suggesting continuous pan- Pacific distribution has been shown by Manning (1963b) to have been based upon an incorrectly labelled American specimen. Evidently Australasian and American populations are geographically distinct. Past American records show a discontinuous antitropical distri- bution, with distinct Californian and Chilean records, the latter including Juan Fernandez (see Schmitt, 1940). One specimen, col- lected by the Allan Hancock Foundation, links these populations by the following measurements: juvenile, anterior half of body only, carapace length 6.0 mm, rostral length 1.7 mm, rostral breadth 1.7 mm, length eye 3.0 mm, length eyestalk 2.3 mm, length cornea 2.5 mm, breadth cornea 2.3 mm, no mandibular palps visible. Other data are: Jicarita Is., Panama, dredging east side, 24 fms, shelly gravel, coll. W. L. Schmitt, Hancock Galapagos Exped., Velero Sta. 240-34, Feb. 20, 1934, USNM 76381. This specimen is obviously a Hemisquilla as evidenced by (1) the raptorial claw having a single terminal tooth and not being swollen basally, (2) the articulation between the merus and ischium being terminal, (3) the carapace lacking carinae, and (4) the detailed structure of the narrowest part of the cornea (see p. 10) being identical with that of specimens of H. ensigera. Apart from the unlikely event of the first specimen of an undescribed species being a damaged juvenile, it must belong to H. ensigera. Although only 37 specimens have been available for study, it is evident that the species is common in the Californian region. The California Department of Fisheries and Game advise that it is fre- quently caught on rod and line (one was so caught in the author’s presence) and by skin divers. Verbal confirmation has been received from various unofficial sources. The contrast between apparent abundance and numbers available for study from this region suggests the possibility that additional specimens may be recovered from Central American waters in future years. Present records certainly extend the known southern limit of the Californian population, as evidenced by the following specimens all recently obtained from Mexican waters: (1) o&, east side of North Coronados Is., lower Calif., fishing line at 100 ft (833 m), Aug. 20, 1949, coll. John L. Perkins, RI 6.2-6 (SIO). (2) o, Ensenada, lower Calif., Feb. 10, 1958, RI 6.2-11 (SIO). (3) 2%, 5.4 mi E Morro Redondo Pt., Cedros Is., 41 fms (74 m), trawl, Apr. 20, 1951, Velero Sta. 2030-51 (AHF). 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 (4) &, Sebastian Vizcaino Bay, lower Calif., coll. Jack Little- page, acc. no. 153 (AHF). (5) 2 #o’, 3 99, San Cristobal Bay, lower Calif., 41 fms (74 m), trawl, Apr. 27, 1950, Velero Sta. 1949-50 (AHF). (6) o, 2, 1 juvenile, 27°24’ N, 114°40’ W, San Cristobal Bay, lower Calif., 40 fms (72 m), Aug. 20, 1960 (1800-1820 hrs.), otter trawl, 6008B, 123.37, coll. Fred Berry (SIO). (7) &@, Turtle Bay, lower Calif., in purse seine inside bay dur- ing daytime, Stella Maris, coll. Ben Fukuzaki (AHF). (8) 9, 6.25 mi SSW San Hipolito Pt., 36 fms (65 m), trawl, Apr. 29, 1950, Velero Sta. 1952-50 (AHF). The following specimen, kindly loaned by Dr. John C. Yaldwyn, is the first recorded from New Zealand: 9, New Zealand waters, DM reg. no. Z.Cr.1493. Morphological Differences—Initial Study Only four possible distinguishing features were noted on preserved specimens from different areas. In each case there was considerable variability and such overlap between the series that statistical analyses were deemed necessary. Initially, comparisons were made between Australasian and Ameri- can material. Chilean and Californian collections were pooled for the following reasons: (1) the single Panamanian specimen tends to bridge the biogeographic gap; (2) probably the northern population extends in strength beyond the known southern limit as evidenced by the fact that 15 of the 37 ‘‘Californian” specimens have been collected in recent years in Mexican waters; and (3) the small number of Chilean specimens (9 only) available for study. In each case a quantitative study of a feature was made and group means were computed. In most cases deviations from means ex- hibited by each of the individuals in a group were plotted as frequency distribution histograms; in other cases raw data were plotted similarly. These histograms sometimes indicate differences in the constitution of populations from the two areas (figs. 1-3). Histograms showing normal distributions are not given. Variabilities of populations were computed in terms of standard deviations divided by means; these again indicate some population differences. Finally, ¢ tests of the significance of differences between group means were carried out (Lacey, 1953, p. 114; Fisher and Yates, 1957). It is appreciated that the applicability of this test is affected by ab- normal frequency distributions; however, the low p values obtained NO. 3564 HEMISQUILLA ENSIGERA—STEPHENSON 5 in all relevant cases (p<0.001) leave no doubt of the significance of the differences. In certain cases inspection of raw data indicated the possibility of sexual dimorphism. Means for Australasian males and females and for American males and females, respectively, were computed and ¢ tests carried out in “promising” cases within the locality groupings. In other cases it was suspected that the measured ratios depended upon specimen size. Using all specimens from a given area, cor- relation coefficients were computed between ratios and carapace length (i.e., the measure of specimen size) and were tested for sig- nificance (Fisher and Yates, 1957). Regression coefficients were computed in several cases. MANDIBULAR PALP.—Kemp (1913) noted two-segmented palps in an Australian specimen and from one to three segments in Chilean specimens. Schmitt (1940) noted three segments on the right and two on the left side of a single Southern Californian specimen. NUMBER OF INTERMEDIATE TELSON LOBES TTT ll AUSTRALASIAN OUT AUSTRALASIAN —————j AMERICAN B = | AMERICAN Ficure 1.—Frequency distribution histogram showing percentages of specimens with varying numbers of: A, segments of mandibular palp; B, intermediate telson lobes. Figure 14 shows that the Australasian population gives an approx- imately normal distribution curve with a mode of about two segments, but the American is strongly skewed to the right. Table 1 indicates approximately equal variability in the two populations. On the difference between means, ¢ tests give p<0.001. There is no doubt that the specimens belong to populations that are statistically separate. 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Inspection of raw data gave no indications of differences between sizes or sexes of specimens, and analyses were not attempted. A minor cause of variation is related to loss and regrowth of palps. While many of the palps missing from the specimens may have been removed during collection and preservation, the occasional presence of unusually small palps, typically weakly segmented, suggests that natural loss and regrowth can occur. This could reflect varying environmental stresses, apart from any genetical differences. TaBLE 1.—Comparative data of Australasian and American specimens No. of segments of Intermediate telson L/B rostrum mandibular palp lobes Aust. Amer. Aust. Amer. Aust. Amer. Mean 2. 3558 2. 6777 1. 3559 1. 0220 1.1021 1. 3037 n 104 90 118 91 59 44 Range 1-3 1-3 0-3 1-2 0.88-1.26 1.10-1. 54 Standard deviation 0. 473 0. 491 0. 612 0. 147 0. 0821 0.134 SiS sal as 0. 201 0.183 0. 451 0.143 0. 0745 0. 103 mean Probability of differences (t test) (0. 001 ((0. 001 (0. 001 L carapace L/B cornea L eye L rostrum B cornea Aust. Amer. Aust. Amer. Aust. Amer. Mean 4.7746 4.1534 1. 2013 1. 3090 1. 3230 1. 4679 n 59 44 55 41 54 42 Range 4.0-6.0 3.50-5.0 1. 08-1.36 1.04-1.55 1.08-1.46 1.12-1.92 Standard deviation 0. 442 0. 371 0. 0589 0.340 0. 0733 0. 283 Slander demalien 0.0926 0. 0893 0. 0490 0. 260 0. 0554 0.193 mean Probability of differences (t test) (0. 001 (0. 001 ((0. 001 INTERMEDIATE LOBES OF TELSON.—Kemp (1913) noted that an Australian specimen had two lobes on each side but the then known Chilean specimens had single lobes. Schmitt (1940, p. 181) noted single lobes on Southern California specimens and suggested that these differences might be ‘‘of more significance than may seem justified at present.” In counting numbers of lobes, difficulties were encountered over two American and one Australian specimen. These possessed on each side one normal lobe and one very small lobe just lateral to it; the latter were regarded as vestigial and were not counted. In addition, one side of an American specimen had a malformed telson, and this was excluded from the count. NO. 3564 HEMISQUILLA ENSIGERA—STEPHENSON 74 Figure 1B shows that the Australasian population gives a skewed curve but the American an approximately normal curve. (This is the opposite of results shown in figure 14). Table 1 shows that the Australasian data vary more than the American. Differences between means are again significant with p < 0.001, and the populations are again statistically distinct. /\ @ SI B Figure 2.—Dimensions of Hemisquilla ensigera: a, rostrum; B, eye, lateral view; c, cornea, anteroventral view (l1=length, b=breadth, le=length of eye, ls=length of eyestalk; drawn from male, Queensland Mus. reg. no. W1779). RosTRAL PROPORTIONS.— This distinction was suggested initially by comparison of Australian specimens with Kemp’s figure (1913, pl. 7, fig. 84) of a Chilean specimen. All specimens were measured for lengths and breadths of rostrum and carapace lengths. Measuring rostral breadths caused occasional difficulty and basal breadths on the upper surface of the line of articulation with the carapace were measured in preference to maxi- 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 mum breadth (fig. 24). From these measurements, the ratios L/B rostrum and L carapace/L rostrum were computed. Selected data show a hint of bimodality in the Australasian data (fig. 34) and a much less definite hint in the American (fig. 38). The 84 L/B ROSTRUM A-AUSTRALASIAN a NUMBERS b | 1 IL -30 re) 10 20 30 DEVIATION FROM MEAN XIOO 4 L/B ROSTRUM B-AMERICAN 3 wo (on ld m2 => =>} | | | —30 -20 -10 20 30 ie) 10 DEVIATION FROM MEAN XIO0O Figure 3.—Frequency distribution histograms showing numbers of Australasian and American specimens with varying deviations from group mean L/B ratios. small ‘separate’ groups with higher deviations are not distinctive as regards sex ratio or size; to simplify analyses, hints of polymorphism in L/B rostrum data were ignored. There are no comparable hints in L carapace/L rostrum data. NO. 3564 HEMISQUILLA ENSIGERA—STEPHENSON 9 Table 1 shows that there is equivalent variability in Australasian and American material and that the rostra of American specimens are significantly longer than that of the Australasian (p. < 0.001). Inspection of data indicated possible sexual dimorphism and size effects. Relevant data on sexual differences are shown below. L/B rostrum no. mean weighted mean Australasia 60S 1.1017 1.1021 239 9 1.1026 America 280'o' 1.3086 1.3027 169° 1.2925 L carapace/L rostrum no. mean weighted mean Australasia 36d0S 4.7292 4.7746 2399 4.8457 America 28'S 4.1289 4.1534 169° 4.1963 For L carapace/L rostrum data the differences between male and female means is significant almost at the 0.02 level on Australasian material but not significant on American (p ca. 0.3). No tests were carried out on L/B rostrum data because of the small differences between means of Australasian males and females, respectively, and the reversed tendency of Australasian versus Californian data. The effects of size upon the ratios were only investigated upon L/B rostrum data, which appeared to offer the more ‘“ promising” indications. Correlation coefficients between carapace length and L/B rostrum for Australasian and American material are —0.070 and +0.330, respectively. The former is not significant (p >0.1), while the latter is significant at about the 0.03 level. Regression coefficients between carapace length and L/B rostrum are —0.00156 and +0.00479, respectively. Summarizing these analyses, the American and Australasian specimens clearly belong to different populations with shorter rostra on the Australasian material. There are indications of sexual di- morphism, with a smaller L carapace/L rostrum ratio in males, but this is significant only on Australasian material. There are also indications of a slight size effect, which is significant only on American material. Eyrs AND EYESTALKS.—This possible distinction, like the last, arose initially by comparison of Australasian specimens with Kemp’s figure (1913, pl. 7, fig. 84) of a Chilean specimen. Measurements were made on all undamaged eyes of length of eyestalk from lateral view, length of entire eye from lateral view, 221-528 67-2 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 length of cornea from anteroventral view (looking at cornea ‘‘end on’’), and breadth of cornea in similar view (see figs. 3B, C). Choice of the portions of the eye to be measured was conditioned partly by ease of precise recognition on successive occasions. Thus minimal corneal breadth was measured because recognition is aided by slight concave curvatures in the centers of the corneas, by the presence of larger facets in this area, and by transverse pigment lines in pigmented specimens. In all measurements there are subjective errors, and the accuracy of results decreases, possibly to +0.2 mm. From these measurements L/B cornea and L eye/B cornea were computed as the most accurate and meaningful of the possible ratios. Histograms of deviations from means give normal frequency distri- butions in each case. Values of standard deviation/mean (table 1) indicate that for L/B cornea data only the American material varies more than the Australasian. For both L/B cornea and L eye/B cornea data American specimens give higher values than Australasian, with the differences significant at values of p<0.001. Inspection of data indicated possible sexual dimorphism and possible size effects. Relevant data on sexual differences are shown below. L/B cornea no. mean weighted mean Australasia 382010! 1.2028 1.2013 PRONG. 1.1991 American 250'o' 1.3092 1.3090 162 2 1.3088 L eye/B cornea no. mean weighted mean Australasia 3ldéd UPL 1.3230 232 2 1.3143 American 2530'S 1.4772 1.4679 U7 ere 1.4512 For L/B cornea data, differences between the means of males and females are not significant either upon Australian material (p ca. 0.6) or American (p>0.9 level). For L eye/B cornea Australian data significance is at the 0.1 level but American data are not significantly different (p>0.9). Size effects were investigated only on L/B cornea data. The correlation coefficient between carapace length and L/B cornea for Australasian specimens is 0.296 (p ca. 0.02) and for American 0.476 (p<0.001). Regression coefficients are 0.00145 and 0.00562, respec- tively. Summarizing these analyses, the American and Australasian specimens clearly belong to different populations as indicated by the NO. 3564 HEMISQUILLA ENSIGERA—STEPHENSON 11 dimensions of eyes and eyestalks (broader cornea in Australasian material). Of four possibilities of sexual dimorphism, only one is significant—L eye/B cornea in Australasian males, greater than in females. The ratio L/B cornea increases significantly with specimen size. The general summation of the results of the present section is that Australasian and American material are significantly different in all features selected for examination. In the six sets of data, differences between means are significant at p levels of <0.001. Apart from differences in means there are also differences in the form of frequency distributions of deviations from mean (e.g., skewness) and in hetero- geneity within groups. There are indications of sexual dimorphism in certain features and stronger indications of proportions changing with specimen size. Since none of these effects are as noteworthy as those depending upon locality, they were neglected in further analyses. Comparisons Between Californian, Chilean, and Australasian Material Because differences between American and Australasian populations had proved so highly significant and because in many respects mean values for Chilean material lay between Californian and Australasian extremes, data were reconsidered under the three main locality group- ings. Results, given in tables 2 and 3, show that differences between Californian and Australasian data are highly significant throughout. Further consideration is restricted to Australasian/Chilean and Chilean/Californian comparisons. MANDIBULAR PALP.—The mean Chilean value is roughly midway between Australasian and Californian means, with an approximately equal probability of the differences being significant (p ca. 0.1 and 0.2 — 0.1, respectively). The three populations have approximately equivalent variability. The Californian data is more skewed than the Chilean data. INTERMEDIATE LOBES OF TELSON.—The mean Chilean value lies further from the Australasian than the Californian and differs sig- nificantly from the Australasian (p ca. 0.01) but not from the Cali- fornian (p 0.4 — 0.3). The Chilean data resembles the Californian in the form of its frequency distribution; both differ from the skewed curve for Australasian data. Variability in the Australasian data is much higher than in the Chilean, which resembles more that of the Californian. RostRAL PROPORTIONS.—L/B rostrum: The mean Chilean value is closer to the Australasian than to the Californian. It is not sig- nificantly different from the former (p ca. 0.2) but is from the latter (p <0.001). The variability of the Chilean values is of the same 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 order of magnitude as for the Australasian and is considerably smaller than the Californian. L carapace/L rostrum: Again the mean Chilean value is closer to the Australasian than to the Californian. It is just significantly different from the former (p ca. 0.1) but 1s highly significantly different from the latter (p <0.001). Variability is approximately the same throughout. EYES AND EYESTALKS.—L/B cornea: The mean Chilean value is midway between Australasian and Californian values, with differences significant in both cases (p ca. 0.001 and <0.05, respectively). Varia- TABLE 2.—Comparative data of Australasian, Chilean, and Californian specimens No. of segments of Intermediate telson L/B rostrum mandibular palp Aust. Chile Calif. | Aust. Chile Calif. | Aust. Chile Calif. Mean 2.3558 2.5500 2.7083 | 1.3559 1.0 1.0274 | 1.1021 1.1744 1.3357 n 104 18 72 118 18 73 59 9 35 Range 1-3 2-3 1-3 0-3 1 1-2 0.88- 1.10- 1.10- 1. 26 1. 29 1.54 Standard deviation 0.473 0.497 0.484 0. 612 0 0.163 | 0.0821 0. 0678 0.112 Standard deviation 0.201 0.194 0.179 | 0.451 0 0.159 | 0.0745 0.0577 0.0839 mean L carapace L/B cornea Leye L rostrum B cornea Aust. Chile Calif. | Aust. Chile Calif. | Aust. Chile Calif. Mean 4.7746 4.5388 4.0543 | 1.2013 1.2689 1.3203 | 1.3220 1.5056 1.4576 n 59 9 35 55 9 32 54 9 33 Range 4. 0- 4.1- 3.50- 1.08- 1.18- 0.95- 1.08- 1.41- 1.12- 6.0 5.15 4. 57 1.36 1.38 1.55 1. 46 1. 65 1.92 Standard deviation 0.442 0.3376 0.3046} 0.0589 0.0617 0.0931 | 0.0733 0.0782 0.3123 Standard deviation mean 0.0926 0.0744 0.0750 | 0.0490 0.0486 0.0705 | 0.0554 0.0519 0.2140 bilities of Australasian and Chilean material are roughly the same and lower than Californian. L eye/B cornea: The Chilean mean is further from the Austra- lasian than is the Californian; p values emphasize the difference between Chilean and Australasian material (p <0.001) and the simi- larity between Chilean and Californian (p 0.6 — 0.5). The varia- bility of the Californian population is much greater than that of either of the others. Summarizing these results, of the six determinations upon the four selected structural features, in four the mean Chilean values lie between the extremes of the Australasian and the Californian, and in the other two they lie further from the Australasian than do the NO. 3564 HEMISQUILLA ENSIGERA—STEPHENSON 13 Californian. In significance tests in 5/6 cases, differences between Chilean and Australasian means are significant, and in one case (L/B rostrum) they just fail to be significant (p ca. 0.2). In 2/6 cases differences between Chilean and Californian means are not significant, m one case the difference just fails to be significant, and in two cases differences are highly significant. Comparisons Between H. ensigera and H. braziliensis In deciding the status to be given to the three populations of H. ensigera, two considerations are relevant: (1) Distinctions between stomatopod taxa are becoming finer from the generic level downward (Seréne, 1962; Manning, 1963a). (2) H. ensigera is very close to H. braziliensis (Moreira, 1903a, b), the only other species of the genus. The only conspicuous difference in preserved material is that in H. braziliensis the lobes of the telson TABLE 3.—Significance of differences among three populations of H. ensigera No. of segments Inter- L/B L carapace L/B Leye of man- mediate rostrum | J rostrum cornea B cornea dibular | telson lobes palp Australasia t 1. 