PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 121(1):72–84. 2008. New genera in the family Sergestidae (Crustacea: : Penaeidea)

David C. Judkins and Brian Kensley (DCJ) 3132 NE 51st Ave Portland, Oregon 97213, U.S.A., e-mail: [email protected]; (BK, deceased), Department of Invertebrate Zoology, National Museum of Natural History, Washington, D.C. 20013-7012, U.S.A.

Abstract.—The sergestid genus Sergestes is restricted in definition, and five new genera erected: Allosergestes, Deosergestes, Eusergestes, Neoser- gestes,andParasergestes. These are defined in terms of 23 morphological characters, including features of the appendages, luminescent organs, and petasmata. The species belonging to each are listed, and the type species for each designated. Diagnoses for the remaining genera in the family, , Peisos, Petalidium, Sergia and Sicyonella are supplied. A key to the 11 currently recognized sergestid genera is provided.

The meso- and bathypelagic shrimp generally agreed upon (see Pe´rez Far- genus Sergestes H. Milne-Edwards, 1830, fante, & Kensley 1997). It has long been has proven to be speciose and diverse in recognized that both ‘‘genera’’ could be the world’s oceans. In 1860, Stimpson divided into groups of closely related applied the name Sergia to a late masti- species [e.g., Yaldwyn (1957), Foxton gopus sergestid larva, which Burkenroad (1972), Judkins (1978), Vereshchaka (1945) determined belonged to that group (2000)], but apart from the unpublished of species of Sergestes generally placed in doctoral dissertation of Judkins’ (1972), the subgenus Sergia.EventhoughOrt- this division was never formally pro- mann (1893) used Sergia as a full genus, posed. until fairly recently most authors regard- The present paper formalizes the sepa- ed Sergestes as having two large subgen- ration of five new genera from Sergestes era, Sergestes and Sergia. Even Yald- as proposed by Judkins (1972). The wyn’s (1957) landmark paper on New creation of these genera is based primarily Zealand sergestids maintained this posi- on the 16 characters used by Judkins tion. The reluctance to elevate the sub- (1972, Table 1), but with some additional genera was perhaps due to their sharing ones. In some cases, e.g., the relatively two important characters: in both, pereo- massive third maxillipeds of three of the pods 4 and 5 exhibit some reduction, new genera, which are clearly a major possessing only 6 podomeres, and both modification related to feeding, the po- bear well-developed gills above pereopod larity seems obvious. For others, howev- 4. Omori (1974) finally re-introduced er, e.g., the setation of the propodus and Sergia as a full genus on the basis of dactylus of pereopod 5, the polarity differences in the luminescent organs. remains uncertain. This division of the genus Sergestes into The division of the opaque-bodied two genera, the transparent-bodied Ser- Sergia group of species, all of which lack gestes s.l., in which the gastrohepatic organs of Pesta, presents more complex gland is modified to form the luminescent problems. Sergia species may or may not organs of Pesta, and the opaque-bodied possess dermal photophores and, when Sergia, which lack organs of Pesta, is now present, these may or may not be VOLUME 121, NUMBER 1 73 equipped with a cuticular lens, but the lensless type], it can be envisioned that otherwise the genus is much more uni- the lens-bearing type could have arisen form in morphology of non-reproductive from the lens-less type on more than one appendages than species in the genera occasion. On the assumption that the separated from the former Sergestes. characters of the petasma, reflecting spe- Judkins (1972) proposed that the three cies isolating mechanisms closely linked opaque-bodied species groups recognized with species-specific mating, are more by Yaldwyn (1957) be elevated to genera: reliable indicators of relationships than Sergia (having lens-less photophores), a photophores or integumental consistency, second genus having photophores with Judkins’ (1972) division of Sergia must lenses, and a third genus having a soft be rejected. sometimes fragile integument and lacking For each genus, the type species with its discernable photophores. However, com- type locality is provided. The latter will parison of the petasmata in Sergia reveals become important should some of the