784 2. 870 1.377 1.651 3. 674 8. 088 v. Chile Pp c. 0.1 ce. 0. O01 c. 0.2 chon ce. 0. 001 «0. 001 Chile v. t 1. 527 0. 932 5. 220 5. 295 2. 032 0. 591 California Dp 0. 2-0. 1 0. 4-0.3 (40. 001 (0. 001 (0. 05 0. 6-0. 5 California t 11. 226 9.176 22. 680 15. 727 13. 538 6. 164 v. Australasia Pp (0. 001 {(0. 001 ((0. 001 ((0. 001 ((0. 001 (0. 001 are spinous (see Schmitt, 1940, figs. 18a, b). There are also possible differences in the form of the basal process of the uropods, in the color of live specimens, and in rostral shape (Moreira, 1905; Lemos de Castro, 1955). Because rostral shape is variable in H. ensigera and because there may be other statistical differences in variable features, most of the measurements upon H. ensigera were repeated upon the nine specimens of H. braziliensis in the USNM. The ranges of values show complete overlap with H. ensigera data, but comparison of mean values gives data as shown below. Number of segments of mandibular palp: 2.8889, cf. 2.7083 for nearest and largest H. ensigera value (Californian), difference highly significant (p <0.001). Intermediate lobes of telson: On the criteria adopted above, the number of lobes is identical with H. ensigera from Chile, being unity throughout; however, most or all specimens of H. braziliensis possess small lobes just lateral to the intermediates, similar to those here 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 regarded as ‘‘vestigial’’ (see p. 6; also Schmitt, 1940, fig. 18b). These were only observed in three specimens of H. ensigera. L/B rostrum: 1.1088, cf. 1.1021 for nearest H. ensigera value (Australasian), difference not significant (p 0.8). L carapace/L rostrum: 3.9650, cf. 4.0543 for nearest and lowest H. ensigera value (Californian), difference not significant (p 0.4-0.3). L/B cornea: 1.0556, cf. 1.2013 for nearest and lowest H. ensigera value (Australasian), difference just not significant (p 0.2—0.1). L eye/B cornea: 1.1544, cf. 1.3220 for nearest and lowest H. ensigera value (Australasian), difference highly significant (p <0.001). Summarizing these differences, H. braziliensis differs from H. ensigera in having (1) in all cases, more spinous telson teeth and (probably) differently shaped basal processes of uropods; and (2) statistically significant differences as follows: a larger number of seg- ments in the mandibular palp, a higher proportion of individuals with “vestigial” intermediate lobes of telson, a relatively broader cornea as indicated by lower L eye/B cornea ratios (and possibly by lower L/B cornea ratios). Discussion There are three populations of Hemisquilla in the Pacific—one Australasian and two American. The latter are geographically sep- arate (apart from a single juvenile from Panama). Four possible morphological differences between the populations have been noted: number of segments in the mandibular palp, number of intermediate lobes on the telson, relative length of rostrum, and relative breadth of cornea. In each case there is considerable vari- ability and overlap between populations. An initial analysis showed that American and Australasian popu- lations are morphologically distinguishable by statistical tests at a highly significant level. (These analyses also show hints of sexual dimorphism in some features and slight alterations of proportions with increasing specimen size.) At this stage one might conclude that the American and Australasian forms belong to two subspecies. In further analyses a three-way comparison has been made, American material being treated under Chilean and Californian headings and compared with the Australasian. Of six features measured (two being different measurements of rostral proportions and two being eye proportions), the numbers of dissimilarities among the three populations are shown below. highly just or not significant doubtfully significant significant Australasian v. Californian 6 0 0 Chilean v. Australasian 3 3 0 Chilean v. Californian 3 1 2 NO. 3564 HEMISQUILLA ENSIGERA—STEPHENSON 15 It is evident on the basis of present data that Chilean material is almost as distinct from the Californian as it is from the Australasian. The best schema to meet this situation is to regard each population as a subspecies. This conclusion is influenced by three factors: (1) Only nine Chilean specimens were available for study. When additional material is measured, it seems likely that differences be- tween Chilean and other material will become more significant. (2) There is the possibility that detailed examination of colors of living specimens may afford further diagnostic criteria. Manning (pers. comm.) stresses the importance of color differences in other genera of stomatopods. In the present case it is possible that Chilean specimens may differ from Californian in having less yellow color on various appendages. Thus Nicolet’s plate (1854, pl. 2, fig. 3) shows the following as green or bluish green: antennal scales, propodus of raptorial claws, and endopodites of uropods. They are canary yellow on living and recently preserved Californian specimens. To determine whether the differences are real or are artifacts, requires observations on fresh Chilean material. (3) Distinctions between stomatopods at all levels are becoming increasingly fine, and subspecific distinctions must be finer still. This is emphasized by a comparison between H. braziliensis and the differ- ent groups of H. ensigera. Apart from two “firm” features the differences are of the same order as those between the different groups of H. ensigera. In these respects the Brazilian form shows no closer relationship to any one group of H. ensigera than to any other. In view of the above, three subspecies of H. ensigera are therefore designated: Chilean: Hemisquilla ensigera ensigera Californian: H/. ensigera californiensis, new subspecies Australasian: HZ. ensigera australiensis, new subspecies The distinguishing features, on the basis of present knowledge are as follows: Mandibular palps: In H.e. australiensis 80% of the palps are two segmented and 20% three segmented. In H.e. californiensis the the values are 25% and ca. 75%, respectively, and in H.e. ensigera they are 45% and 55%, respectively. Intermediate lobes of telson: In He. australiensis about half the telsons have single lobes (45%) and about half (52%) two lobes. In He. californiensis over 95% have single lobes, and in H.e. ensigera all specimens examined have single lobes. On this basis H.e. cali- forniensis and H.e. ensigera are indistinguishable. Length/breadth ratio of rostrum: In H.e. australiensis the ratio is low (mean 1.10), in He. californiensis high (mean 1.34), and in Me. 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 ensigera intermediate (mean 1.17). On this basis H.e. australiensis and H.e. ensigera are doubtfully distinguishable. Length carapace/length rostrum ratio: In H.e. australiensis the ratio is high (mean 4.77), in H.e. californiensis low (mean 4.05), and in H.e. ensigera intermediate (mean 4.54). Length/breadth ratio of cornea: In H.e. australiensis the ratio is low (mean 1.20), in He. californiensis high (mean 1.32), and in H.e. ensigera intermediate (mean 1.27). Length of eye/breadth of cornea: In H.e. australensis the ratio is low (mean 1.32), in He. californiensis intermediate (mean 1.46), and in H.e ensigera high (mean 1.51). On this basis He. californi- ensis and H.e. ensigera are indistinguishable. Type specimens of the three subspecies are designated as follows: H.e. australiensis, male, carapace length, 28.5 mm, 12 miles E. Broken Bay, NSW, Australia, presented by H. Arnold, November 1946, AM reg. no. P.11695. H.e. californiensis male, carapace length, 41.1 mm, Southern California, Anton Dohn coll., gift Venice Mar. Biol. Sta. 1909, USNM 50016. H.e. ensigera Manning (pers. comm.) has informed me that the type of H. ensigera is a dry specimen in the collection of the British Museum (Natural History). I have not had an opportunity to study the specimen. Literature Cited BigELow, R. P. 1894. Report on the Crustacea of the order Stomatopoda collected by the steamer “Albatross” between 1885 and 1891, and on other speci- mens in the U.S. National Museum. Proc. U.S. Nat. Mus., vol. 17, pp. 489-550, figs. 1-28, 3 pls. Bucusspaum R., and Mitng, L. J. 1960. The lower animals: Living invertebrates of the world, 303 pp., 315 figs. Dana, J. D. 1852. Crustacea, 1. Jn vol. 13 of United States Exploring Expedition during the years 1838-1842, under the command of Charles Wilkes, U.S.N., viii+685 pp. Epwarps, H. M. 1837. Histoire naturelle des Crustacés, vol. 2, 531 pp.; Atlas, 28 pls. Fasricius, J. C. 1787. Mantissa insectorum sistens eorum species nuper detectas: Adjectis characteribus genericis, differentiis specificis, emendationibus ob- servationibus, vol. 1, xx+382 pp. Fisoer, R. A., and Yartss, F. 1957. Statistical tables for biological, agricultural and medical research, viii+ 138 pp. NO. 3564 HEMISQUILLA ENSIGERA—STEPHENSON 7 Forest, J.. and Guinot, DANIELE 1961. Crustacés décapodes brachyoures de Tahite et des Tuamotu. In Expédition Frangaise récifs coral. Nouveau Calédonie, vol. pré- liminaire, 195 pp., 18 pls. Hansen, H. J. 1895. Isopoden, Cumaceen, und Stomatopoden der Plankton-Expedition. Vol. 2Ge in Hensen, Ergebnisse der in dem Atlantischen Ocean. . ., 105 pp. Hitton, W. A. 1915a. Pseudosquilla from Laguna. Pomona College Journ. Ent. Zool., vol. 7, no. 1, pp. 77-78 [unnum.], text fig. p. 78. 1915b. Pseudosquilla from Laguna. Pomona College Laguna Mar. Lab. Rep., no. 3, art. 20, 1 text p., 2 figs. Kemp, 8. 1913. An account of the Crustacea Stomatopoda of the Indo-Pacific region based on the collection in the Indian Museum. Mem. Indian Mus., vol. 4, no. 1, 217 pp., 10 pls. Lacny, O. L. 1953. Statistical methods in experimentation: An introduction, ix-+ 248 pp. Lamarck, J. B. P. A. DE M. DE 1818. Histoire naturelle des animaux sans vertébres.. ., Ist ed., vol. 5, 612 pp. LATREILLE, P. A. 1825. Histoire naturelle: Entomologie, ou histoire naturelle des Crustacés, des Arachnides et des Insectes. Vol. 10 in Encyclopédie methodi- que, 832 pp. [Not seen.] Lemos DE Castro, A. 1955. Contribuicao ao conhecimento dos Crustdceos da ordem Stomatopoda do litoral Brasileiro: (Crustacea, Hoplocarida). Bol. Mus. Nace. Rio de Janeiro, new series, Zoologica, vol. 128, 68 pp., 28 pls. Mannine, R. B. 1963a. Preliminary revision of the genera Pseudosquilla and Lysiosquilla with descriptions of six new genera (Crustacea: Stomatopoda). Bull. Mar. Sci. Gulf Caribbean, vol. 13, no. 2, pp. 308-328. 1963b. Hemisquilla ensigera (Owen, 1832) an earlier name for H. bigelowi (Rathbun, 1910) (Stomatopoda). Crustaceana, vol. 5, no. 4, pp. 315-317. 1967. Notes on some Australian and New Zealand Stomatopod Crustacea, with an account of the species collected by the F.I.8. ‘‘“Endeavour’’. Rec. Australian Mus., vol. 27, no. 4, pp. 79-137, 10 figs. Miers, E. J. 1880. On the Squillidae. Ann. Mag. Nat. Hist., vol. 5, no. 5, pp. 1-380, 108-127, pls. 1-3. Morerra, C. 1903a. Crustdceos, estudos preliminares: Companhas do pesca do _ hiate “Annie” dos Srs. Bandeira e Bravo. Lavoura, Biol. Soc. Nac. Agric. Brasil, vol. 7, nos. 1-38, pp. 60-67. 1903b. Crustdiceos da Ponta de Pharol em S4o Francisco do Sul, no estado de Santa Catharina. Arch. Mus. Nac. Rio de Janeiro, vol. 12, pp. 119-123. 1905. Campanhas de Pesca do ‘Annie’: Crustdéceos. Arch. Mus. Nace. Rio de Janeiro, vol. 13, pp. 123-145, pls. 1-5. 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Nicouet, H. 1849. Araneidae. Jn vol. 3 of Zoologia in Gay, Historia fisica y politica de Chile. . ., pp. 115-318. 1854. [Crustd4ceos.] pls. 1-4 in vol. 2 of Atlas in Gay, Historia fisica y politica de Chile. Owen, R. 1832. Characters of some new species of stomatopus Crustacea, collected by Mr. Cuming. Proc. Zool. Soc. London, no. 2, pp. 5-6. RANDALL, J. W. 1839. Catalogue of Crustacea brought by T. Nuttall and J. K. Townsend from the W. Coast of N. America, and the Sandwich Is.,. . Journ. Acad. Nat. Sci. Philadelphia, ser. 1, vol. 8, pp. 106-147. RatuBun, Mary J. 1910. The stalk-eyed Crustacea of Peru and the adjacent coast. Proc. U.S. Nat. Mus., vol. 38, pp. 531-620, pls. 36-56. Scumitt, W. L. 1940. The stomatopods of the west coast of America based on collections made by the Allan Hancock Expeditions, 1933-38. Allan Hancock Found. Pacific Exped., vol. 5, no. 4, pp. 129-255, 33 figs. SERENE, R. 1962. Révision du genre Pseudosquilla (Stomatopoda) et définition de genres nouveaux. Bull. Inst. Oceanogr. Monaco, no. 1241, pp. 1-27. STEPHENSON, W. 1953. Notes on Australian Stomatopoda (Crustacea) in the collections of the Queensland Museum. Mem. Queensland Mus., vol. 13, no. 1, pp. 40-49. 1954. Notes onstomatopod Crustacea from Victoria and Tasmania. Mem. Nat. Mus. Victoria, no. 19, pp. 1-4. STEPHENSON, W., and McNett, F. 1955. The Australian Stomatopoda (Crustacea) in the collections of the Australian Museum, with a check list and key to the known Australian species. Rec. Australian Mus., vol. 23, no. 5, pp. 239-265. WHITELEGGE, T. 1900. Crustacea, 1 Jn no. 2 of vol. 1 in Scientific results of the trawling expedition of H.M.C.S. Thetis. Mem. Australian Mus., vol. 4, pp. 185-199, pls. 32-35. U.S. GOVERNMENT PRINTING OFFICE: 1967 get na hin mihi RN ‘ ; P Wahi: ‘ee ; ad ti hii ; , ‘owall i " en mr Nihal ‘Dial Qik ata pr iy leh Minha J iB gg’ aa sia tas bbs lutart WP th the even) im. wen ee a ihe Be ng ‘ty ee Ve Rian. & ane Ay ie ie Ws :. nihil La i es: Pe i! lg mm " abies ii eee. oe ier RD TR RT IN md amet ee Mabe . ky Gia allitilehe, ae) ek ve in } | ia elaltant Ro my i : 7 7 | x e. i" i a y on , neh oe mM e eas v wine p ar ea er * ¥ yt ie e ae Pee 7 ‘ ; ve, 4, : ay : . | rT i a ae Mth, @ y: Are & i Wy, “) ; gre, + Cea hind eee x: ee me Sa Sn ek ; na Wie, Aiwa >a oe ‘ay oni : ieeheleadll in 1 _ mine ie, Mien Fai Amr ree ea vy lies i Tiny nna " Pete, (i ealaliea lle alaheil sui Loki pee EO) he [ lpi : ahaa ee at 4 me ri itd Proceedings of the United States National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. Volume 120 1967 Number 3565 THE BENTHIC POLYCHAETA AND AMPHIPODA OF MORRO BAY, CALIFORNIA By Donatp J. RetsH anp J. Laurens Barnarp! This paper records the present fauna of polychaete worms and amphipod crustaceans of Morro Bay, California, and reports upon their zoogeographic relationships to areas previously studied by the authors. Prior work in Morro Bay includes a study of the distribu- tion of the wood-boring isopod Limnoria and the Nebaliacea (Menzies and Mohr, 1952) and amphipods (Barnard, 1952). Although altered by dredging, the installation of wharfs, and the establishment of oyster farming, the environment of Morro Bay is of interest because of the sparsity of other embayments on the long coastline of central California. Previous studies on the benthic invertebrates of Californian bays and harbors include the following: Tomales Bay (Pitelka and Paulson, 1962), San Francisco Bay (Packard, 1918; Hartman, 1954a; Filice, 1954, 1958, 1959; Jones, 1961), Elkhorn Slough (MacGinitie, 1935), Los Angeles—Long Beach Harbors (Reish, 1959b), Alamitos Bay (Reish and Winter, 1954; Reish, 1961, 1963a), Newport Bay (Barnard and Reish, 1959), San Diego Bay (Anon., 1952), and Catalina Harbor, Santa Catalina 1Reish: Department of Biology, California State College, Long Beach, California; Barnard: Associate Curator, Division of Crustacea, Smithsonian Institution. Barnard’s work completed at Beaudette Foundation, Moss Land- ing, California. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Island (Reish, 1964). All of these Californian embayments have been altered to some extent by man so that the aboriginal faunas are unknown; presumably the biotas have been modified by the introduction of cosmopolitan bay forms (Barnard, 1961), by means of shipping, oyster culture, and other transmitting agents. Description oF Morro Bay.—Morro Bay (fig. 1) lies on the southern middle portion of the California coast, just north of the Zz Te) ° fo} N 35°2I' W MORRO BAY, CALIFORNIA station locations 2000 feet scale Ficure 1.—Station locations of benthic survey, August 25, 1960. zoogeographical boundary of Point Conception. Open-sea water temperatures approximate the southernmost extension of cold- temperate conditions and the rainfall is sufficient to permit a slight dilution of bay waters. This situation is in contrast to the embay- NO. 3565 POLYCHAETA AND AMPHIPODA—REISH AND BARNARD 3 ments south of Point Conception that belong with the warm- temperate, low-rainfall regime. Except where dredging has deepened some channels, Morro Bay is exceedingly shallow; much of the bay floor is exposed during minus tides. Two streams drain into the bay: Chorro Creek enters the Bay through a channel in the marshland (Stations 28 to 25) and Los Osos Creek enters near Station 29. The triangular marshland, extending from Stations 28 to 25 and then landward to the right in figure 1, constitutes the Morro Bay State Park Bird Reserve. Changes by man in Morro Bay include the dredging of channels, the construction of a small boat harbor at Stations 11 and 12, dock facilities along the shore extending from about Stations 3 to 11, jetties at the Bay entrance, and fencing for oyster beds. Small docks have been built at other localities of the bay. No pollution is known to exist (Barrett, 1963). A steam-generating plant near Station 3 takes in sea water from this area for use in cooling con- densors, but the water is discharged by a channel into the open sea north of the Bay. Although Morro Bay is important as a port for both commercial and sport fisheries, only the oyster, accounting for a value of $24,467 in 1961, is harvested directly from the bay. The total value of commercial landings for 1961 (Anon., 1963) was $632,324, with abalone accounting for over one-third of the total. The history of oyster farming in Morro Bay has been summarized by Barrett (1963). The Pacific oyster, Crassostrea gigas (Thunberg), was first introduced from Japan to Morro Bay in 1932. The beds extend throughout much of the middle part of the bay from about the region of Station 5 to Stations 21, 22, and 29 (fig. 1). MATERIALS AND METHODS.—Bottom samples, taken on August 25, 1960, from a small skiff using a size one Hayward orange-peel bucket (taking an area of %_ sq. meter), were obtained from all stations indi- cated in figure 1 with the exception of Stations 1, 2, 5, and 6 where the substrate was a hard-packed sand. The samples were washed on shore through a size 24 screen (0.7 mm openings) and preserved in formalin diluted with seawater. The polychaetes and amphipods were separated from the other organisms by Mr. Harold Pope. The polychaetes were identified by Reish and the amphipods by Barnard. The dissolved oxygen, salinity, and water temperature were determined on August 26, 1960, from 14 representative stations at surface and deeper levels; each varied according to depth of water. The modified Winkler method was used for dissolved oxygen meas- urement (Barnes, 1959), and the chlorinity was determined by the Mohr method (Barnes, 1959). Water temperature was measured with a thermistor-type thermometer. 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 CHEMICAL AND PHYSICAL DATA.—Chlorinity was that of normal sea water throughout the sampled areas of Morro Bay. The chlor- inity probably is lowered in the vicinities of Chorro and Los Osos Creeks during the winter rainy season. Dissolved oxygen of the water ranged from 4.2 to 7.1 ppm with the lowest value measured at Station 10 and the highest at Station 25. The oxygen values of the deeper waters were generally slightly higher than those of the surface readings. Temperature of the water ranged from 13.4° to 17.8° C on August 26, 1960. The lower measurements were recorded at the entrance of Morro Bay and the higher values at Stations 23, 25, and 26. Sediments were not analyzed for particle size, but their condition was noted at the time of collection. In general, the sedimentary characteristics of Morro Bay may be divided into three catagories. Sands predominate at the entrance of the bay and extend to about the region of stations 7 and 8. Fine sediments and extensive patches of eel-grass, Zostera marina, and sea lettuce, Ulva sp., occur throughout much of the rest of the bay. The sediments of Chorro Creek, es- pecially at Stations 27 and 28, consisted primarily of gravels with a thin layer of silts and fine sands on top. THE POLYCHAETE FAUNA The polychaetes collected from the benthos of Morro Bay are listed in table 1 and are compared with those of other embayments of California and Baja California for which data are available. Of the 34 species taken, 18% or 55% also occur in San Francisco Bay. Less than 50% of the species are common with areas studied south of Morro Bay. New distributional data are recorded for seven species. The northern distribution is herein extended for three species from southern California, namely, Prionospio pygmaeus, Spiophanes massionensis and Pista alata. Cossura longocirrata is extended southward from Vancouver. Pseudopolydora kempi is reported from the eastern Pacific Ocean for the first time. With reservation, Magelona papil- licornis is reported from the Pacific Ocean for the first time. Chone infundibuliformis was reported from Elkhorn Slough by Berkeley and Berkeley (1935). No endemic polychaete was found in Morro Bay. Sphaerosyllis hystrix, Typosyllis fasciata, Boccardia polybrancha, and Spirorbinae have not been reported previously from the bay environments in the eastern Pacific Ocean. Four species account for over 70% of the 9127 specimens of poly- chaetes collected from the bay: Capitella capitata (2434), Streblospio benedicti (1999), Heteromastus filiformis (1078), and Exogone lourer NO. 3565 POLYCHAETA AND AMPHIPODA—REISH AND BARNARD 5 TaBLE 1.