several groupings that are not congruent species currently thought to have an with the photophore/integument-based almost cosmopolitan distribution in the divisions. Vereshchaka (2000) divided oceans, prove to be complexes of cryptic, the species of Sergia into nine groups or closely related species, as in the caridean isolated species based primarily on pho- Acanthephyra purpurea group elucidated tophores and the petasma, but also on by Kemp (1939). A list of species belong- the hepatic tubercle/spine, the ocular ing to each of the genera is provided. In a papilla, the endopod of the first maxil- few cases, due to poor descriptions or lack liped, and the posterior branchial lobe of material, these placements may be on somite XII. Species of Vereshchaka’s inaccurate; only good fresh material will S. gardineri, S. phorca and S. robusta solve these problems. groups possess lens-less phophotophores, The genus , although at an early and his S. prehensilis, S. challengeri,and stage placed in a separate family (Lucifer- S. lucens groups have lensed photo- idae Dana, 1850), has often been treated phores. The isolated species S. tenuiremis as a member of the Sergestidae. However, and S. inoa, and species of the S. japonica in a key to the dendrobranchiate families group all lack photophores but do not and genera, Burkenroad (1983:283) uses appear to be a monophyletic primitive such a powerful group of synapomor- group. In some cases the absence of phies to define the Luciferidae as to photophores may be the result of loss in leave no doubt regarding the validity of response to environmental conditions. this family (also see Pe´rez Farfante, & The occurrence of lensed photophores in Kensley 1997.) the S. prehensilis, S. challengeri,andS. lucens groups and the similar epibenthic New and Restricted Genera of Sergestes habits of most of their species (Veresh- sensu lato chaka 1994, 2000) suggest a common ancestry. However, the petasma bauplan Allosergestes, new genus of the S. lucens group is very different Fig. 1A from that of the S. prehensilis and S. Diagnosis.—Cuticle semitransparent; challengeri groups, which are closer in photophores absent. Carapace with small petasma plan to species with lens-less hepatic and supraorbital spines present. photophores or no photophores. Given Organs of Pesta present, antero- and the similarity of structure of the two types posterolateral organs spheroid. Ocular of photophores [see Terao (1917) for lens- tubercle present. Antennular peduncle bearing photophores, Dennell (1940) for with first and third articles subequal in 74 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 1. Petasmata of selected species of genera formally in Sergestes sensu lato. A, Allosergestes sargassi; B, Deosergestes curvatus;C,Eusergestes arcticus;D,Neosergestes edwardsii;E,Parasergestes armatus;F, Sergestes atlanticus. c - capitulum; la - lobus armatus; lc - lobus connectens; li - lobus inermis; lt - lobus terminalis; pu - processus uncifer; pv - processus ventralis. VOLUME 121, NUMBER 1 75 length; stylocerite absent. Maxillule with 7 articles. Ischium of pereopods 1 and 2 palp. Maxilliped 1 with palp. Maxilliped 3 bearing spine. Fingers of chela of pereo- much longer than pereopod 3; dactylus pod 2 unequal. Pereopod 3, coxa having at consisting of 5 articles. Ischium of pereo- least one mesial tooth; posterior arthro- pods 1 and 2 bearing spine. Fingers of branchia above pereopod 3 well devel- chela of pereopod 2 unequal. Pereopod 3, oped. Pereopods 4 and 5 each having 6 posterior arthrobranchia lamellar. Pereo- podomeres. Branchiae present above pe- pods 4 and 5 each having 6 podomeres. reopod 4. Two distal podomeres of Branchiae present above pereopod 4. Two pereopod 5 setose on both margins. distal podomeres of pereopod 5 setose on Petasma with pars media elongate; pro- both margins. Petasma with pars media cessus ventralis distally expanded, bearing elongate; processes somewhat elongate, fringe of spines; lobus armatus strong, consisting of lobus inermis, lobus termi- usually straight, occasionally curved, nalis, lobus armatus, and processus ven- bearing several hooks; distalmost lobus tralis; processus ventralis distally expand- inermis, lobus terminalis, and proximal ed, fringed, or bearing stellate spines. lobus connectens characteristically in tri- Appendix masculina of pleopod 2 with fid arrangement. Appendix masculina of proximalmost spines on medial margin pleopod 2 with proximalmost spines on much longer than those immediately medial margin much longer than those following. Outer margin of lateral uropo- immediately following. Outer margin of dal ramus setose for distal 60–80% of lateral uropodal ramus setose for distal length, lacking tooth. 