—Comparison of the polychaetes from Morro Bay with those from other California bays San Los Alamitos New- Catalina San Fran- Angeles Bay 3 port MHarbor® Quintin Species cisco Long Bay 4 Bay 8 Bay ! Beach Harbors? Anattides williamsi x x Eteone californica x E. dilatae Hypoeulalia bilineata Exogone lourei Sphaerosyllis hystrix Typosyllis fasciata Nereis latescens N. procera Platynereis bicanaliculata Nephtys caecoides Lumbrineris zonata Stauronereis articulata Haploscoloplos pugettensis Scoloplos acmeceps Nerinides acuta Boccardia polybranchia P. ligni Prionospio cirrifera P. pygmaeus x Pseudopolydora kempi Streblospio benedicti Spiophanes missionensis x Magelona ? papillicornis Cossura longocirrata Ctenodrilus serratus Armandia bioculata Capitella capitata Heteromastus filiformis Axiothella rubrocincta Pista alata x Chone infundibuliformis Spirorbinae Number of species in common 18 15 16 16 12 15 Percent in common 55 45 48 48 35 45 rd rr dd aia Mr PM Mm Me MM Pm Pd PX PPS PP Pd Pd OP at eh ad aia! » Pos m4 rm a rs a Po Pd PS PS Mm MR ia rr Part PM x x | Hartman, 1954a; Jones, 1961. ? Reish, 1959b, 1961. * Reish, 1961, 1963a. ‘ Reish, 1959a. * Reish, 1964. 6 Reish, 1963b. (886). Of the 26 stations from which polychaetes were taken, E. louret was present at 19, H. filiformis at 17, C. capitata at 15, and S. benedicti at 14. The community structure of the polychaetes follows the general pattern of the sediment characteristics discussed above. The sandy 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 fauna is nearest the channel entrance and is dominated numerically by Armandia bioculata. The central part of the bay consists of fine sediments and is dominated numerically by Heteromastus filoformis and Exogone lourei. At individual stations C. capitata and Pasta alata may occur in large numbers and thus alter the community structure. The Chorro and Los Osos Creeks fauna is dominated numerically by S. benedicti and C. capitata; these two species are known to be tolerant of brackish waters elsewhere (Woodwick, 1953). Dr. Keith H. Woodwick, Fresno State College, identified the specimens of the genus Polydora, sensu lato, for which aid we are grateful. Family Phyllodocidae Anaitides williamsi Hartman Anaitides williamsi Hartman, 1936, p. 126, figs. 33-35.—Reish, 1963b, p. 408. MATERIAL.—Stations 9(1), 10(6), 11(8), 17(1), 19(1). DistRiBpuTION.—Oregon to Baja California. Eteone californica Hartman Eteone californica Hartman, 1936, p. 131, figs. 43-46; 1961, p. 12. MATERIAL.—Station 29(11). DistTRIBUTION.—Central and southern California. Eteone dilatae Hartman Eteone dilatae Hartman, 1936, pp. 130-131, figs. 40-42.—Reish, 1963b, p. 408. MATERIAL.—Stations 8(1), 12(1), 14(4), 16(1), 22(8), 25(14), 26(8), 30(2). DistRipuTIoN.—Central California to Baja California. Hypoeulalia bilineata (Johnston) Phyllodoce bilineata Johnston, 1840, p. 227. Eulalia bilineata (Johnston).—Pettibone, 1963, pp. 86-88, fig. 20. Hypoeulalia bilineata (Johnston).—Hartman, 1961, p. 13.—Reish, 1963b, p. 423. MATERIAL.—Stations 8(1), 11(4). DistrRisuTIoN.—British Columbia to southern California, Nova Scotia to North Carolina, Arctic Ocean, Europe, Japan, and South Africa. Family Syllidae Exogone lourei Berkeley and Berkeley Exogone lourei Berkeley and Berkeley, 1938, pp. 44-47, figs. 6-10.—Reish, 1959b, p. 80. MATERIAL.—Stations 7(1), 8(1), 9(81), 10(102), 11(7), 12(4), 13(), 15(17), 16(260), 17(1), 18(18), 19(1), 21(1), 22(808), 23(22), 24(5), 26 (2), 29(102), 30(2). DistrRisuTION.—British Columbia, California, and Acapulco, Mexico. NO. 3565 POLYCHAETA AND AMPHIPODA—REISH AND BARNARD i Sphaerosyllis hystrix Claparéde Sphaerosyllis hystrix Claparéde, 1863, p. 45, pl. 13, figs. 36-37.—Berkeley and Berkeley, 1948, p. 80, fig. 119.—Hartman, 1961, p. 16.—Pettibone, 1963, pp. 136-137, fig. 35g. MATERIAL.—Stations 9(6), 12(15), 16(12), 19(2), 20(1), 21(1), 22(150); 27(1). DistrisuTion.—British Columbia to southern California, Con- necticut, Europe. Typosyllis fasciata (Malmgren) Syllis fasciata Malmgren, 1867, p. 43, pl. 8, fig. 47.—Berkeley and Berkeley, 1948, pp. 74-75, figs. 109-110. Syllis (Typosyllis) fasicata.—Pettibone, 1954, pp. 254-255, figs. 28 c-e. Typosyllis fasicata (Malmgren).—Imajima and Hartman, 1964, pp. 135-136, pl. 33, figs. j-o. MATERIAL.—Station 11(8). Distrisution.—North Atlantic Arctic, Japan, China, Alaska south to southern California. Family Nereidae Nereis latescens Chamberlin Nereis latescens Chamberlin, 1919, pp. 10-11.—Hartman, 1961, p.18. MATERIAL.—Station 11(1). DistrisutTion.—Central and southern California. Nereis procera Ehlers Nereis procera Ehlers, 1868, p. 557-559, pl. 23, fig. 2—Reish, 1959b, pp. 81-82. MATERIAL.—Station 24(2). Distrisution.—Alaska to southern California. Platynereis bicanaliculata (Baird) Nereis bicanaliculata Baird, 1863, p. 109. Platynereis bicanaliculata—Hartman, 1954b, pp. 36-39, figs. 38-39.—Reish, 1963b, p. 424. MaTERIAL.—Stations 7(6), 8(6), 9(9), 10(3), 11(25), 12(1), 15(53), 18(7), 21(46), 22(6), 24(24), 26(9), 27(1), 29(1). Distrisution.—Northeast Pacific Ocean from British Columbia to Baja California, Hawaii, Australia. Family Nephtyidae Nephtys caecoides Hartman Nephtys caecoides Hartman, 1938b, pp. 148-149, fig. 63; 1950, pp. 101-102. MATERIAL.—Stations 13(4), 17(1), 20(3), 22(2). DistRiBUTION.—British Columbia to Baja California. 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Family Lumbrineridae Lumbrineris zonata (Johnson) Lumbriconereis zonata Johnson, 1901, pp. 408-409, pl. 9, figs. 93-100. Lumbrineris zonata (Johnson).—Hartman, 1944a, pp. 146-147. MaTERIAL.—Stations 7(8), 8(1), 9(11), 10(21), 11(8), 12(1), 16(8), 17(2). DistrIBuTION.—Washington to Baja California. Family Dorvilleidae Stauronereis articulatus Hartman Stauronereis articulatus Hartman, 1938a, pp. 101-102. Dorvillea articulata Hartman, 1944a, p. 189.—Reish, 1963b, p. 426. MaTERIAL.—Stations 9(1), 10(8), 15(8), 16(2), 18(8), 21(1), 22(6), 24 (4). DistrisutTion.—Central California to Baja California. Remarxs.—Stauronereis Verrill (1900) was re-established by Petti- bone (1961). Pettibone (1963) placed S. articulatus in synonymy with S. rudolphi (Delle Chiaje), but we are retaining the name SV. articulatus for the eastern Pacific population until detailed compari- sons can be made with collections from European waters. Family Orbiniidae Haploscoloplos pugettensis (Pettibone) Scoloplos elongata Johnson, 1901, pp. 412-413, pl. 10, figs. 105-110. Haploscoloplos elongatus (Johnson).—Hartman, 1944b, p. 257; 1957, pp. 273-275, pl. 26, figs. 1-11.—Reish, 1963b, p. 426. Scoloplos (Scoloplos) pugettensis Pettibone, 1957, p. 162. MaTERIAL.—Stations 11(8), 16(1), 17(2), 20(2). Distripution.—Alaska to Baja California. Remarxs.—Scoloplos elongata Johnson, 1901, is a junior homonym of S. elongata Quatrefages, 1866. Pettibone (1957) renamed Johnson’s species as S. (Scoloplos) pugettensis. Earlier Hartman (1944b) re- ferred S. elongata Johnson to the genus Haploscoloplos Monro, 1933, because of the absence of thoracic uncini in this genus. Pettibone, in 1954 (p. 279) and again in 1957 (p. 160) indicated the difficulty in separating worn thoracic capillary setae in the genus Haploscoloplos from the thoracic uncini of the genus Scoloplos. This viewpoint was not followed by Hartman (1957) nor Berkeley and Berkeley (1958). We are therefore retaining the new name given by Pettibone, as re- quired by the rules of nomenclature, and placing the species in the genus Haploscoloplos. No. 3566 POLYCHAETA AND AMPHIPODA—REISH AND BARNARD 9 Scoloplos acmeceps Chamberlin Scoloplos acmeceps Chamberlin, 1919, pp. 15-16.—Hartman, 1957, pp. 282-283, pl. 30, figs. 1-7. MATERIAL.—Stations 7(1), 8(1), 9(13), 10(81), 11(2), 19(2). Distrisution.—Alaska south to Mazatlan, Mexico. Family Spionidae Nerinides acuta (Treadwell) Spio acuta Treadwell, 1914, pp. 199-201, pl. 11, figs. 14-20. Nerinides acuta (Treadwell).—Hartman, 1941, pp. 294-296, pl. 45, figs. 1-8; pl. 47, fig. 29; 1954a, p. 10. MATERIAL.—Stations 9(8), 10(45), 11(4), 12(1), 19(2), 22(15), 23 (2), 27(6), 29(1). DistrisutTion.—Central and southern California. Boccardia uncata Berkeley Boccardia uncata Berkeley, 1927, p. 418, pl. 1, figs. 9-13.—Hartman, 1961, p. 28. Polydora (Boccardia) uncata.—Berkeley and Berkeley, 1952, pp. 14-15, figs. 18-21.—Reish, 1963b, p. 427. MATERIAL.— Stations 15(1), 16(23), 23(14), 24(10), 27(2). DistrisuTiIon.—British Columbia to Baja California. Boccardia polybranchia (Haswell) Polydora polybranchia Haswell, 1885, p. 275. Polydora (Boccardia) polybranchia.—Berkeley and Berkeley, 1952, pp. 16-17, figs. 24-25.—Reish, 1959b, p. 38. Boccardia polybranchia (Haswell).—Hartman, 1959, p. 375. MatTERIAL.—Station 25(130). DistrisuTion.—Europe, Japan, Australia, British Columbia, and central and southern California. Polydora ligni Webster Polydora ligni Webster, 1879, p. 119.—Hartman, 1941, pp. 309-310, pl. 48, figs. 47-49.— Berkeley and Berkeley, 1952, p. 19, figs. 31-33. MaTERIAL.—Stations 15(3), 21(2), 25(8), 27(14), 28(192), 30(83). Distrisution.—Both sides of United States, British Columbia, Mexico. Prionospio cirrifera Wiren Prionospio cirrifera Wiren, 1883, p. 409.—Berkeley and Berkeley, 1952, pp. 28- 29, figs. 52-53. MATERIAL.—Stations 23(1), 25(1). DistRiBUTION.—Europe, Arctic, India, Bering Sea to southern California. Prionospio pygmaeus Hartman Prionospio pygmaeus Hartman, 1961, pp. 93-95.—Reish, 1963b, p. 427. MareEriAL.—Stations 10(1), 15(1). 221-529—67——_2 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 DisTRIBUTION.—This species was known previously only from southern California and Baja California. Pseudopolydora kempi (Southern) Polydora (Carazzia) kempi Southern, 1921 p. 636, pl. 28, fig. 20.—Fauvel, 1953, pp. 317-318, fig. 167 a-c.—Chlebovitsch, 1961, pp. 199-200. Polydora kempi.—Day, 1957, pp. 99-100. Pseudopolydora kempi (Southern).— Hartman, 1959, p. 387. MaTERIAL.—Stations 10(44), 13(1), 17(1), 18(8), 19(2), 22(74), 25(1), 26(25), 27(29), 29(84), 30(2). DistRiBuTION.—India, Japan, Kurile Islands, and South Africa. This is the first report of the species from the eastern Pacific Ocean. Spiophanes missionensis Hartman Spiophanes missionensis Hartman, 1941, pp. 296-298, pl. 46, figs. 17-21.—Reish, 1963b, p. 427. MaATERIAL.— Station 12(1). DisTRIBUTION.—Known previously from southern California to Baja California. Streblospio benedicti Webster Streblospio benedicti Webster, 1879, pp. 120-121.—Hartman, 1944b, p. 260; 1945, p. 34, pl. 6, fig. 4. MaTERIAL.—Stations 11(2), 12(45), 13(83), 14(8), 19(6), 21(5), 22(216), 23(2), 25(1), 26437), 27(740), 2843), 29(255), 3011): DistRipuTION.—New Jersey, Massachusetts, North Carolina, cen- tral and southern California. Family Magelonidae Magelona ?papillicornis (Miiller) ? Magelona papillicornis (Miiller).—fide Fauvel, 1927, pp. 64-65, fig. 22 a-k. MATERIAL.—Station 4(1). Remarxks.—This specimen comes closest to M. papillicornis (fide Fauvel, 1927), but since only one worm in poor condition was collected and since it has not been reported previously from the Pacific Ocean, it is referred to this species with reservation. Family Cirratulidae Cossura longocirrata Webster and Benedict Cossura longocirrata Webster and Benedict, 1887, p. 743.—Berkeley and Berkeley, 1956, pp. 544-545.—Reish, 1965, p. 145. MATERIAL.—Stations 12(41), 13(80), 26(1). Distrisution.—North Atlantic, Maine, Russian Pacific, Bering and Chukchi Seas, and Washington. This record of the species from Morro Bay is a southern extension of its distribution in the eastern Pacific Ocean. NO. 3565 POLYCHAETA AND AMPHIPODA—REISH AND BARNARD Ala Family Ctenodrilidae Ctenodrilus serratus (Schmidt) Parthenope serratus Schmidt, 1857, p. 363, pl. 5, fig. 18. Ctenodrilus serratus (Schmidt).—Fauvel, 1927, pp. 108-109, figs. 38a-e.—Hartman, 1961, p. 32. MATERIAL.—Stations 11(2), 26(2). DistTRIBUTION.—Cosmopolitan. Family Ophelidae Armandia bioculata Hartman Armandia bioculata Hartman, 1938a, pp. 105-106, figs. 51-54.—Reish, 1963b, p. 428. MATERIAL.—Stations 3(1), 7(62), 8(29), 9(78), 10(4), 11(9), 18@3), 19(1), 20(2), 21(1), 22(12), 26(11). Distrisution.—Northern California to Baja California. Family Capitellidae Capitella capitata (Fabricius) Lumbricus capitatus Fabricius, 1780, p. 279. Capitella capitata (Fabricius).—Hartman, 1947, pp. 404-405, pl. 43, figs. 1-2. MaATERIALS.—Stations 7(166), 8(149), 10(1), 13(8), 18(179), 21(5), 22 (25), 23(11), 24(2), 25(27), 26(190), 27(1628), 28(14), 29(5), 30(29). DistTRIBUTION.—Cosmopolitan. Heteromastus filiformis (Claparéde) Capitella filiformis Claparéde, 1864, p. 509, pl. 4, fig. 10. Heteromastus filiformis (Claparéde).—Hartman, 1947, pp. 427-428, pl. 52, figs. 1-4. MATERIAL.—Stations 8(42), 9(32), 10(181), 11(89), 12(23), 13(84), 14(1), 16(3), 17(44), 18(34), 19(101), 21(1), 22(173), 23(1), 25(5), 26 (26), 29(288). DisTRIBUTION.—Widely distributed in both hemispheres. Family Maldanidae Axiothella rubrocincta (Johnson) Clymenella rubrocincta Johnson, 1901, pp. 418-419, pl. 13, figs. 128-1383. Aziothella rubrocincta (Johnson).—Berkeley and Berkeley, 1952, pp. 51-52, figs. 105-106.—Reish, 1963b, p. 429. MATERIAL.—Stations 18(1), 20(1), 22(89). DistRIBsuTION.—British Columbia to Baja California. Family Terebellidae Pista alata Moore Pista (Scionopsis) alata Moore, 1909, pp. 273-275, pl. 9, figs. 48-51. Pista alata.—Reish, 1963b, p. 430. 12 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 120 MATERIAL.—Stations 9(2), 12(1), 13(1), 15(15), 18(125), 20(1), 21 (6), 2281). Disrrinution.—Heretofore known only from southern California and San Quintin Bay, Baja California. Family Sabellidae Chone infundibuliformis Kréyer Chone infundibuliformis Kréyer, 1856, p. 33.—Berkeley and Berkeley, 1952, p. 133, figs. 252-253.—Pettibone, 1954, pp. 338-339, figs. 39a-j. MATERIAL.—Stations 7(7), 10(88), 11(29), 12(1), 14(2), 17(7), 18(1), 19(89), 20(69), 22(4). DistrirputTion.—North Atlantic, Arctic, British Columbia, Cali- fornia. Family Serpulidae Subfamily Spirorbinae MATERIAL.—Stations 12(8), 15(512), 16(8). Remarks.—No attempt was made to identify these specimens. Many were observed attached to blades of the eel grass Zostera marina. THe AmpPHIPOD FAUNA In contrast to the other well-known bays of the Californias, Bahfa de San Quintin and Newport Bay, the amphipod fauna of Morro Bay is characterized by several boreal elements and by a few species associated with brackish water. Although the waters of Morro Bay were not brackish at the time of this survey, the presence of Corophium spinicorne and the abundance of C. wenor and C. ache- rusicum suggest a history of dilution. Of course these species may also be linked with environments which are abnormal in factors other than dilution, such as natural and artificial pollution and wide ranges in thermal regime. ‘The boreal elements known to occur in the Morro Bay fauna, such as Ampithoe lacertosa, Allorchestes angustus, Aoroides columbiae, Pontogeneia rostrata, Ampithoe valida, and possibly Hohaus- torius washingtonianus, which are lost or diminished in Newport Bay (Barnard, 1959) and Bahia de San Quintin (Barnard, 1964), may not be due so much to the supposed increase in average temperatures of the southern bays as they are to the greater yearly range of tempera- tures there. An analysis of the ecological and zoogeographical distribution of the amphipod species encountered showed that all of the species have been derived from the open sea, or have had to find a pathway through it. Of all species encountered, Corophium spinicorne is the most firmly bound to estuarine conditions. Those species that are considered as primarily of estuarine habit, because of the absence or near absence of NO. 2565 POLYCHAETA AND AMPHIPODA—REISH AND BARNARD 13 their populations in the open sea at these latitudes, include Oorophium uenoi, Allorchestes angustus, Hlasmopus rapaz (sensu stricto), Ampithoe valida, Pontogeneia minuta, and Ampithoe longimana. These are pri- marily confined to the innermost reaches of the bay, in or near marsh- land channels. Generally, they are also among the most abundant species of any of the samples, as the following ranked list of species and numbers of individuals in 26 samples shows: Corophium uenoi (1131), Aoroides columbiae (808), Corophium spinicorne (333), Ampi- thoe lacertosa (242), Microdeutopus schmitti (205), Pontogeneia rostrata (194), Corophium acherusicum (187), Allorchestes angustus (116). Note, however, that Hlasmopus rapaz, Ampithoe valida, Pontogeneia minuta, and Ampithoe longimana are rare in these samples. The dis- continuity of many of the typical open-sea species from the sea to the inner reaches of Morro Bay is probably due to the unsuitable, coarse, sandy substrates, rapid tidal currents and lack of benthic algae at the seaward stations. Species and genera known to occur in Bahia de San Quintin (table 2) that were not discovered in Morro Bay and that are considered to be absent because of geographic thermal differences are: Acwmino- deutopus, Rudilemboides, Lembos macromanus, Amphideutopus, Orcho- mene magdalenensis, and Pontogeneia quinsana. Other San Quintin species that, in the open sea, extend to the north of Morro Bay and probably are excluded from the estuary for reasons other than thermal structure are Ampelisca compressa and Hyale frequens. The absence of both species is striking and unexplainable. The most unusual difference between Bahia de San Quintin and Morro Bay is in the species of Ampelisca that dominate the benthos; in San Quintin Ampelisca compressa is the dominant; this species occurs northward in the open sea to Puget Sound, Washington, but in Morro Bay (and in Tomales Bay, material at hand), the principal species is A. cristata. Both A. compressa and A. cristata, however, are known to occur in tropical Pacific and tropical Atlantic America, so that neither can be considered a warm or a cool-sea species. Other species of Ampelisca occur only incidentally in Newport Bay. Although the plant is widespread in Morro Bay, stands of the eel- grass, Zostera marina, do not seem as dense in Morro Bay as in Bahia de San Quintin. That eel-grass is not so prominent in Morro Bay as in San Quintin is shown by the absence or low rank of Hyale, Ponto- geneia, and Ericthonius, genera of amphipods which characterize the San Quintin eel-grass beds (Barnard, 1964). On the other hand, a relatively higher incidence of algae is indicated in Morro Bay by the high rankings of Corophium, Aoroides, and Ampithoe. The influence of the dense populations associated with pilings in Newport Bay is now confirmed in comparing the fauna of that bay 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 120 with that of San Quintin Bay and Morro Bay. The bottom fauna at Newport has a number of highly ranked species (table 2) that ap- parently represent extravagants from the piling fauna: Leucothoides pacifica, Colomastix pusilla, Jassa falcata, and both species of Podocerus. TABLE 2.—Dominant amphipods listed by rank according to their frequencies in Morro Bay, Newport Bay, and Bahfa de San Quintin Morro Bay Newport Bay Bahfa de San Quintin (herein) (Barnard, 1959, 1961) (Barnard, 1964) Corophium uenot Elasmopus rapax Ampelisca compressa Aoroides columbiae Acuminodeutopus Ericthonius brasiliensis heteruropus Corophium spinicorne Rudilemboides steno- Rudilemboides steno- propodus propodus Ampithoe lacertosa Ampitthoe plumulosa ! Acuminodeutopus heteru- ropus Microdeutopus schmitti Ampithoe pollex Hyale frequens (as nigra) Pontogeneia rostrata Leucothoides pacifica Aruga holmesi Corophium acherusicum Corophium acherusicum Microdeutopus schmitti Allorchestes angustus Corophium bacont Paraphozus obtusidens Heterophoxus oculatus Hyale frequens Lembos macromanus Ampelisca cristata Paraphozus spinosus Amphideutopus oculatus Photis brevipes Ericthonius brasiliensis Orchomene magdalenensis Synchelidium shoemakert Colomastix pusilla Corophium uenoi Elasmopus rapax Amphideutopus oculatus Listriella melanica Ampithoe valida Corophium uenot Elasmopus rapax Synchelidium rectipalmum Amphilochus neapolitanus Uristes entalladurus Batea transversa Maera simile Corophium bacont Paraphoxus stenodes Microdeutopus schmitti Pontogeneia quinsana Pontogeneia minuta Jassa falcata Corophium acherusicum Ampithoe longimana Podocerus brasiliensis Paraphoxus heterocuspi- datus Corophium baconi Podocerus fulanus Ampithoe plumulosa 1 Table 1 of Barnard (1961) entry ‘‘4’’ should read Ampithoe plumulosa not A. pollex. Family Ampeliscidae Ampelisca cristata Holmes Ampelisca cristata Holmes, 1908, pp. 507-508, figs. 16, 17.—Barnard, 1954a, pp. 26-29, pls. 17-18; 1959, p. 18. MaTERIAL.— Stations 10(1), 12(14), 20(5), 21(10), 22(2), 25(3). DiIstRIBUTION.—Caribbean Sea; eastern Pacific from Ecuador to Tomales Bay, Calif. NO. 3565 POLYCHAETA AND AMPHIPODA—REISH AND BARNARD 15 Family Ampithoidae Ampithoe lacertosa Bate Ampithoe lacertosa Bate, 1858, p. 362.—Gurjanova, 1951, pp. 895-897, fig. 622.— Barnard, 1954b, pp. 31-33, pls. 29-30.—Nagata, 1960, pp. 175-176, pl. 16, figs. 95-96. MaTERIAL.—Stations 9(29), 16(206), 18(7). DistTRIBUTION.—Japan; Kodiak, Alaska to Magdalena Bay, Baja California. Ampithoe longimana Smith Ampithoe longimana Smith, 1873, pp. 563-564.— Barnard, 1959, pp. 36-37, pl. 12; 1964, p. 111. MATERIAL.—Stations 9(1), 21(4). DistRiBuTION.—Atlantic coast of the United States and Bermuda; eastern Pacific Ocean only in bays (Morro Bay, Newport Bay, Bahia de San Quintin). Ampithoe valida Smith Ampithoe valida Smith, 1873, p. 563.—Barnard, 1954b, pp. 34-35, pl. 31.—Nagata, 1960, p. 176, pl. 16, figs. 97, 98. MATERIAL.—Stations 26(6), 28(4). Distrisution.—Atlantic coast of United States; Japan; eastern Pacific Ocean from Coos Bay, Oreg., to Morro Bay. Family Aoridae Aoroides columbiae Walker Aoroides columbiae Walker, 1898, p. 285, pl. 16, figs. 7-10.—Barnard, 1954b, pp. 24-26, pl. 22; 1959, p. 33.—Nagata, 1960, p. 175, pl. 16, fig. 94.—Barnard, 1961, p. 180; 1964, p. 110. MATERIAL.—Stations 7(15), 8(10), 9(473), 10(34), 15(8), 18(266), Bilt) ol). DistRIBuTION.—Puget Sound to Bahia de San Quintin, Baja California. Microdeutopus schmitti Shoemaker Microdeutopus schmitti Shoemaker, 1942, pp. 18-21, fig. 6.—Barnard 1959, pp. 32-33, pl. 9; 1961, p. 180; 1964, p. 110, chart 13. MATERIAL.—Stations 15(114), 21(52), 24(39). DistrRiBuUTION.—Monterey Bay to Cape San Lucas, Baja Califor- nia. Family Bateidae Batea transversa Shoemaker Batea transversa Shoemaker, 1926, pp. 13-18, figs. 8-11. MATERIAL.—Stations 15(5), 18(1). 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 DistTRIBUTION.—Southern California at Morro Bay, Catalina Island, and Point Loma. Family Corophiidae Corophium acherusicum Costa Corophium acherusicum Costa, 1857, p. 232.—Shoemaker, 1947, p. 53, figs. 2, 3; 1949, p. 76.—Barnard, 1954b, p. 36; 1959, p. 38; 1961, p. 182; 1964, p. 111, chart 5. MaTERIAL.—Stations 9(1), 15(5), 16(1438), 21(1), 23(37). DisTRIBUTION.—Cosmopolitan in temperate and tropical waters, especially in bays and harbors. Corophium baconi Shoemaker Corophium baconi Shoemaker, 1934, pp. 356-359, fig. 1; 1949, p. 82.—Barnard, 1959, p. 38; 1961, p. 182; 1964, pp. 111-112, chart 16. MAaTERIAL.—Station 15(4). DistTRIBUTION.—Peru to the Bering Sea. Corophium spinicorne Stimpson Corophium spinicorne Stimpson, 1857, pp. 514-515.—Shoemaker, 1949, pp. 74-76, fig. 6—Barnard, 1952, p. 33; 1954b, pp. 36-37. MATERIAL.—Stations 26(327), 27(5), 28(1). Distrisution.—Alaska to Morro Bay, Calif. Corophium uenoi Stephensen Corophium uenoi Stephensen, 1932, pp. 494-498, figs. 3, 4—Barnard, 1952, pp. 28-32, pls. 8, 9; 1959, p. 39.—Nagata, 1960, p. 178.— Barnard, 1961, p. 183; 1964, p. 112, chart 16. MATERIAL.—Stations 16(260), 17(8), 21(2), 22(1), 23(31), 26(18), 27(291), 28(517), 29(4), 30(4). DistriBpuTIon.—Japan; Morro Bay to Bahia de San Quintin in the Californias. Ericthonius sp. [cf. brasiliensis (Dana)] MATERIAL.—Station 27 (1 juv.). Probably this specimen is the juvenile of E. brasiliensis. See Sars (1895, pl. 215) for illustration and Barnard (1959 and 1964) for references and distribution of this species. DistTrIBUTION.—Cosmopolitan (except in polar regions) in bays and harbors and the open-coast intertidal. Family Eusiridae Pontogeneia minuta Chevreux Pontogeneia minuta Chevreux, 1908, pp. 1-3, fig. 1—Barnard, 1959, p. 23, pl. 3; 1964, p. 106, figs. 21 B, C. MATERIAL.—Stations 26(4), 27(1). NO. 3565 POLYCHAETA AND AMPHIPODA—REISH AND BARNARD Ly) DistrisutTion.—In the Pacific found in Morro Bay, Newport Bay, and Bahia de San Quintin, Baja California. The identification of the Pacific forms with Chevreux’s species is open to question. Pontogeneia rostrata Gurjanova Pontogeneia rostrata Gurjanova, 1938, p. 330, fig. 39.—Barnard, 1962b, p. 81. MatTERIAL.—Stations 4(1), 9(87), 10(21), 15(5), 18(79). Distripution.—Bering Sea, Okhotsk Sea, Japan Sea to Bahfa de San Quintin, Baja California. Family Gammaridae Elasmopus rapax Costa Elasmopus rapax Costa, 1853, p. 175.—Sars, 1895, pp. 521-522, pl. 183.—Shoe maker, 1942, p. 12.—Barnard, 1959, pp. 23-24; 1962b, pp. 94-96, figs. 16, 17, 1964, p. 108, chart 9. MAaATERIAL.— Stations 27(9), 28(1). DistTRIBuTION.—Pantropical with penetration to warm-temperate and rarely cold-temperate waters in the northeastern Atlantic. This is a bay form; however, an open-sea Californian form has been de- scribed by Barnard (1962b). Family Haustoriidae Eohaustorius washingtonianus (Thorsteinson) Haustorius washingtonianus Thorsteinson, 1941, pp. 61-62. pl. 4, figs. 39-51. Eohaustorius washingtonianus Barnard, 1957, p. 82, pl. 16.—Gurjanova, 1962, pp. 404-405, fig. 135. MATERIAL.—Station 4(2). DistriputTion.—Puget Sound to just southeast of Point Conception, Calif. Family Hyalidae Allorchestes angustus Dana Allorchestes angustus Dana, 1856, p. 177.—Barnard, 1952, pp. 20-23, pl. 5, figs. 2-6; 1954b, pp. 21-23, pl. 21; 1959, p. 28. MATERIAL.—Stations 16(33), 26(77), 28(6). DistRiBUTION.—Japan; Coos Bay, Oreg., to Newport Bay, Calif. Family Ischyroceridae Jassa falcata (Montagu) Jassa falcata (Montagu).—Sexton and Reid, 1951, pp. 30-47, pls. 4-30 (with synonymy).—Barnard, 1959, p. 37; 1964, p. 118. MATERIAL.—Station 9(1). DistTRIBUTION.—Cosmopolitan in cold-temperate and_ tropical waters, especially in bays and harbors. 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Family Oedicerotidae Synchelidium rectipalmum Mills Synchelidium rectipalmum Mills, 1962, pp. 17-19, fig. 5. MatTERIAL.—Stations 15(3), 21(4), 22(1). DistRiBUTION.—British Columbia to middle Baja California. Synchelidium shoemakeri Mills Synchelidium shoemakeri Mills, 1962, pp. 15-17, fig. 4. MaTERIAL.—Stations 7(1), 14(1), 20(10). DistrRiBpuTION.—British Columbia to middle Baja California. Family Photidae Photis brevipes Shoemaker Photis brevipes Shoemaker, 1942, pp. 25-27, fig. 9—Barnard, 1962a, pp. 31-33, sohed, Jhile MarTERIAL.—Stations 7(1), 8(2), 9(24), 12(3), 18(3). DistripuTION.—Coos Bay, Oreg., to Magdalena Bay, Baja Cali- fornia. Family Phoxocephalidae Heterophoxus oculatus (Holmes) Harpinia oculata Holmes, 1908, pp. 521-523, fig. 28. Heterophoxus oculatus (Holmes).—Barnard, 1960, pp. 320-324, pls. 59-61; 1964, p. 102. MarTERIAL.—Stations 12(5), 15(66), 16(2), 24(25). DistriBuTION.—Puget Sound to Panama, an open-sea deep-water species, 13-1785 meters, occasionally penetrating shallow bays such as Morro Bay and San Quintin Bay. Paraphoxus epistomus (Shoemaker) Pontharpinia epistoma Shoemaker, 1938, pp. 326-329, fig. 1. Paraphoxus epistomus (Shoemaker).—Barnard, 1960, pp. 205-209, pls. 6-8. MaTERIAL.— Station 14(1). DistriBuTION.—Mendocino County, Calif., to Panama; Atlantic Ocean from New Hampshire to South Carolina. Paraphoxus spinosus Holmes Paraphorus spinosus Holmes, 1905, pp. 477-478, fig. 12.—Barnard, 1959, p. 18; 1960, pp. 243-249, pls. 29-31; 1964, p. 105. MaTERIAL.— Station 14(1). DistrRipuTion.—Western Atlantic Ocean; Pacific Ocean from Puget Sound to the Gulf of California. NO. 3565 POLYCHAETA AND AMPHIPODA—REISH AND BARNARD 19 Paraphoxus stenodes Barnard Paraphozrus stenodes Barnard, 1960, pp. 221-224, pls. 17, 18. MaTERIAL.—Stations 4(2), 14(1), 20(2). DistrisuTion.—Point Conception, Calif., to San Cristobal Bay, Baja California. Miscellaneous Amphipod Specimens Ampithoe species, juveniles MaTERIAL.—Stations 9(9), 10(1), 15(5), 17(1), 23(1), 24(1), 27(2). Corophium species, juveniles MaTERIAL.—Stations 7(2), 9(1), 16(10), 18(1), 21(1), 23(3), 24 (1), 2 (32), 28(386). Ericthonius species, juveniles MATERIAL.—Station 27(1). Photis species, juveniles MatTERIAL.—Stations 10(2), 15(3). Literature Cited ANONYMOUS 1952. Report upon the extent, effects and limitations of waste disposal into San Diego Bay. San Diego Regional Water Pollution Control Board, pp. 1-95. 1963. The California marine fish catch for 1961. California Fish and Game Fish Bull., no. 121, pp. 1-47, figs. 1-8. Barrp, WILLIAM 1863. 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Scumipt, O. 1857. Zur Kenntnis der Turbellaria rhabdocoela und einiger anderer Wiirmer des Mittelmeeres. Akad. Wiss. Wien Sitzber., vol. 23, pp. 347-366, pls. 1-5. Sexton, E. W., and Reip, D. M. 1951. The life-history of the multiform species Jassa falcata (Montagu) (Crustacea Amphipoda) with a review of the bibliography of the species. Journ. Linnean Soc. London, vol. 42, Zool., pp. 29-91, pls. 4-30. No. 3565 POLYCHAETA AND AMPHIPODA—REISH AND BARNARD 25 SHOEMAKER, C. R. 1926. Amphipods of the family Bateidae in the collection of the United States National Museum. Proc. U.S. Nat. Mus., vol. 68, pp. 1-26, figs. 1-16. 1934. Two new species of Corophium from the west coast of America. Journ. Washington Acad. Sci., vol. 24, pp. 356-360, figs. 1-2. 1938. Two new species of amphipod crustaceans from the east coast of the United States. Journ. Washington Acad. Sci., vol. 28, pp. 326-332, figs. 1-2. 1942. Amphipod crustaceans collected on the Presidential Cruise of 1938. Smithsonian Misc. Coll., vol. 101, no. 11, pp. 1-52, figs. 1-17. 1947. Further notes on the amphipod genus Corophium from the east coast of America. Journ. Washington Acad. Sci., vol. 37, pp. 47-63, figs. 1-12. 1949. The amphipod genus Corophium on the west coast of America. Journ. Washington Acad. Sci., vol. 39, pp. 66-82, figs. 1-8. Smita, S. I. 1873. Crustacea. Pp. 545-580 in Isopoda in Verrill, Report upon the in- vertebrate animals of Vineyard Sound and the adjacent waters with an account of the physical characters of the region, pt. 1 in U.S. Commission of Fish and Fisheries, Report on the condition of the sea fisheries of the south coast of New England in 1871 and 1872, pp. 295-778, pls. 1-38, figs. 1-4. SouTHERN, R. 1921. Polychaeta of the Chilka Lake and also of fresh and brackish waters in other parts of India. Mem. Indian Mus. Calcutta, vol. 5, pp. 563-659, pls. 19-31, figs. 1-18. STEPHENSEN, K. 1932. Some new amphipods from Japan. Annot. Zool. Japon. [sic], vol. 13, pp. 487-501, figs. 1-5. Stimpson, W. 1857. On the Crustacea and Echinodermata of the Pacific shores of North America. Boston Journ. Nat. Hist., vol. 6, pp. 444-532, pls. 18-23. THORSTEINSON, E. D. 1941. New or noteworthy amphipods from the North Pacific coast. Univ. Washington Publ. Oceanogr., vol. 4, pp. 50-96, pls. 1-8. TREADWELL, A. L. 1914. Polychaetous annelids of the Pacific coast in the collection of the zoological museum of the University of California. Univ. Califor- nia Publ. Zool., vol. 13, pp. 175-234, pls. 11-12. Wa ker, A. O. 1898. Crustacea collected by W. A. Herdman F.R.S., in Puget Sound, Pacific coast of North America, September, 1897. Proc. Trans. Liverpool Biol. Soc., vol. 12, pp. 268-287, pls. 15-16. Wesster, H. E. 1879. Annelida Chaetopoda of the Virginia coast. Trans. Albany Inst., vol. 9, pp. 101-128, pls. 1-11. Wesster, H. E., and Benepict, J. E. 1887. The Annelida Chaetopoda from Provincetown and Wellfleet, Massa- chusetts. Rep. U.S. Fish Comm. 1885, pp. 707-755, pls. 1-8. 26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 WIREN, AXEL 1883. Chaetopoder from Sibiriska Ishafvet och Berings Haf insamlade under Vega-Expeditionen 1878-79. In Vega-Exped.-Vetenskapliga Iakttagelser, vol. 2, pp. 383-428, pls. 27-32. Woopwick, K. H. 1953. Polydora nuchalis, a new species of polychaetous annelid from Cali- fornia. Journ. Washington Acad. Sci., vol. 43, pp. 381-384, fig. 1. U.S. GOVERNMENT PRINTING OFFICE:1966 Proceedings of the United States National Museum SMITHSONIAN INSTITUTION + WASHINGTON, D.C. Volume 120 1967 Number 3566 SUPPLEMENTARY DESCRIPTION OF THE MYODOCOPID OSTRACOD EUPHILOMEDES MULTICHELATA FROM THE GREAT BAHAMA BANK By Louis S. KorNIcKER Associate Curator, Division af Crustacea The original description of Philomedes multichelata Kornicker, 1959, was based solely on males collected in the vicinity of the Bimini Islands, Great Bahama Bank. Some details of the carapace and appendages, of value in taxonomic classification, were not included in the original description. 'To remedy these omissions, the holotype and paratypes have been restudied and described more completely in this paper. In addition, a female of the species, obtained from sedi- ment in the Bimini area, is described and illustrated. E. M. Poulsen (1962, p. 343) proposed the genus Huphilomedes for species of the subfamily Philomedinae having the following diagnostic characters: Maxilla with three endites; some of the secondary claws of the furca alternating with main claws; frontal organ not ringed, and shell without small horns at dorsal margin. Philomedes multi- chelata Kornicker, 1959, which agrees with the above diagnosis, is herein referred to Huphilomedes. I wish to thank Doctors Raymond B. Manning, Marian H. Petti- bone, Meredith L. Jones, Roger F. Cressey, and Mr. I. Gregory Sohn for reviewing the manuscript and Miss Caroline Bartlett for final preparation of the illustrations. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Family Cypridinidae Baird, 1850 Subfamily Philomedinae G. W. Miller, 1912 Genus Euphilomedes Poulsen, 1962 Philomedes (part).—Miiller, 1912.—Skogsberg, 1920. Euphilomedes Poulsen, 1962. Type spectes.—LHuphilomedes nodosa Poulsen, 1962, by subsequent designation, Kornicker (1967). GENERIC CHARACTERS.—Only a few additions are needed to Poul- sen’s diagnosis of the genus (1962, pp. 359-361) as amended by Kornicker (1967) to include E. multichelata. Sixth limb: The end joint of H. multichelata is comprised of two lobes; the proximal lobe bears a short, slender, hirsute bristle followed by three stout, plumose bristles. Seventh limb: The male of EL’. multichelata bears only five bristles on the seventh limb (4 distal, 1 lateral), the same number as reported on FE. oblonga (Juday, 1907). Euphilomedes multichelata (Kornicker, 1959) Fiaures 1-6 Philomedes multichelata Kornicker, 1959, pp. 230-232, figs. 46, 3A, B; 50, A-E; 51, A-D. Ho.toryre.—Specimen no. 686Z-6, Columbia University, Depart- ment of Geology. Gender: male. PARATYPES.—USNM 112940, 10 males. Hypotyrre.—USNM 112985, 1 female. TypE LocaLity.—Holotype and paratypes were collected at night off the end of the Lerner Marine Laboratory on North Bimini, B.W.I. The female described in this paper (USNM 112985) was from sediment approximately a mile east of the Bimini Islands. Draanosis (revised) —Euphilomedes with five primary claws and six to nine (usually eight) secondary claws. Second joint of exopodite of sixth limb bilobed, with small bristle followed by three large bristles on proximal lobe. Seventh limb of male with one lateral and four distal bristles. Anterior surface of valves with irregular polygons distributed in shingle-like pattern. DEscRIPTION OF MALE.—Shell (figs. 1, 32): oval, elongate with greatest height near middle, prominent rostrum and broad rostral incisure (figs. la, 6); anterior margin of rostrum and anteroventral margin of shell with scalloped outline formed by crescent-like marginal denticulations. Anterior surface of valves with irregular polygons distributed in shingle-like pattern (fig. 1c). Posterior dorsal margin with linear hinge depressed below outline of shell; hinge not visible NO. 3566 EUPHILOMEDES MULTICHELATA—KORNICKER 3 in lateral view except by transmitted light. Posterior hinge ele- ment of each valve consisting of angular sclerotized process (figs. le, h); medial hinge element straight; anterior hinge element not prominent. Left valve broadly overlapping right valve along an- terodorsal margin (fig. 16). Fifteen or more individual muscle scars clustered near middle of valve (fig. 1a); large scar situated some distance above others. Inner lamella broad, terminating at each end of dorsal hinge. Line of concrescence about half-way between middle and outer margin of inner lamella. Selvage with wide, corrugated, lamellar pro- longation, having fringe of slender spines along outer margin; no parallel striations observed on inner lamella. Six or seven long hairs, some bearing secondary spines, forming row on inner lamella be- hind rostrum (figs. 1d, 7); inner lamella below rostrum with small hair followed by wide space and then about four hairs (fig. 1/); about 16 hairs on posteroventral part of inner lamella (figs. le, h). Marginal pore canal with minute tapered hair between first and second crescent-like denticulation on anteroventral margin of ros- trum; another pore canal with minute hair near 12th to 15th denticle (figs. 1d, 2). Additional marginal pore canals more or less evenly spaced along ventral and posteroventral margin. Hairs with either pointed or blunt ends distributed on rostral surface (fig. 1g), some forming row near margin of shell; long slender hairs with pointed ends forming row that is closer to edge posteriorly and is near ven- tral and posteroventral margins of valves; additional hairs coming from normal pore canals; a long tapered hair projects backward from base immediately behind posterior hinge element of each valve (figs. le, h). Dimensions: Holotype length 1.04 mm, height 0.60 mm, width 0.57 mm. Length of specimens ranges between 0.96-1.04 mm (Kornicker, 1959, p. 231). First antenna (fig. 2a): First joint with about seven clusters of short hairs on lateral surface. Second joint with clusters of long hairs on medial surface; spines projecting from ventral margin and distally with a dorsal and lateral bristle; a distal ventral bristle observed only on one of five specimens examined; all bristles bare or with short hairs distally, none with wreaths of long hairs. Third joint with one ventral and two dorsal bristles, the longer dorsal bristle provided with wreath of long hairs near middle and short hairs distally; other dorsal bristle bare or with short hairs distally; ventral bristle with short hairs on most specimens. Fourth joint with two dorsal bristles, each with wreaths of long hairs, and one short and three long ventral bristles subdistally; bristles bare or with short hairs distally. Fifth joint inferred to be inserted ventrally between 4. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 fourth and sixth joints, bearing sensory bristle with broad base and provided with numerous filaments. Sixth joint bears distally a long dorsal bristle, which is bare or with short hairs distally. End joints with five slender bristles and two long stout c- and f-bristles. Second antenna (figs. 2b, c): Exopodite: first jomt elongate with small medial spine distally; second joint about one-third length of first; third joint about twice length of second; distal margins of second to eighth joints with medial comb of short spines; basal spines not observed. First joint without bristle; bristle of second joint about three-fourths length of joint, bearing short marginal spines. Bristles on joints three to eight bearing natatory hairs, without marginal spines. End joint provided with four stout bristles with natatory hairs, and two short slender bristles without hairs. Endop- odite 3-jointed; first jot with five short basal bristles and one long distal bristle, all bare, except for single spine observed near middle of long bristle on some specimens; second joint elongate, with two bristles provided with short spines (one specimen had three bristles on one appendage only); third joint elongate, arcuate, with two short bare annulate bristles and about five serrated ridges near tip (appendage which had three bristles on second joint had only one short bristle near tip of third joint). Mandible (figs. 2d—f): No coxale endite. Basale: ventral margin with four short slender bristles with short spines, and two long bristles with wreaths of long hairs. Dorsal margin with three bristles, one near middle and two at distal corner, middle bristle and one of two at distal corner bearing short spines distally. Medial surface with five short bristles proximally near ventral corner and one short bristle at middle near ventral margin. Exopodite (fig. 2e) almost reaching middle of first endopodite joint with two terminal bristles, distal about one-half length of proximal bristle, both with short hairs dis- tally on some specimens; tip of joint with blunt hirsute process. Endopodite: first joint with one bare short bristle and three long hirsute ventral bristles distally. Second joint dorsal margin with proximal group of two, and distal group of six bare bristles; ventral margin with two bare bristles distal to middle, one short annulate bristle, and two clawlike bristles subdistally. End joint (fig. 2f) with two large subequal claws, one short dorsal claw, and three bare annu- late bristles. Medial surfaces of basale and first and second joints of endopodite provided with groups of hairs. Maxilla (figs. 2g, h): Very small. Exopodite with one proximal and two distal bristles; proximal bristle bare, annulate, about one- third length of longest distal bristle; longest distal bristle with hairs, annulate, longer than combined length of second and third joints of endopodite; second distal bristle nonannulate, semitransparent, with NO. 3566 EUPHILOMEDES MULTICHELATA—KORNICKER 5 few stiff spinelike hairs, about one-half length of other distal bristle. Coxale with bare, annulate anterior bristle distally. Three small endites, each with about five bare, nonannulate, semitransparent bristles (crowding of bristles makes accurate count difficult); one bare, annulate bristle about same length as proximal bristle of exopodite, located near base of third endite. Anterodistal end of basale with long, stout, annulate bristle with long hairs; posterodistal end with bare, nonannulate, semitransparent bristle. Distal ends of first and second endopodite joints with about 14 bare, nonannulate semitrans- parent bristles; one bare, nonannulate, semitransparent bristle lo- cated medially near middle of first endopodite joint; second semi- transparent bristle located medially on distal margin. Surface of endopodite and basale with long hairs; precoxal with fringe of long hairs along anterior margin; endopodite surface with short spines. Fifth limb (figs. 3a, 6): Epipodial appendage with about 39 plumose bristles. First endite with two bare bristles, one long; second endite with one long, stout bristle and about four shorter, semitransparent bristles of varying length, all bare; third endite with one long, stout, annulate bristle provided with short hairs, and about eight bare, shorter, semitransparent bristles. First exopodite joint with about seven bare, semitransparent bristles; second exopodite joint with one broad bladelike and three or four stubby, semitransparent bris- tles, all bare; third exopodite joint with long, annulate, plumose bris- tles on outer lobe and about three bare, semitransparent bristles of unequal lengths on inner lobe; end joints with about six bristles, two longest bristles hirsute, annulate, one stout bristle also annulate, but without hairs, remaining three bare and semitransparent. Sixth limb (fig. 3c): Protopodite: first endite with one bristle, bare or with short spines; second endite with one bare, proximal bristle and three terminal bristles; third endite with six terminal bristles, three medial, three lateral; fourth endite with five bristles. Second joint of exopodite narrow with two lobes, distal lobe provided with five stout plumose bristles, proximal lobe with three stout, plumose bristles followed by one short bristle; no bristles in place of epipodial appendage. Surface with clusters of short hairs; joint separations not well marked. Seventh limb (figs. 3d, e): Cleaning bristles: four in distal group, two ventral, two dorsal, each with two to four bells; one proximal bristle with two or three bells; bristles bare or with short marginal spines. Terminal comb: fan shaped with marginal teeth (in side view teeth difficult to see, and fan gives false appearance of being single large tooth). One small fan-shaped comb with slender marginal spines on each side of terminal comb. Long slender peg with rounded end, and sharp spine opposite terminal comb. 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Copulatory organ (fig. 37): Long, slender, divided into three lobes, each hirsute with two annulate bristles; one lobe with large curved tooth. Furca (figs. 3f-h): Each lamella with 12 to 14 (usually 13) claws consisting of 5 primary and 7 to 9 secondary claws; primary claws numbers 1 and 2 followed by 3 to 5 (usually 4) secondary claws, 1 primary claw, 3 to 5 (usually 4) secondary claws, and finally 2 small primary claws; medial and lateral rows of teeth on first 3 primary claws; lateral sides of proximal 2 primary claws each with comb of TABLE 1.—Distribution of primary (P) and secondary (S) claws on furcal lamellae Claw Specimen 1 2 Shs Ole 7 8 9 10 il 12 13 14 Total 686Z-A Left ) SANS caeiaels lumi ey < ede belie cae ks ts IMS Wado ial hE) ie 13 Right|-P PS VS S'Se HPS: OSS aks aie 13 686Z-12 Left PIP cisiaS Shas Sea Sot Si es h. sonet cme 13 686Z-13 Left Pe Pe 4 Sees ib HO Seo Ste ene ees 13 Right PB. (SS S28 WS. PEE es b4 8. Jo ee ees 14 686Z-14 Left Po oP SS: Se 8 ee 88) Loon) ene 12 Right.|- bP. SoS sR SS ao eee 12 686Z-4 ? PoP SVs 48S. Pawo SS) SiS Site eae 14 686Z-B_ Left PunPor S838) SRS) Seis) sseiee eee 13 Right. Ps cP Sco. JSsdP)) | Sip Ba tS Stee 13 686Z-R_ Left PP S'S. 8) S.P S18. 1S) oS) bare 13 Right |) PPS 7S) S87 PS Se a eee 13 686Z-J Left PoP 8:48 Ss" 32 nS: iS. eS Sr ieee 13 Right | Pe cP TS 8s SebsP) 48 °Sh US So we Eeaee 13 686Z-H Left Py SP af8i88 oSei8: 42 w8e us) Sako eee 13 RightuleP ae Es) ASA Sis So eeek Sak pone 13 686Z-K Left Po iP? TS. 8 8: (8 Pes) SS) Specs 13 Right. Pi PS" S*Si 8 8) SiS) aoe eee 14 686Z-L Left PP LS" Ss 8 PS) sy 48) AS eka 13 Right (oP. Poi Sh SSS oP SS ES Ss) a eee 13 short spines extending obliquely onto lamella; secondary claws with spines along anterior and posterior margins. Primary claw number 1 with cluster of long hairs near base; lateral surface of each lamella with numerous clusters of short hairs. The distribution of primary and secondary claws on several specimens is shown in table 1. Eyes: Lateral eyes large; about 17 ommatophores visible in side view, each weakly divided by a suture into two parts (fig. 3k). Medial eye pigmented (fig. 37). Frontal organ (fig. 37): Elongate, two jointed; short spines on sur- face of end joint and distal end of basal joint. NO. 3566 EUPHILOMEDES MULTICHELATA—KORNICKER 7 DESCRIPTION OF FEMALE.—Shell (figs. 4, 5a,b): oval in lateral view, highest near middle, widest behind middle; prominent rostrum, broad rostral incisure (figs. 4a-c). Rostrum anterior margin and _ shell anteroventral margin with scalloped outline formed by crescent-like marginal denticulations of flange (figs. 4f, 5b). Posterior one-third of dorsal margin with linear hinge depressed below outline of shell (fig. 47) ; posterior hinge element of each valve with angular sclerotized process; straight medial hinge element; anterior hinge element not prominent. Left valve broadly overlaps right valve along antero- dorsal margin (fig. 4b). Numerous muscle scars in front of middle of valve, located more anterior than on male (figs. 4a, e). Shell surface with irregular polygons distributed in shingle-like pattern, anterior to muscle scars (figs. 4e, 5a). Inner lamella broad, with vestibule; line of concrescence (identified as inner margin of clear zone of inner lamella) near middle of inner lamella anteriorly and ventrally, coming closer to outer margin posteriorly (figs. 4f-h). Selvage with wide, corrugated, lamella pro- longation with fringe of slender spines along margin. Faint striations along anteroventral inner lamella. About six long hairs, bearing secondary spines, in row on inner lamella behind rostrum (figs. 4f, 50). Small tapered hair on inner lamella below rostrum (figs. 4f, 56) fol- lowed by four hairs with secondary spines. Numerous tapered hairs on posteroventral part of inner lamella (fig. 4g). Numerous radial pore canals and false radial pore canals along anterior, ventral, and pos- terior margins (figs. 4f-h). Hairs with blunt and pointed ends (fig. 4d), scattered on rostral surface; tapered hairs of various lengths in row along ventral and posteroventral margins; additional hairs in normal pore canals, scattered over shell surface. Long tapered hair projecting backward from behind posterior hinge element of each valve (fig. 49). Dimensions: Length 1.07 mm, height 0.66 mm, width 0.51 mm. First antenna (fig. 5c): First and second joints with surface hairs. Second joint with distal bristles, one dorsal, one lateral bristle. Third joint with one ventral and one dorsal bristle, dorsal bristle with short hairs. Fourth joint with two dorsal bristles and one short and two long bristles near ventrodistal corner. End joints with eight bristles. Second antenna (figs. 5d—h): Exopodite with nine joints (fig. 5e); first joint elongate without bristle; second to ninth joints trapezoidal decreasing in width distally, without basal spines. Distal margin of second joint only with comb of long spines or stiff hairs (fig. 5h). Second and third joints each with bristles with denticulation along ventral margin, no natatory hairs (fig. 5g); fourth to eighth joints with long bristles with natatory hairs; ninth joint with four bristles (fig. 5f): long bristle with natatory hairs; shorter bristles bare, one about one- 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 third length of long bristles, two extremely short. Endopodite with two joints (fig. 5d): basal joint with a long stout bristle with wreath of hairs and a short, bare annulated bristle (additional bristles normally present on this joint in subfamily were not observed); distal joint elongate with stout terminal bristle with wreath of long hairs near middle. Mandible (figs. 5i, 7): Coxal endite large, bifurcate, with rows of spines. Basale: ventral margin of left appendage with four short bristles followed by one long bristle near distal end; right appendage with two additional bristles near middle (fig. 57), one with base on medial side, other on lateral side; dorsal margin of both appendages with three bristles, one near middle and two at distal corner; medial surface with one bristle at middle near ventral margin and four shorter bristles near proximoventral corner; short spines and long hairs on ventral margin near proximal end. Exopodite short with two bristles and hairy process at tip, inner bristle with short marginal hairs. Endopodite: distal end of first joint with two long and two short bristles ventrally; dorsal margin of second joint with proximal eroup of two and distal group of about five bristles; ventral margin with two subdistal bristles and distally with one short, annulate bristle and two clawlike bristles; distal joint with two long, stout claws of subequal length and three bare, annulate bristles. Medial surface of basale and second joint of endopodite with hairs. Maxilla (figs. 54, 6a-c): Precoxa and coxa with marginal fringe of fine hair, anterior margin of coxa with one short, bare, annulate bristle. Basale with three distal bristles: anterior bristle with wreaths of long hairs, medial bristle bare, posterior bristle broken off at base of specimen examined. Anteroventral margin of first endopodite joint with annulate bristle with wreath of hairs near middle and marginal hairs. Terminal end of endopodite with 13 bristles (fig. 6b). Exopodite with two long bristles and one short bristle (fig. 5k). Three endites (fig. 6c): first endite with six bristles; second endite with four bristles; third endite with six distal bristles and one proximal bristle. Fifth limb (figs. 6d—f): Main tooth of first joint of exopodite com- prised of several constituent teeth (fig. 6d): distal tooth rectangular; following tooth longer, slender with few low secondary teeth; next tooth shorter than distal tooth, slender, pointed with few secondary teeth proximally; next tooth short, bare. Spine with row of secondary spines along dorsal margin following fourth tooth. Triangular tooth anterior to rectangular distal tooth low, curving upward. Distal margin of first joint with two centrally located bristles with wreath of stiff hairs near middle; lateral margin with low node, bearing short bristle with wreath of stiff hairs near middle (fig. 6d). Second joint with large tooth with low crenulations along inner curvature; two NO. 3566 EUPHILOMEDES MULTICHELATA—KORNICKER 9 bristles on base near posteroproximal margin, one spinose, one bare (fig. 6e). Third joint with two long, plumose bristles on outer margin and two or three on inner margin. End joints with about seven bristles. Endites with numerous bristles. Sixth limb (figs. 6g-h): Protopodite: first endite with one short, annulated, bare bristle; second endite with three terminal bristles with wreath of stiff hairs near middle, and one short, annulated, bare bristle on posterior margin; third endite with six terminal bristles, all with wreath of stiff hairs near middle; fourth endite of left sixth limb with five bristles, all with wreath of stiff hairs near middle, right sixth limb with bifurcating bristle resulting in total of six bristles on endite (fig. 6h). Second joint of exopodite with two distinct lobes; distal lobe with five bristles on left sixth limb, six on right, all with wreaths of stiff hairs; proximal lobes on both limbs with three long, stout, plumose bristles followed closely by one short, bare, annulate bristle. Seventh limb (fig. 67): Quite similar to male. Terminal comb consisting of fan with marginal teeth, in side view appearing as large single tooth. Small fan with marginal teeth located on each side of terminal comb. Slender peg and sharp spine opposite comb. Four of five distal bristles and one proximal bristle; bristles bare or with short marginal spines distally. Frontal organ (fig. 67): 2-jointed; apparent additional joints near proximal end probably resulting from wrinkles and folds. Eyes (fig. 67): Median eye well developed, pigmented; lateral eyes absent. Furca: Missing from specimen. Eggs: About five oval eggs in brood pouch. Remarxks.—LHuphilomedes multichelata is closely related to Eu- philomedes oblonga (Juday, 1907). Specimens of the latter were unavailable for study. According to the description and figure by Juday (1907, p. 145, pl. 20, fig. 6), EL. oblonga has fewer secondary claws (5 claws) on the caudal furea than FE. multichelata has. In order to determine whether the difference in number of secondary claws could be due to variability in that character, the number of secondary claws on the furca was counted on 11 specimens of £. multichelata (table 1). No specimen examined had fewer than seven secondary claws, most had eight, a few nine; consequently it is con- cluded that Z. multichelata and E. oblonga are distinct. Literature Cited Barrp, W. 1850. The natural history of the British Entomostraca, 364 pp., 36 pls. [Ostracoda: pp. 188-182, pls. 18-23.] JupDAY, CHANCEY 1907. Ostracoda of the San Diego region, 2: Littoral forms. Univ. Cali- fornia Publ. Zool., vol. 3, no. 9, pp. 135-150, pls. 18-20. KornickeEr, L. 8S. 1959. Ecology and taxonomy of recent marine ostracodes in the Bimini area, Great Bahama Bank. Publ. Inst. Mar. Sci., vol. 5 (1958), pp. 194-399, figs. 1-89. 1967. Euphilomedes arostrata, a new myodocopid ostracod from the Maldive Islands, Indian Ocean. Proc. U.S. Nat. Mus., vol. 120, no. 3563, pp. 1-21. Miu.ier, G. W. 1912. Ostracoda. Pt. 31 in Das Tierreich, 434 pp., 92 figs. PovutsENn, E. M. 1962. Cypridiniformes-Cypridinidae. Pt. 1 in Ostracoda-M yodocopa, 414 pp. [Dana-Report No. 57.] SxoasBereG, T. 1920. Studies on marine ostracods, part 1 (Cypridinids, Halocyprids, and Polycopids). Zool. Bidr. Uppsala, suppl., vol. 1, 784 pp., 153 figs. 10 NO. 3566 EUPHILOMEDES MULTICHELATA—KORNICKER 11 Ficure 1.—Euphilomedes multichelata, male: a, right lateral view, showing position of muscle spots and scalelike appearance of reticulations on anterior part of shell (holotype); b, dorsal view of shell, anterior on right (holotype); c, detail of surface pattern, anterior part, right valve from inside (686Z-12); d, medial view, anterior extremity, left valve (686Z-13); e, medial view, posterior extremity, right valve (686Z-12); f, medial view, anteroventral margin, left valve (686Z-13); g, detail of blunt, tapered hair on rostrum of right valve from inside (686Z-12); h, medial view, posterodorsal part, left valve (686Z-13); 7, medial view rostrum, right valve (686Z-13). (Same scale, in microns: a,b; c, g; Ger lty ts fe) 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 eSasee Ficure 2.—Euphilomedes multichelata, male: a, lateral view, left first antenna (686Z-12); b, medial view, right second antenna (686Z-14); c, medial view endopodite, right first antenna (686Z-13); d, lateral view, left mandible (686Z-12); ¢, medial view exopodite, left mandible (686Z-13); f, medial view, distal end, left mandible (686Z-14); g, medial view, right maxilla (686Z-13); h, medial view, exopodite of left maxilla (686Z-13). (Same scale, in microns: a, b, d; c; e-h.) NO. 3566 EUPHILOMEDES MULTICHELATA—KORNICKER 113; S LIF = is iS ea, NY 50 Ficure 3.—Euphilomedes multichelata, male: a, 5th limb (686Z-14); b, distal end, 5th limb (686Z-13); c, medial view, left 6th limb (686Z-12); d, distal part, 7th limb (686Z-14); é, distal end, 7th limb (686Z-13), f, medial view, right lamella of furca (686Z-13); g, lateral view, right lamella of furca showing surface hairs (686Z-13); h, lateral view, two proximal primary claws on left lamella of furca (686Z-13); i, copulatory organ (686Z-13); j, frontal organ and medial eye (686Z-13); k, lateral eye (686Z-14); 1, opened valves showing position of adductor muscles, left valve at top (686Z-12). (Same scale, in microns: 4,f,k; b,c,e,g—1; d,j; 1.) 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 S60 e\e-0/piaiove Ficure 4.—Euphilomedes multichelata, female (USNM 112985): a, lateral view, showing muscle scars; b, dorsal view with polygons shown on right valve; c, ventral view, anterior to right; d, three hairs from shell: long tapered hair from posterior margin, short tapered hair from radial pore canal on posterior margin, blunt hair from rostrum; e, muscle scars and portions of ornamentation in vicinity of scars, right valve from inside, anterior to left; f, medial view, anterior right valve; g, medial view, posterior right valve; h, medial view, ventral section, right valve; i, medial view, hinge section, right valve. (Same scale, in microns: a-c; d; ¢,i; f-h.) NO. 3566 EUPHILOMEDES MULTICHELATA—KORNICKER 15 2 | = o ‘2, Hh }-}\ ‘o ] ‘es ry - Ficure 5.—£uphilomedes multichelata, female (USNM 112985): a, medial view, polygons of left valve anterior to adductor muscle attachment, anterior of valve to right; b, medial view, anterior left valve; c, lateral view, right first antenna; d, endopodite, second antenna; e, medial view, right second antenna; f, detail of 8th and 9th joints of second antenna; g, detail of sections of bristles on 2nd, 3rd, and 4th joints of 2nd antenna; h, detail showing comb of stiff hairs or spines on distal margin of 2nd joint of second antenna; 7, lateral view, left mandible; 7, lateral view of basale of right mandible; k, medial view, exopodite of maxilla. (Same scale, in microns: a,d,f—h,k; b,c,1,7; e.) 16 PROCEEDINGS OF THE NATIONAL MUSEUM Steer Praccaccana Ficure 6.—Euphilomedes multichelata, female (USNM 112985): a, medial view, right maxilla; b, medial view, distal end, endopodite of maxilla; c, medial view, endites I, II, III of maxilla; d, posterior view, Ist joint of right 5th limb; e, posterior view, 2nd joint of right 5th limb; f, part of epipodial appendage of 5th limb, fine hairs shown on one bristle, other bristles similar; g, lateral view, left 6th limb; h, medial view, 4th endite, right 6th limb; 2, distal end, 7th limb; 7, medial eye and frontal organ. (Same scale, in microns: a,F,8,95 b-e,h,i.) U.S. GOVERNMENT PRINTING OFFICE; 1967 VOL. 120 Proceedings of the United States National Museum SMITHSONIAN INSTITUTION +- WASHINGTON, D.C. Volume 120 1967 Number 3567 TAXONOMY, DISTRIBUTION, AND POLYMORPHISM IN THE LABIDOCERA JOLLAE GROUP WITH REMARKS ON EVOLUTION WITHIN THE GROUP (COPEPODA: CALANOIDA) By ABRAHAM FLEMINGER ' Introduction Acceptance of the idea that the prevailing mode of speciation in sexually reproducing animals begins with genetic divergence of spatially separated stocks is widespread and rightly so. With regard to the pelagic biotope of the oceans, however, this concept is based more on faith than on documentation, although several noteworthy efforts have been published (e.g., Bowman, 1955; Brinton, 1962; Ebeling, 1962; Davies, 1963). Study of evolution in pelagic metazoans is impeded by a combination of factors, including (1) confluence of oceans and the relative obscurity of isolating barriers; (2) uncharted variability of major current systems with climatic changes over time; (3) general lack of detailed knowledge regarding the biology and range of pelagic organisms; (4) scarcity of information on the kinds and extent of morphological variation within and between populations; (5) extensive economic and logistic re- quirements for collecting oceanic species; (6) continuing prevalence of _ 1Scripps Institution of Oceanography, University of California, San Diego, Calif. 1 2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 typology in taxonomic accounts of pelagic invertebrates (cf. Simpson, 1961, pp. 45-50); and (7) paucity or absence of a fossil record for most taxa. Many of these obstacles can be lessened materially by concentrating ; upon predominantly neritic taxa. Among the planktonic copepods; there are a number of large, essentially neritic genera from which to) select. One of these, the genus Labidocera (family Pontellidae), | offers several notable advantages for examining ecological, distribu- ‘| tional, and morphological relationships between closely related con-- geners. Species are relatively large in size and can be collected ini abundance. Sexually modified anatomical features are species-- specific and remarkably well developed. Ranges, which are limited to warm-temperate to tropical latitudes, | are sharply defined and tend to coincide with major zoogeographic as; well as inshore-offshore boundaries (Fleminger 1957, 1959; Heinrich, , 1960; Voronina, 1962, 1964; Sherman, 1963, 1964). Range limits) conforming to oceanic barriers on the one hand and latitudinal | zonation on the other are relatively simple to determine and monitor? for an extended period of time. This lattermost feature confers: added advantages to the study of neritic Labidocera in the Western Hemisphere, where several distinctive species groups are found.. These species groups are adapted to coastal waters. Within a group, . the distribution and taxonomic relationships of its species constitute) a series of succeeding ranges like successive links in a chain. Taxo- nomic, morphological, and distributional relationships within and! between species groups of Labidocera found in the Western Hemisphere * are under study, one objective being to disclose patterns of evolu-- tionary consequence operating interspecifically on planktonic copepods, | Seven species of Labidocera are known to occur on the Pacific Coast: of the Americas (Giesbrecht, 1892; Esterly, 1905, 1906; Johnson, | 1935; Wilson, 1942, 1950; Sherman, 1963; Fleminger, 1964a).? Of these, L. acutifrons and L. detruncata are typically oceanic, whereas L. acuta is more closely associated with continental and insular land ! masses. Three, L. trispinosa, L. johnson, and L. jollae, have been) found chiefly in inshore waters off California and northern Mexico / while the remaining species, L. lubbockii, is known only from the type’ locality, Golfo de Guayaquil, Ecuador. In the present study wo. additional species have been found. The two new species are closely related to LZ. jollae, and the three together comprise the subject of this paper. One of the new species | is especially noteworthy in that it is conspicuously dimorphic in the: i 2Some of the records of Labidocera published by Wilson (1950) have been | omitted for reasons discussed elsewhere (Fleminger, 1965). | NO. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER 3 male sex. Following description of the three species, ranges and interspecific relationships are discussed with special regard to their bearing on evolution within the group. The genus Labidocera was established by Lubbock (1853) for a new species, L. darwinia Lubbock, collected off Argentina at latitude 38°5’ S. More than 40 species have subsequently been described. Although substantial revision of this complex is necessary, it must be delayed due to the lack of appropriate collections for study. In this report the genus Labidocera is used in the sense of Giesbrecht (1892) and refers to those pontellids with (1) one pair of dorsal sub- cuticular lenses in forehead; (2) rostrum deeply bifurcated and lacking a lense; (3) medioventral eye protuberant and extending antero- ventrad between rostral prongs; (4) first pair of swimming legs with two-segmented endopod; (5) male right antennule with at least four separate segments distal to hinge between segment 18 and fusion segment 19-21; (6) maxillule with basipod about twice the length of endite 2, maxilliped with six distinct segments. Roughly 20 percent of the species of Labidocera, including the jollae group, bear lateral head-hooks, which are obscure structures best seen in dorsal view. All are coastal in habitat. Most are restricted to the Indian and western Pacific Oceans although one is found in the northeast Atlantic and the jollae group occurs along the west coast of North America. Their phylogenetic relationships are not yet established; however, zoogeographically reasonable groups can be sorted out using segmentation of the antennule, sexually modified appendages, and female urosome segmentation. On this basis the jollae group can be regarded as being monophyletic and widely separated from the other head-hooked species. Extrapolating from ‘present day ranges, no other head-hooked Labidocera has occupied any portion of the Western Hemisphere—at least since late | Pleistocene. _ This research was partially supported by National Science Founda- tion Grants G 19417 and GB 2861. A large number of plankton ‘samples used in this study were obtained through the generous cooperation of colleagues with Scripps Institution of Oceanography, the Bureau of Commercial Fisheries, La Jolla Laboratory, and the Inter-American Tropical Tuna Commission. Especially notable contributions of coastal plankton were made by T. Matsui, S. Kato, R. H. Rosenblatt, F. H. Berry, B. Zahuranec, W. Klawe, and M. Gilmartin. I have been greatly helped with preparation of the manu- script by the thoughtful council of C. R. Stasek, who read an early draft, and by T. E. Bowman and several members of the curatorial staff of the U.S. National Museum, who read the final draft. 4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Methods MEASUREMENTS.—In general, the procedures of Fleminger (1957) were followed. All measurements were taken with an ocular microm- - eter from specimens mounted in glycerine; prosome and urosome > were measured at 62.5X magnification using a stereoscopic micro- - scope; individual segments were measured at 80 magnification { using a compound microscope. No allowance for telescoping or ° expansion at joints has been made, but damaged specimens have been omitted. Total length (TL) was routinely measured and the | prosome-urosome length ratio (PUR) was calculated. Details of © special measurments are presented in the text. All figures were : drawn with the aid of a camera lucida. Co.tiEctions.—Plankton samples used in this study were obtained by three kinds of tow. Most collections were CalCOFI (California | Cooperative Oceanic Fisheries Investigations standard plankton tow) oblique tows (140 meters to the surface), taken with a one-meter — net (1m/O; cf. Berner and Reid, 1961, or Fleminger, 1964b, for more | extensive description). Some collections were obtained from aseries ; of vertically stratified horizontal net tows taken at various depths | by one-meter, Leavitt-type, opening-closing nets (1m/H; cf. Ahlstrom, 1959, for description). Much of the coastal material from the Gulf of California was collected by standard conical nets (no. 333 Nytex mesh) roughly %m in diameter (4m). A %m conical net constructed of monel mesh (¥%m/HS) and patterned after the cone of the Gulf-III Sampler (Gehringer, 1952) was used at some stations off southern Baja California. The %m and \m/HS nets were towed within five miles of the intertidal between the surface and five meters for periods of 5 to 15 minutes depending on locality. Some collections were taken by vertical tows and are referred to in the tables by ‘‘V” placed after net type. Station locations are shown in figure 2. Morphological Terminology The abbreviations of Giesbrecht and Schmeil (1898), later repeated ' by Rose (1933), afford a practical means for dealing with calanoid morphology in the light of the unresolved problem of homologizing | it with malacostracan morphology (Illg, 1958). With widespread usage these abbreviations would simplify descriptive sections, but, unfortunately, they have not been utilized as frequently as their merits warrant. To reduce copepod morphological ambiguities, Gooding (1957, 1960) has tabulated and carefully defined terms and abbreviations frequently misused. His definitions are followed in the * present paper. It seems useful to review briefly the terminology used in this and | future papers of this series on Labidocera. The calanoid body is | | NO. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER 5 divisible into three primary tagmata, the cephalosome, the metasome, and the urosome. ‘The cephalosome consists of the cephalic somites bearing the antennules (A1), the antennae (A2), the mandibles (Mnd), the maxillules (Mx;), and the maxillae (Mx:), and the first true thoracic somite bearing the maxillipeds (Mxp). The metasome con- tains the five thoracic somites (TI-V) that bear the swimming legs (P1—-5), the last pair (P5) at least being sexually modified. The cephalosome and metasome jointly make up the prosome, i.e., all body segments anterior to the major articulation between TV and the genital segment. The urosome begins with the genital segment and contains four or less abdominal segments, the last one being the anal (perianal) segment bearing the caudal rami. The swimming legs consist of two basal segments, B1 proximal and B2 distal, together referred to as the protopodite, and two distal rami, the lateral exopod (Re) and the medial endopod (Ri). The segments of the rami are numbered in sequence from proximal to distal joint; e.g., second segment of the exopod of the fourth pair of legs would be referred to as Re2 of P4. The larger articulating elements on the segments of appendages are lateral (Se), medial (Si), or terminal (St), and are numbered proximal to distal, a new sequence beginning with each segment. Spinules and hairs (cilia) also articulate or arise from sockets. Nonarticulating structures, or those imperfectly separated from the parent element by a weak line of constriction but lacking an apparent line of flexure, are treated as a general class of processes with appropriate descriptive adjectives to denote form. The genital pore or antrum (Fahrenbach, 1962) is a cuticular-lined depression on the sternum of the genital segment into which the ovi- ducts (gonopores) open. The gonopores are incompletely shielded by a genital plate articulating along its anterior margin and possibly free along its posterior margin. Dentition of the mandibular gnathobase appears to be a useful phylogenetic tool, at least for the higher taxa of calanoid copepods. Relationships have been noted between shape and arrangement of the teeth and the kind and condition of the food observed in gut analyses of a number of different species (Fleminger, 1956; Beklemishev, 1959; Anaraku and Omori, 1963). Giesbrecht (1892, pl. 8, figs. 4, 8, 11) did not propose a formal nomenclature but sometimes numbered the teeth in a ventral to dorsal sequence. Beklemishev (1959) has employed a similar system of letters and numbers. Interfamily homologies in dentition are not established, but an empirical system for Pontellidae has been derived from a survey of pontellid genera (Fleminger, unpubl.) and is used in the descriptions that follow. For the subfamily Eupontellinae, three groups of teeth can be distinguished. Group 1 consists of the two 6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 ventralmost teeth, the apical and subapical, which are the largest, usually unicuspidate, and widely separated by a deeply cut, rounded notch. Group 2 includes the three medial teeth, which are usually bicuspidate and separated by narrower, shallower notches, and a minute intercalary tooth, which may occur between medials 1 and 2 and medials 2 and 3. Group 3, the dorsalmost teeth, are referred to as basals. They extend dorsomediad into the mouth field instead of medial like the others. Basal teeth are usually acuminate, unicus- pidate, and closely spaced; they vary in number from one to four and often present useful specific characters. A seta is also present in the subfamily Eupontellinae and is located dorsal to the basals. Several rows of scalelike accessory spines and patches of spinules occur ad- jacent to the dentition, but they have not been used in this system. Holotypes, allotypes, and reference specimens have been deposited in the U.S. National Museum (USNM). Reference specimens have also been deposited in the Marine Invertebrate Collections of Scripps Institution of Oceanography (SMIC). Labidocera jollae Esterly Fiagures 1, 10 Labidocera jollae Esterly, 1906, p. 74, pls. 9 (figs. 22, 23), 12 (figs. 50-53), 14 (figs. 89, 91).—Johnson, 1935, p. 411, pls. 3 (figs. 1-3), 4 (figs. 1, 3, 11, 18, 17- 19, 20-21), 5 (figs. 3-8) [moult stages described]. LOCALITIES, MATERIALS.—See table 1. LENGTH AND BODY PROPORTIONS.—Adult and stage V copepodite measurements, excluding caudal setae, of specimens selected at random are as follows: Prosome-Urosome Length Total Length (mm) Ratio no. mean range st. dev. median range adult 9 63 2.69 2.07-3.07 0.152 4.1:1 3.6-4.9:1 adult 60 2.26 1.93-2.61 0.121 2.9:1 2.5-3.3:1 stage V cop. 2 14 2.02 1.81—2.23 4,2:1 4.0-4.6:1 stage V cop. o& 15 1.80 1.72-1.92 Baie 3.0-3.5:1 Description.—Adult female: Overall appearance stocky, urosome short relative to prosome, latter roughly four times length of former, forehead armed with pair of lateral hooks. A1 short relative to prosome, when extended posteriad reaching articulation between TIII and TIV. Urosome two segmented; genital segment large, asymmetrical, anal segment exceedingly short. In dorsal view (fig. 1d) genital segment with two anterolateral swellings: one on right side larger, knoblike, one on left side a low boss; genital pore lateral, opening on right side. Caudal rami asymmetrical, right ramus NO. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER < larger, inflated, left ramus somewhat reduced, setation concentrated at distal end; setae of both rami relatively short, less than length of urosome, proximal portions not expanded. TIV and TV separated by a distinct suture. In dorsal view TV asymmetrical, left posterior corner larger than right, both corners strongly flared, maximum width between apex of left and right TV corners about 1.2 times larger than maximum width of preceding seoment (fig. 1d). P5 with exopod bearing two lateral denticles and one short apical spine; endopod small, imperfectly separated from B2 by weak constriction (fig. 1h). Spermatophore, in general, resembles that found in Labidocera mirabilis (Fleminger, 1957). Coupler (‘‘Koppler,’’ Heberer, 1932= accessory ornamentation, Fleminger, 1957) relatively large, cemented to ventral side of urosome; in ventral view (fig. 1f) appearing as large, somewhat rectangular plate obscuring the urosome. Left and right sides of coupler extending laterodorsad in ovate lobes that flank right and left sides of urosome (fig. le). Left lobe longer and higher than right lobe, both extending dorsad well above uppermost limit of genital segment (fig. 1g); occasionally, lobe tending to flare laterally, decreasing height relative to genital segment. Spermato- phore sac short, about as long as, and located ventral to, right caudal ramus (fig. 1g). Adult male: Head-hooks present. TIV and TV separate. TV corners symmetrical, not expanded, each side ending in a minute denticle; caudal rami comparatively long, equal to combined lengths of preceding four segments (fig. 1b); rostrum asymmetrical, right prong reduced, left prong as in female (figs. 1/7, 10). Hinged right Al (fig. 1a), in dorsal view, with segment 17 expanded anteriad in an arched ridge lightly sculptured with irregular ribbing; ridge lacking denticles. Segment 17 wider (anterior-posterior limits) than long (proximal-distal limits), mean of ratio W/L=1.190, range 1.0-1.5 (22 specimens selected at random). Segment 18 with two parallel rows of slender, closely spaced denticles; fusion segment 19-21 with single row of blunter, shorter denticles, row extending three- quarters length of segment; fusion segment 19-21 roughly twice length of segment 18. Pi with unusual asymmetry in external spines on exopod of right leg; Sel of Rel, Re2, and Re3 elongate, distal portion swollen; same spines on exopod of left lez normal (fig. Im). Right P5 (fig. 1c) with large chela; both segments comprising chela elongate, slender, crescent shaped and lacking pronounced lamelliform ridges; straight line distance between origin and apex of distal segment about equal to length of right protopodite; proximal and distal 8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 \ SCALE 05 mm figs. b,d,e,f,g 02 mm figs. c,h,J,k,m 0.1 mm figs.i, | Ficure 1.—Labidocera jollae Esterly, adult male: a, right Al, setation incomplete; J, | right lateral view, appendages incomplete; c, P5, posterior view; /, rostrum, posterior view; m, Pl, posterior view. Adult female: d, posterior half of body, dorsal view; e, posterior half of body with spermatophore, dorsal view; f, posterior half of body with spermatophore, ventral view (terminal segments of P5 omitted); g, posterior half of © body with spermatophore, right lateral view; h, P5, posterior view; 7, right mandibular gnathobase, anterior view. Penultimate stage, P5, posterior view: 7, female; &, male. NO. 3567 LABIDOCERA JOLLAE GROUP—-FLEMINGER 9 segments articulate at medialmost extension of chela; when chela closed, apex of distal segment crosses proximal segment near articula- tion with B2 between prominant ovate swelling and lateral attenuated prong. Left P5 (fig. 1c) relatively short, pediform; apical segment hirsute on medial face, bearing two short distal spines, one lateral spine at midlength. No appreciable variation was observed in the proportions of the spermatophore, the flaring of the TV corners in the female, the male P5 and the male right Al. ADDITIONAL DESCRIPTION.— Right Al of adult male with partial to complete tagmosis of the following segments: 8-9, 10-12, 13-14, 19-21, 24-25 (based on numbering system of Giesbrecht, 1892). A1 of adult female and left Al of adult male with 23 perfectly separated segments; segments 8-9 imperfectly fused, segments 24-25 perfectly fused. Gnathobase of Mnd (fig. 12) with apical and subapical teeth acuminate; median teeth 1-3 bicuspidate, each cusp pointed. Three large basal teeth present, basals 1 and 2 usually bifid, basal 3 simple, acuminate, Johnson (1935, pl. 4, fig. 11) noted only seven teeth on the mandibu- lar gnathobase of jollae; his illustration lacks one of the three basals found in the present study. He has suggested (personal communica- tion) that the third basal tooth may have been overlooked since he used relatively low magnifications to examine the mandible. Adult female with symmetrical rostrum (fig. 106), prongs robust, proximal portion expanded, in frontal view prongs separated by angular notch. Diameter of subcuticular dorsal eye lenses similar in both sexes, not exhibiting the strong sexual dimorphism common to most other species of the genus. Penultimate stage female: Urosome with three segments. Corners of TV not expanded, similar to penultimate stage male. P5 in general resembles that in adult but with relatively smaller processes (fig. 19). A1 extends to posterior end of TIT. Genital segment occasionally with precocious, more or less incompletely developed genital pore. Penultimate stage male: Urosome with four segments. Right Al with middle segments moderately expanded proximal to site of subse- quent hinge. Right P5 (fig. 1k) with B2 bearing mediodistal, knoblike protuberance regarded as the endopod precursor. Left B2 with smaller, less distinct protuberance. Right and left B1 fused, ventral margin with asymmetrical notch closer to left leg. Right exopod uniseg- mented with constriction just distal to proximalmost spinous process. Left exopod unisegmented, shorter, width in posterior view essentially constant from origin to first lateral denticle. Johnson (1935, pl. 4, fig. 20) shows endopods of penultimate stage male P5 separated from 10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 B2 by suture or marked constriction. This condition was not found in the material examined during the present study. CotoraTiIon.—Two sets of prominent carmine-colored bodies in the prosome and the red pigment cup of the ventral eye generally persist for some time in material kept in 5 percent solution of formalde- hyde in sea water. ‘The carmine-colored bodies occur in pairs lateral to the midline, the anterior pair being straplike and lying between TI and TII immediately dorsal to the sternum. The posterior pair vary in shape and in the number of lobes or particles. One clump is found on each side of the gut usually at midlength of TIII. In the adult male the left member of the TIII pair is typically absent. The cuticle is generally colorless except for a faint blue tint in the swimming legs. Alive, adults are blue green, the intensity of the hue over the entire body varying from light to heavy. The blue-green color is located under the cuticle. It is not restricted to the epidermis but appears diffused throughout the outermost muscle layers. Lateral pairs of reddish-orange chromatophores to the right and left of the midline were observed in the prosome. The chromatophores are uniformly distributed, there being in general two pairs per pair of prosomal appendages. One pair is ventrolateral and can be observed in lateral view where it is seen near the ventral margin of the pleuron. The other pair seen in ventral view is associated with the appendage’s Bil. Movement of pigment in these chromatophores was observed to be slow. About one minute was required for a chromatophore to change from the reticulate stage of total pigment dispersion to the punctate state of extreme concentration. I have observed somewhat similar chromatophores in Pontellopsis occidentalis and Labidocera acuta. There is reason to believe that pigmentation will play a useful role in studies of habitat and mate selection in Pontellidae and such data should be systematically gathered and made available. Typrs.—KEsterly (1906, p. 75) refers to two specimens of jollae, a female ‘‘taken on the surface three and one-half miles from La Jolla, June 26, 1905,” and a male collected “in the cove of La Jolla, Janaury 2, 1906,” as the types that he deposited in the collections of the University of California. None of the specimens of jollae that I found in the Esterly Collec- tion held in the Scripps Marine Invertebrate Collections were desig- nated by label as a type specimen. Collecting data on the label of one male, however, matches the collecting details presented in the original description. Some of the appendages are missing (right Al, P1-5). The same kind and number of appendages are mounted NO. 3567 LABIDOCERA JOLLAE GROUP—-FLEMINGER ial on a slide in the Esterly Collection that bears the name of the species, but lacks other distinguishing remarks. The close similarity of Ksterly’s figures of the male P5 with the position of the P5 on the slide suggests that the slide and the tubed specimen comprise the male component of Esterly’s original specimens. One of the two females found in the Esterly Collection may be the other syntype. The two females were together in the same vial, labeled “‘Labidocera jollae Esterly, 2 ad., 1158.” Tow number 1158 refers to a surface tow taken on June 29, 1906 between 0505-0520 hrs. at latitude 32°52.3’ N, longitude 117°16.4’ W (off San Diego). It is possible and even likely that only one of the two females was taken at station 1158, and that the other specimen is the missing syntype from La Jolla cove (3.5 miles off La Jolla, June 26, 1905); but there is no way of distinguishing one from the other. The male specimen is, therefore, designated the lectotype. The lectotype and one of the females are deposited in the USNM, 110762. The other female and a male specimen collected by Esterly at a later date (tow 2104, Mar. 17, 1910, 1934 hrs., lat. 32°50.6’ N, long. 117°29.2’ W) are deposited in the SMIC, XVIII-3012 and XVIII-3011, respectively. DistrRiIBUTION.—Localities and collecting data are presented in table 1. These records comprise all of the captures observed in examination of several hundred plankton samples taken in the California Current, the eastern-equatorial Pacific Ocean, and the Gulf of California (fig. 2). JL. jollae occurred chiefly in inshore samples characterized by a coastal-neritic assemblage of zooplankton. The species has not been reported from any region outside of the range established by the records in table 1. It was not found in earlier coastal water studies undertaken north of Point Conception (Esterly, 1924; Campbell, 1929a, 1929b, 1930; Johnson, 1932; Davis, 1949; Le Brasseur, 1955, 1956a, 1956b; Legare, 1957; Cameron, 1957). It is not in the Aleutians list (Anraku, 1954) nor has it been reported from the northwest Pacific (Brodsky, 1950). Turning to the eastern equatorial Pacific it is absent from both Giesbrecht’s (1895) and Wilson’s (1942, 1950) lists. The present range of L. jollae, then, apparently does not extend north of Cape Mendocino, California, nor south of Cabo San Lucas, Baja California. Within these limits it is confined to near-shore waters. Most of the geographical records fall between Point Con- ception and Sebastian Viscaino Bay (table 1, fig. 11). North of Point Conception, and extending west along the Aleutians to Asia, jollae is replaced by Epilabidocera longipedata (Sato) [= E. amphitrites 12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 (McMurrich)]. South of Magdalena Bay, Baja California, jollae is replaced by Labidocera diandra, new species, which is described below. Ficure 2.—Location of plankton samples examined for Labidocera. CalCOFI stations (parallel lines perpendicular to the coasts of California and Baja California) have been occupied regularly since 1950; from them one or more samples from each seasonal quarter of one year were examined. Labidocera kolpos, new species Figures 3-5, 10 LOCALITIES, MATERIALS.—See table 2. LENGTH AND BODY PROPORTIONS.—Adult and stage V copepodite NO. 3567 LABIDOCERA JOLLAE GROUP—-FLEMINGER 13 measurements, excluding caudal setae, of specimens selected at ran- dom are as follows: Prosome-Urosome Length Total Length (mm) Ratio no. mean range st. dev. median range adult @ 54 2.66 2.18-2.97 0.152 4.3:1 3.6—-5.0:1 adult o 61 2.10 1.91-2.30 0.089 3.0:1 2.8-3.6:1 st. V cop. ? 6 1.99 1.90—2.07 4.2:1 4.0-4.6:1 st. V cop. @ o 1.75 1.63-1.88 3.4:1 3.2-3.6:1 Draenosis.—An allopatric twin species of L. jollae, morphologically similar in general features, but showing several constant differences in sexual structure. The name kolpos [Greek] refers to the geograph- ical region (Gulf of California) inhabited by the new species. Adult female: Greatest width of TV, in dorsal view, approximately equal to greatest width of preceding segment (fig. 3f). TV corners weakly flared, maximum width between corners about 15 percent less than that in jollae of equal prosome length (fig. 4d). P5 with vestigial endopod perfectly fused to B2 (figs. 37, 1). Mandibular gnathobase with cusps of median teeth rounded at apex (fig. 3k). Spermatophore, in lateral view, with left and right lobes low, closely appressed to dorsolateral portions of genital segment, lobes not extending beyond dorsal limit of genital segment (figs. 3g-i); in jollae lobes larger and not flattened but extending conspicuously above genital segment. Rostrum as in jollae (fig. 10c). Adult male: Chela of right P5 similar in form but smaller in pro- portion to protopodite and left ramus than in jollae; shortest length of distal segment, measured along imaginary straight line between proximal and distal ends, excluding spine, equal to lengths of B2 plus half of B1 (fig. 3c); in jollae this width equal to total length of both segments of protopodite. Inner circumference of chela about half the length of that in jollae, although protopodite only slightly smaller (fig. 5). In closed chela, apex of distal segment crosses proximal segment near articulation with B2 between weak swelling and stout lateral prong. Rostrum as in jollae (figs. 3n, 10c). Right Al (fig. 3a) in dorsal view with segment 17 almost twice as long as wide, mean of ratio W/L=0.57, range .50-.64 (22 specimens), anterior edge produced in shallow ridge; ridge more weakly sculp- tured than in jollae; segment 18 with two rows of teeth as in jollae; fusion segment 19-21 with row of small denticles extending about half length of total segment; segment 19-21 about one and one-half times length of segment 18. Compared with jollae, length of segment 17 is slightly greater, length of segment 18 is slightly less, length of segment 19-21 is considerably less (figs. 4a-c). ADDITIONAL DESCRIPTION.—L. jollae and kolpos share the fol- lowing characters: Female Al and male left Al with 23 perfectly 14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 SCALE 05mm figs. b,d,e,f,g,h,i,] 0.2 mm figs. c,l,m 0.1 mm figsk,n Ficure 3.—Labidocera kolpos, new species, adult male: a, right Al, setation and dentition approximated; b, right lateral view, appendages incomplete; c, P5, posterior view; d, dorsal view; m, rostrum, posterior view. Adult female: ¢, right lateral view, appendages incomplete; f, posterior half of body, dorsal view; g, posterior half of body, dorsal view, with spermatophore; h, posterior half of body, right lateral view, with spermatophore; 1, posterior half of body, ventral view, with spermatophore; j, P5, posterior view; , right mandible, gnathobase, anterior view. Penultimate stage, P5, posterior view: J, female; m, male. NO. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER 15 separated segments, segments 8-9 imperfectly fused, segments 24-25 perfectly fused; right Al of adult male with partial to complete fusion of the following segments: 8-9, 10-12, 13-14, 19-21, 24-25. Exopod of P5 in adult female occasionally with extra lateral spinous prong as in figure 3). Right P1 of adult male with Se2 of Re3 not swollen, Sel of Rel-3 swollen as in jollae. Penultimate stage male with right leg of P5 (fig. 3m) bearing 2-3 minute, lateral denticles and one terminal prong. P5 of penultimate stage female resembles that of adult except for proportional relation- ships of segments (fig. 3/). - ° 2500 ° ° OF Or Bo [ °° Bos e@ 008 02 eee @ 000-3 % 0990-3 0°88 05 oO ce 2-920 eok2 0 e 132 a ee 100 125 150 175 150 175 .200 .225 .250 .275 300 .325 .350 .375 400 425 .450 475 .500 525 Segment |7 (mm) Segment 18 (mm) Segment 19-21 (mm) Ficure 4.—Comparisons of selected structures in Labidocera jollaeO and L. kolpos @: a-c, length of adult male right Al segments 17-21 plotted against length of prosome; d, maximum width of TV plotted against length of prosome in adult female. (Specimens chosen at random from selected localities.) 62 66 70 74 78 62 .86 Adult ¢ TY Width (mm) Cotoration.— Preserved specimens (5 percent formaldehyde solu- tion in sea water) exhibit two sets of carmine-colored structures, one between TI and TII, the other in TIII, as described above for jollae. The swimming legs also show a faint tint of blue in the cuticle. Living specimens were found to be uniformly blue-green as in ollae. Examination for chromatophores was carried out only on preserved material. The results suggest a chromatophore array similar to that in jollae but, due to fading of pigments, this should be confirmed by examination of living material. Typrs.—Female holotype, USNM 110747, allotype, USNM 110748, selected from material collected at station II-1 (lat. 28°03’ N, long. 111°16.5’ W), located a few yards offshore of a steeply sloped sand beach in a small cove, roughly 20 miles north of Guaymas, 16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Sonora, Mexico. About 100 adult specimens collected with the holo- type are deposited in the USNM, 110749. A series of reference specimens representing the known range in the Gulf of California has been deposited with SMIC XVIII-3. DistriputTion.— Along the Mexican mainland the species has been taken from Puerto Pefiasco to Isla Macapule, just south of Topolo- bampo. On the Baja California shore of the Gulf the species has been collected from San Felipe south to Bahia San Carlos, located just north of Santa Rosalia. Within these geographical limits (fig. 11) kolpos has been found most frequently and in the greatest abundance in near-shore waters. The species has not appeared in samples from the southernmost third of Baja California. It is also absent in the tows taken between Mazatlin and Ecuador. A related species, described next below, replaces kolpos in southern Baja California and along the mainland coast, at least as far south as Puerto Vallarto, Mexico. 1.88 1,80 1,72 1,64 1,56 1,48 Prosome Length: (mm) 1,40 52 60 68 76 84 92 1.00 108 LI6 124 1,32 30 38 46 P5 Chela, Inner Circumference (mm) P5 Protopodite Length (mm) Ficure 5.—Comparison of P5 in adult males of Labidocera jollae O and L. kolpos @ chosen at random: a, length of inner circumference of chela plotted against prosome length (measurements of chela made with planimeter from camera lucida sketches of posterior view, 80X magnification; circumference limits are from apex of distal segment to apex of lateral prong of proximal segment); b, length of protopodite plotted against prosome length (protopodite measured along lateral margin at 80 magnification). Labidocera diandra, new species Figures 6-10 LOCALITIES, MATERIALS.— See table 3. LENGTH AND BODY PROPORTIONS.—Adult and stage V copepodite measurements, excluding caudal setae, of specimens selected at random are as follows: NO. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER iL 7 Prosome-Urosome Length Total Length (mm) Ratio no. mean range st. dev. median range adult 9? 144 2.95 2.57-3.49 0.140 3.5:1 3.1-4.0:1 adult A 135 2.95 2.53-3.25 0.148 3.3:1 3.0-3.8:1 adult 7 B 122 2.89 2.51-3.25 0.152 3.4:1 2.9-4.1:1 st. V cop. ? 25 2.13 2.00—2.32 3.631 3.4-3.9:1 st. Vcop.A co 25 2.12 1.93-2.31 3.4:1 3.2-3.7:1 st. Vcop.B of 25 2.09 1.91-2.24 3.4:1 3.0-3.6:1 Diaenosis.— Distinguished from other species by (1) details of sexually modified structures, (2) secondary loss of head-hooks in the mature stage of both sexes, and (3) occurrence of two morphologically different kinds of adult male (morph A and morph B), the differences being concentrated in sexually modified appendages. The name diandra [Greek] refers to the two morphologically distinct kinds of males comprising this new species. Adult Al segments 24—25 fused as in genus (fig. 6d), segments 8-9 imperfectly fused as in jollae and kolpos; segments 6 and 7 perfectly separate. Forehead rounded, no head-hooks in adult, dorsal sub- cuticular eye lenses large in male, left and right lenses touching (fig. 8k); dorsal eye lenses in female small, well separated (fig. 6d). TIV and TV not fused. Rostrum as in jollae except lacking asym- metry in male (fig. 10a). Adult female: TV in dorsal view produced posteriad in subequal lobes; in lateral view lobes flanking proximal end of genital segment (figs. 6e,f), reminiscent of detruncata; robust prong ventral to right lobe and extending posterolaterad (figs. 6d,f), left side of TV without prong. Urosome asymmetrical, in dorsal view with relatively long genital segment curving to right; anal segment small; caudal rami asymmetrical, left ramus larger, second innermost seta moderately expanded in proximal portion (fig. 6d). Genital segment with robust hooked process extending from anterior portion of left side (figs. 6d,e); posterodorsal portion of genital serzment with swollen dorsal surface lightly sculptured. Genital pore displaced to right of mid- longitudinal axis (fig. 6f). P5 with apex of exopod deeply bifurcated, weakly asymmetrical, right exopod longer; one or two small, lateral denticles occasionally present on exopod; endopods asymmetrical, left endopod apically bifurcated, right endopod spinelike with minute lateral prong (fig. 65). Adult male: Both male forms essentially identical in prosome and urosome; TV corners symmetrical with simple rounded margins not flaring outward (fig. 7c). Caudal rami elongate, length exceeding combined lengths of preceding three urosomal segments (fig. 87). Forehead rounded, lacking head-hooks and crest (fig. 8h). 221-531—67—_-2 18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Right Al in dorsal view with segment 17 lacking denticulated ridge, anterior surface rounded, not flattened, and bearing small, low protuberance just proximal to midlength of segment. Segment 18 with prominent denticulated ridge; ridge extended proximad, projecting along axis of antennule and in dorsal view overlapping SCALE 05mm figs.c,d,e,f,g,h r o2mm { figs. b,i I 0.1mm \ fig.a Ficure 6.—Labidocera diandra, new species, adult female: a, right mandible, gnathobase, anterior view; b, P5, posterior view; c, cephalosome, right lateral view; d, prosome and urosome, dorsal view; ¢, posterior half of body, left side; f, posterior half of body, right side; g, posterior half of body, right side, with spermatophore; h, posterior half of body, dorsal view, with spermatophore. Penultimate copepodite: 4, P5, posterior view. No. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER 19 anterior edge of preceding segment, surface abutting segment 17 not flattened. Fusion segment 19-21 with well-developed denticu- lated ridge extending length of segment. Segment 22 with distal spiniform process about as long as segment 23, in dorsal view process somewhat appressed to front edge of segment 23. P3 and P4 asymmetrical, right leg with Sel of Rel and Re2 enlarged and extending perpendicular to axis of leg (figs. 7g, 8%); in right lateral view enlarged spines (Sel) of P3 and P4 projecting obliquely posteriad (fig. 7c); P3 with Sel of Re2 bearing large irregular lamella on distal border, Sel usually coated with amber colored material resembling resinous (cement-like) secretion occurring on females where spermatophore is fixed to urosome. Basic characteristics of P5, as in jollae and kolpos, are short left leg; long, robust right leg bearing large, ovate chela; left leg with terminal segment lacking prominent spines and processes, terminal segment roughly perpendicular to protopodite; ventral face hirsute; in general, left P5 also reminiscent of that in L. mirabilis (Fleminger, 1957, figs, 32, 33). Morph A differs from morph B in details of the right Al and chela of right P5. Morph A: Right Al with denticulated ridge of segment 18 ex- tending proximad on an elongated process that reaches distal half of fusion segment 13-14 (fig. 7a). Segments of chela slender, elon- gated; proximal segment with perpendicular digitiform process extend- ing from proximal end; process bearing knob on posterior face, ridge on anterior face. Slender distal segment (finger) opposes proximal segment (thumb), and closure of chela provided by finger crossing digitiform process of thumb (figs. 7d-h). Proximal segment of chela with large proximal seta; distal segment with three setae, one approximately one-third distance from origin of segment, second at midlength, third nearer apex. Morph B: Segment 18 of right Al with ridge extending proximad to suture between seements 16 and 17 (fig. 8a). Chela ovate, compact (figs. 8d, e, h), proximal segment roughly semicircular in lateral view and bearing a flaccid lamella with distal margin broadly notched; angle of lamella relative to segment varying from acute to perpendicu- lar. Large seta originating at base of lamella resembling similarly positioned seta in male A and suggesting that lamella homologous to rigid digitiform process on proximal segment of chela in male A. Spinule located adjacent to distal side of lamella. Distal seg- ment in ventral view somewhat lanceolate, with concave palmar surface opposing proximal segment; apex of distal segment terminating in small denticle; three setae on distal segment, one located close to 20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 origin of segment, one about one-fourth of length from origin, and one at midlength; number similar to those in male A. Terminal segment of left P5 differing slightly in proportions from male A but similar in organization and armament (figs. 8f, q). ADDITIONAL DESCRIPTION.—Cephalic appendages similar in most re- spects to Giesbrecht’s (1892) account of L. wollastoni but showing several noteworthy differences. A2 with exopod almost as robust as endopod. Figure 7.—Labidocera diandra, new species, adult male, morph A: a, right Al; , dentition of fusion segment 19-21, right Al; c, posterior half of body, right side; d, P5, posterior view; e, P5, terminal segment of left leg, ventral surface; f, P5, terminal segment of left leg, lateral view; g, P3, posterior view; h, P5, chela, anterior view. Penultimate male: 1, P5, posterior view (male B); 7, right Al, dorsal view (typical of both male forms). NO. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER PAL A es SCALE 05 mm figs. b,c,j,k,| 0.5 mm figs. a,i 0.2 mm figs. d,e,f,g,h Ficure 8.—Labidocera diandra, new species, adult male, morph B: a, right Al, dorsal! view; b, posterior half of body, right side; c, cephalosome, right side; d, P5, posterior view; e, P5, chela, lateral view; f, P5, left leg, lateral view; g, P5, terminal segment of left leg, ventral surface; h, P5, chela, medial view; i, P3, posterior view; 7, posterior half of body, dorsal view; k, forehead, dorsal view. Penultimate stage: 1, forehead, dorsal view. 22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Mandibular gnathobase with eight teeth, apical and subapical teeth typical, median teeth 1-3 biscuspidate, basal teeth 1 and 2 usually bicuspidate, occasionally with single cusp as in basal 3; basal seta well developed (fig. 6a). Mx, with B2 bearing clump of three setae followed by clump of two setae on inner margin (wollastoni has two setae in each clump), lobe 3 with four setae. Large, spinelike setae of Mx bearing about 1% times more pairs of perpendicular spinules, distance between spinules less than in wollastont. Mxp with B2 bearing row of denticles as in all species of Labidocera examined. G . / / } Sy Figure 9.—Labidocera diandra, stage V copepodites: a, female, posterior half of body, right lateral view; b, male, morph B, posterior half of body, right lateral view; c, male, morph A, posterior half of body, right lateral view; d, female, P5, posterior view; ¢, male, morph B, P5, posterior view; f, male, morph A, P5, posterior view. Stage IV copepodites, P5, posterior view: g, female; h, male. In adult female, tissue within right TV corner and ventral prong eranular; lobe flanking genital segment with large clear space. Tissues underlying tergum of genital segment also glandular in appearance. Spermatophore in diandra essentially similar to that in jollae and kolpos but differing in several fundamental characteristics from that in other coastal species of east Pacific Labidocera (Fleminger, 1964a). NO. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER 23 Coupler in jollae and kolpos consists of shield ventral to urosome; right and left portions of shield turned dorsad flanking urosome and extending upward as more or less free, curved, lobiform or lamelliform processes. In diandra, right and left portions of shield appressed to genital segment. Sac in jollae and kolpos remarkably short and arising from left side of coupler; separation of coupler and sac poorly demarcated; beginning at genital pore, neck extending to left, ventral to genital segment, and posterior along left side of coupler before expanding into short sac. In diandra, beginning at genital pore, neck extending to left side ventral to genital sezment and turning posteriad, as in jollae and kolpos, but then extending dorsad and to WW A WW Ficure 10.—Front view of rostrum (top, females; bottom, males): a, Labidocera diandra (A, B); b, L. jollae; c, L. kolpos. a 2 OL 2M A right above anal segment; elongated sac appearing over right caudal ramus, and extending posteriad before abruptly turning about 180° in horizontal plane and extending toward prosome (figs. 6g,h); about three-fourths of sac length projecting forward toward right corner of TY. Flexure of spermatophore sac projecting it away from usual pos- teriorly directed attitude occurs in several other species of Labidocera. In detruncata, neck leaves coupler middorsal on genital segment, broadening into sac and turning horizontally counter-clockwise about 150°, distal portion of sac extending to right more or less perpendicular to longitudinal axis of urosome. In acutifrons, neck bending abruptly about 180° under left caudal ramus and sac projecting anteroventrad to left of urosome. In minuta, neck extending to left of midventral 24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 genital pore, turning posteriad and crossing ventral to anal segment to right side of urosome; portion of sac free of coupler extending anterolaterad. Two types of spermatophore placement observed in diandra: type 1 placement considered normal; type 2 placement less common and appears to be nonfunctional. In type 1, described above (figs. 6g, h), coupler envelops genital segment posterior to hook on left side and extends back to cover anal segment. Actual site of attachment of neck not observed but appears to emanate from genital pore. Type 2 placement with spermatophore similar to that in type 1, but coupler cemented to left corner of TV, 180° bend in sac anterior to terminal portion of sac which extends posteriad; resulting orienta- tion of spermatophore sac and coupler 180° out of phase with type 1 placement. Neck cemented to left corner of TV, totally lacking contact with genital pore. Co.toraTion.—Living and preserved specimens (5 percent formal- dehyde solution in sea water) exhibit the same sets of carmine struc- tures between TI and TII and in TIII as described above for jollae. The cuticle on the ventral side of the prosome and on the swimming legs is tinted blue. Otherwise the species is translucent and shows no pronounced coloration. Typrs.—Female holotype, USNM 110758, allotype, USNM 110759, selected from material collected at station F—5 (lat. 24°14’ N, long. 111°18’ W), bottom depth about 20 m, located roughly 1 mile offshore of Boca Flor de Malba, Baja California, an estero flanked on either side by moderately sloping sandy beach coastline. Additional refer- ence specimens are USNM 110760 and SMIC XVIII-2. DistriputTion.—Plankton samples containing L. diandra (table 3) were collected in coastal waters usually within 1 mile of shore. The northernmost positive station (F—5) is located on the Pacific coast of Baja California just south of Almejas Bay (24°20’ N). Along the Gulf of California coast of Baja California, the species occurred as far north as Santa Rosalia (27°19’ N). From the eastern shore on the Mexican mainland we have four records, all coastal and from the region between Banderas Bay (20°26’ N) and Tenacatita Bay (19° 15.3’ N). The distribution of L. diandra is shown in figure 11. During portions of the year the range will probably be found to extend farther north (approximately a hundred miles) on both shores of the Gulf. The only remaining broad question of range concerns how far south the species extends along the Central American coast. In general, the calanoids found in samples containing diandra were a mixed assemblage of coastal, neritic, and oceanic species which have a wide distribution in the eastern Equatorial Pacific. Samples rich in diandra were also rich in the coastal and neritic species of this NO. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER 25 assemblage, but the actual composition varied somewhat from tow to tow. Perhaps the only consistent feature is the relatively high abun- dance of inshore coastal species such as Acartia tonsa. Most in- teresting is the observed trend in the occurrence of other species of 125° 120° 115° 110° 105° 40° 40° © Labidocera jollae ¢ Labidocera diandra 4 Labidocera kolpos Sor 30° 25° 20° 125° 120° 115° 110° 105° Ficure 11.—Location of plankton samples containing species of the jollae group; no sample yielded more than one species. °6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 Labidocera. Within north-south limits of the diandra range, L. detruncata was very rare and decidedly offshore; L. acutifrons was abundant chiefly in the more offshore tows from blue, highly trans- parent water; and L. acuta was the numerically dominant species of the genus in blue to bluish-green water especially within 10-20 miles of the coast. Wherever the latter two species were abundant, diandra was rare or absent. The abundant catches of diandva were made in green to greenish-blue, relatively turbid water. REMARKS ON IDENTIFICATION.—Adults of diandra are distinguished from other species, except jollae and kolpos, by (1) partial fusion of segments 8 and 9 and separation of 6 and 7 in the nongeniculated A1; and (2) the short robust rostrum in which the base of each prong is swollen in lateral and frontal views. Adult diandra are easily separated from kolpos and jollae by the absence of head-hooks. The male is further characterized by (1) the terminal segment of the left P5 lacking prominent spines, hooks, or modified setae; (2) the right Al with segment 18 bearing a proximal process overlapping at least the entire length of segment 17, superficially as in L. aestiva, but differing in that abutting surfaces of the process and segment 17 are rounded and the joint separating segment 17 from 18 is not hinged as in aestiva superspecies (for aestiva superspecies, see Giesbrecht, 1892, pl. 23, figs. 3, 4); and (3) the asymmetry in P3 and P4 of Sel on Re2. In the female, excellent identifying features are found in dorsal view in (1) the combination of the lateral spinous process extending laterad from the right side of TV; and (2) the lateral, hooklike process extending from the left side of the genital segment. Remarks on Spermatophore Placement and Mating About 36 percent of adult females carried spermatophores (table 4). A number of females carried more than one, arranged in a variety of combinations of type 1 and type 2 placements. Some carried more than one spermatophore of the same placement, the couplers being arranged in sequence, one on top of the other, the sacs lying parallel to one another. Type 1 was the most frequent and geographically widespread spermatophore placement observed. In this position the neck is in close proximity to the genital pore although the actual point of con- tact is obscured by the sheath. The less frequent type 2 occurred sporadically and the mouth of the neck is not in contact or even in close proximity to the segment, the coupler and neck being cemented to the left corner of TV. In the material at hand, the frequency of type 1 spermatophore placement was more than three times that of type 2, and the latter, NO. 3567 LABIDOCERA JOLLAE GROUP—-FLEMINGER 27 when present, usually occurrred in conjunction with a type 1; ie., most specimens with a type 2 placement also carried another sperma- tophore in the type 1 position. Type 1 is, therefore, considered to be normal placement on the basis of higher frequency, more wide- spread occurrence, and the reasonable assumption that the opening to the neck of the spermatophore must be closely associated with the genital pore to provide the female with sperm. The Punta Frailes sample (table 5) is distinguished by a high pro- portion of females bearing more than one spermatophore. In this sample, examined in toto, males outnumbered females 1.2 to 1 and 80 percent of females carried one or more spermatophores. This percentage is unusually high in my experience and suggests the time of sampling coincided with a local mating swarm. Sexual swarming, if this in fact occurred, and an excess of males over females, could be the basis for the large number of misplaced spermatophores. In any event, comparison of placement types 1 and 2 provides some insight into the question of sexual behavior in the genus Labidocera. Of primary significance in spermatophore attachment in the Punta Frailes sample is the suggestion of instinctive ritualistic behavior which seems to have a high degree of species-specificity. The com- plexly arranged coupler is cemented to the female genital segment with such precision that spermatophore position, in addition to form, provides a valid criterion for identifying the species. The fact that type 2 placement is the reciprocal of type 1 can be interpreted as evidence that the male accomplishes coupler placement relative to his position on the female. Since the normal type 1 placement is much more frequent and since only one type of alternative placement has been seen, it can be inferred that the male positions himself in a specific manner relative to the orientation of the female. The pre- dominant co-occurrence of type 1 placements on females bearing type 2 placements suggests further that the former type either deters sub- sequent males from assuming a type 1 orientation or that a second male grasped a female on the side opposite the normal position already occupied by another male. Instances of two or more spermatophores of one placement type superimposed on one another could be the product of a single male that fails to release the female after cementing the first spermatophore or the outcome of subsequent matings by other males. Abnormal placement of spermatophores in other species of the jollae group were not observed. In all, spermatophores have been examined in 15 species found in the Western Hemisphere and an additional number from Africa and Asian coastal waters (Fleminger, unpubl.). Each species was found to have its own characteristic coupler—sac morphology and placement. One notable point of interest is the fact that the greater the extent of 28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 range overlap between species, the stronger the interspecific differences in spermatophore morphology. For example in the jollae group, diandra, potentially in contact with both jollae and kolpos, has a surprisingly unique spermatophore, whereas the latter two are al- lopatric and show only weak differences in spermatophore morphology. Complexity and species-specificity in spermatophore morphology and constancy in its orientation with respect to the female urosome are compelling reasons to anticipate ritualistic mating behavior in the genus. Labidocera jollae has been observed briefly in copula on two occasions, once in January 1962 (pers. comm., Dr. Bui Thi Lang, SIO) and again in April 1964 (by the author). At first the two individuals were side by side. They quickly changed positions, pivoting about the region of the genital segment like the hands of a clock and causing the prosomes to be pointed in opposing directions while the urosomes crossed at about the level of the genital segments. Clasping lasted for at least 10 minutes. In both instances the observers were unable to note whether a spermatophore was deposited. More extensive observations on copepod mating behavior, however, are available for other genera. Hill and Coker (1930) provide considerable evidence of ritual asso- ciated with copulation as well as a critical time for spermatophore deposition derived from observations on mating in several species of freshwater cyclopoids. They found that, regardless of which female structure was initially grasped, the male would later shift its position until it had secured itself to the female’s fourth pair of swimming legs; furthermore, in several species, adult males remained close to or clasped alate stage V female and waited for ecdysis, thus guaranteeing that the spermatophore would be implanted on unhardened cuticle. Fahrenbach (1962) found a pronounced mating ritual in the algal- dwelling harpacticoid Diarthrodes cystoecus, while Gauld’s (1957) brief notes on mating in some estuarine-coastal planktonic calanoids (Centropages, Temora, Eurytemora, and Acartia) also suggest ritual prior to attachment of spermatophore and uniformity of orientation of the male during attachment. In Pseudodiaptomus euryhalinus and P. coronatus, copula persists for hours and even days, whereas in Acartia tonsa the act is apparently accomplished in less than a minute (Johnson, 1948; Jacobs, 1961). Jacobs (ibid., p. 445) observed that males were more prone to attack “when they chanced to become situated some 5 mm obliquely behind a female, leaving the impression that the attack might be triggered by the female feeding current.” Johnson (op. cit.) noted that when isolated Pseudodiaptomus males and females are brought together a high percentage of the animals may copulate within a few minutes. NO. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER 29 The suggestion that the female feeding current triggers clasping by the Pseudodiaptomus male is not without merit; however, cephalic appendages appear to beat at relatively low frequencies. Under- water, low-frequency signals are probably subject to serious limita- tions for directionality and for separation from background noise. Such signals could be effective only over relatively short distances. Furthermore, no special organs of hearing are known in Crustacea (Cohen and Dijkgraaf, 1961). Taking note of the commonness of cuticular pores overlying small epidermal glands, the abundance of aesthetasks on the Al and the more extensive development of the latter in adult males of many families, it seems likely that mate recognition, pursuit and receptivity, the mating attack and locating of the female genital pore, etc., are mediated by chemical trail substances, i.e., pheromones. Feeding and swimming currents would serve to disperse these substances. Parker (1902), working on Labidocera aestiva, discusses several simple experiments which support the idea that the male finds the female by means of pheromones. Polymorphism in Labidocera diandra IMMATURE COPEPODITE STAGES Heap-Hooxs.—Head-hooks of the usual form were seen in all immature copepodites (fig. 82) from stages II to V. In copepodite stage I, found only in collections in and around La Paz Harbor, Baja California, in April 1962, both head-hooks and rostrum were absent. Except for the absence of head-hooks in adult diandra these findings agree with Johnson’s (1935) account of rostrum and head-hooks in the ontogeny of L. jollae. Regarding the ontogeny of other species of Labidocera, head-hooks are lacking in all stages of L. trispinosa, a fact established by Johnson (1935) and verified during the present study. For comparative purposes, I have also examined immature copepodite stages of other species of Labidocera at my disposal (table 6). No instance was found that paralleled the secondary loss of head-hooks* seen in 3 J have found two specimens of Labidocera in collections from Mombasa, Kenya (Lusiad Expedition, lat. 4°04’ S, long. 39°39’ E, VII-24-1962, tow 1h 1824-1829 hrs., tow 2, 1852-1859 hrs., % m net, 3-0 m depth) that agree in all particulars with L. kroyeri except that head-hooks are lacking. No younger copepodites, males or normal kroyeri females were present in the two samples. I have seen normal kroyeri in a collection from Ceylon (Lusiad Expedition, VIII-28-1962, 1807-1817 hrs., 1 m net, 3-0 m depth, in Colombo, Ceylon Harbor). Brady (1915) included kroyeri in a list of copepods from Durban Bay, South Africa, but failed to include notes on its appearance. 30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 diandra, but it should be pointed out that the list in table 6 represents less than half of the described species. The presence or absence of head-hooks in pontellids, and especially in Labidocera, has been widely regarded as a feature of considerable taxonomic significance. Obviously, if head-hooks can vary in oc- currence intraspecifically, their usefulness to phylogeny hinges upon a thorough survey of all species in the genus. Head-hooks are found in other pontellid genera (Anomalocera, Pontella, Calanopia, Ivellopsis, Epilabidocera) but not necessarily in all species of these genera. They are characteristically absent in two other genera (Pontellopsis, Pontellina). Based on this distribution, head-hooks may be regarded as an older generalized feature that is losing adaptive value in some of the younger phyletic lines. Certainly, revision of Labidocera is Labidocera diandra, co’, St. Y C = | individual station F-5 ST Pe Ue eT ST TT ae See a ee a ea ie eel el eee ee aS po station F-2 Noy alee Ts Vie elber i cI e (Ee pec aT of 1 AA SA AL SUS 186 1:94 2,02 32:10) 218 226 I | ll eal eGo, 4S) s53lbe 57) 6 65! 1,70 1.78 390 430 .470 .510 550 590 630 670 173 181 489 197 205 213 221 229 Right P5 Length (mm) TL (mm) Ficure 12.—Size distribution of right leg 5 (P5) and total length (TL) in male Labidocera diandra, stage V copepodites, collected at stations F2 and F5 (horizontal hatching= morph B males; heavy line=morph A males; specimens selected at random and measured at 60X magnification). not merely a matter of grouping species according to (1) the presence or absence of head-hooks in adults and (2) urosomal segmentation in adult females, as suggested by Sewell (1956). Not only are head- hooks undependable but the urosome is variously two and three segmented in head-hooked and nonhead-hooked species alike. No. 3567 LABIDOCERA JOLLAE GROUP—FLEMINGER 31 P5.—In the stage V female the P5 exopod (figs. 67, 9d) bears three short, lateral, spinulose processes. ‘The apex is spiniform and the left leg is slightly smaller than the right. The female P5 increases uni- formly with subsequent moults, following appearance of the ap- pendage in stage IV (figs. 9g, 14); lateral spinous processes on the exopod disappear in stage VI (fig. 65). LABIDOCERA DIANDRA STAGE ZW @& 2.34 zgeiO) 2.26 Bae m Ne on) 2.14 ZNO 2.06 Total Length (m 2.02 1.98 1.94 1.90 043 O47 O5| @55 @59 063 0.67 Length (mm) Right P5 Figure 13.—Scatter diagram comparing length of right leg 5 and total length in male Labidocera diandra, stage V copepodites (measurements taken from lateral view at 62.5 X magnification, using undissected specimens, appendage length extending from line of articulation of protopodite with tergite to apex of ramus; specimens selected at random from sample FS). The stage V male P5 (fig. 727) resembles that found in jollae in the arrangement of spinous processes and basic proportions. It differs in lacking rudimentary endopodal lobes and in having a knoblike pro- trusion on the median side of the left ramus. Stage V males can be separated into two groups (morph 1 and morph 2) based on length of P5 (figs. 9e,f). This division, into a longer and 32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 120 a shorter length, occurs irrespective of the individual’s TL (figs. 12, 13, table 7). TL distribution among the two morphs differ only slightly and in the same direction and degree as morph A differs from morph B in adult males (table 8). P85 size distribution in stage IV males does not appear to be bimodal (fig. 14). rere 3.0 2.8 2.6 eaten pales be peestesteaeer 2.4 + ice) @ aie ‘ 3 P5, Chela, Length of arr a Distal Segment (mm) One 2.0 Total Length, (mm) < . SS intersexes © O oO oO oO [e) (©) q i) T, Ta) © S P5 Length (mm) Ficure 14.—Labidocera diandra: at left scatter diagram comparing length of right leg 5 to total length in male A and female @, stages IV and V copepodites and stage VI female; in upper right, scatter diagram comparing length of distal segments of chela with total length in stage VI male (chela measured by planimeter from camera lucida drawing of posterior view; segment length determined along inner perimeter relative to mouth of chela; specimens selected at random from station F2). In the moult to stage VI, the terminal segment of right P5 divides to form the chela, the proximal segment becoming the thumb, the distal segment the finger. Assuming that the lateral spinous processes on this leg in stage V are homologous to setous-like spines in the adult chela, the proximal segment in the adult appears to be a fusion of Rel and Re2, while the distal segment comprises Re3. Immature stages of female and male P5 are compared in figure 9. NO. 3567 LABIDOCERA JOLLAE GROUP—-FLEMINGER 303 Oo aL A 3:0 A | Vice te CR . Ow Be ' ' ! lanai catia Al NK | 2.8 ! | ! ! a anes Rigi oe ! | a eae a ! 1 | ! A ! i nega — | A 1 AGES — cana team 7 = Aine ! as ! es 1 A ; Nh ee Ly = Nae ny > eS EOS = St 7 4 Sra?! Ss oO HK 22 aa a \ (OM | ae | 1 ! i] PAG ah Ubi esa! Mas ! ea ew A ! fie bene ee
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