60–80% of length, lacking tooth. Type species.—By present designation, Type species.—By present designation, Sergestes sargassi Ortmann, 1893. Sergestes curvatus Crosnier & Forest, Type locality.—Florida Current, Sar- 1973. gasso Sea. Type locality.—South-west Indian Ocean Etymology.—Derived from the Greek off South Africa, 35u429S, 24u409E, ‘allos’ meaning other, of another kind or 500 m. race, plus the root generic epithet ‘ser- Etymology.—Derived from the Greek gestes’. ‘deo’ meaning to bind, plus the root Species.—A. index (Burkenroad, 1940); generic epithet ‘sergestes’. A. nudus (Illig, 1914); A. pectinatus (Sund, Species.—D. coalitus (Burkenroad, 1920); A. pestafer (Burkenroad, 1937); A. 1940); D. corniculum (Krøyer, 1855); D. sargassi (Ortmann, 1893); A. verpus (Bur- curvatus Crosnier & Forest, 1973; D. kenroad, 1940). disjunctus (Burkenroad, 1940); D. erectus (Burkenroad, 1940); D. henseni (Ortmann, Deosergestes, new genus 1893); D. nipponensis (Yokoya, 1933); D. Fig. 1B paraseminudus (Crosnier & Forest, 1973); Diagnosis.—Cuticle semitransparent; D. pediformis (Crosnier & Forest, 1973); D. photophores absent. Carapace with he- rubroguttatus (Wood-Mason, 1891); D. patic spine present, supraorbital spine seminudus (Hansen, 1919). present or absent. Organs of Pesta pre- sent, anterolateral organs lobelike, poste- Eusergestes, new genus rior organs fringelike. Ocular tubercle Fig. 1C present. Antennular peduncle with first Diagnosis.—Cuticle semitransparent; article much longer than third; stylocerite photophores absent. Carapace with he- mobile. Maxillule with palp. Maxilliped 1 patic and supraorbital spines present. with palp. Maxilliped 3 subequal in length Organs of Pesta present, anterolateral to pereopod 3; dactylus consisting of 6 or organs lobe-like, posterior organs fringe- 76 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON like. Ocular tubercle absent. Antennular spine. Fingers of chela of pereopod 2 peduncle with first article much longer subequal. Pereopod 3, coxa lacking mesial than third; stylocerite mobile. Maxillule tooth; posterior arthrobranchia above with palp. Maxilliped 1 with palp. Max- pereopod 3 lamellar. Pereopods 4 and 5 illiped 3 subequal in length to pereopod 3; each having 6 podomeres. Branchiae dactylus consisting of 6 articles. Ischium present above pereopod 4. Two distal of pereopods 1 and 2 bearing spine. podomeres of pereopod 5 setose only on Fingers of chela of pereopod 2 subequal. posterior margin. Petasma with lobes and Pereopod 3, coxa with mesial tooth; processes short, rounded, especially lobus posterior arthrobranchia above pereopod connectens and lobus armatus; pars media 3 well developed. Pereopods 4 and 5 each broad; processus ventralis apically acute. having 6 podomeres. Branchiae present Appendix masculina of pleopod 2 with above pereopod 4. Two distal podomeres spines gradually decreasing in size apical- of pereopod 5 setose only on posterior ly. Outer margin of lateral uropodal margin. Petasma with pars media elon- ramus setose for entire length, lacking gate; processus ventralis distally expand- tooth. ed, bearing spines; lobus terminalis distal- Type species.—By present designation, most, lobus connectens triangular and Sergestes edwardsii Krøyer, 1855. bearing numerous hooks, lobus armatus Type locality.—Atlantic Ocean at strong, curved, bearing several hooks 20uN, unknown longitude; 10u229N, along inner margin. Appendix masculina 21u169W; 7uN, 30uW. of pleopod 2 with proximalmost spines on Etymology.—Derived from the Greek medial margin much longer than those ‘neos’ meaning new, young, or recent, immediately following. Outer margin of plus the root generic epithet ‘sergestes’. lateral uropodal ramus setose for distal Species.—N. brevispinatus (Judkins, 30% of length, with tooth present. 1978); N. consobrinus (Milne, 1968); N. Type species.—By present designation, edwardsii (Krøyer, 1855); N. geminus Sergestes arcticus Krøyer, 1855. (Judkins, 1978); N. gibbilobatus (Judkins, Type locality.—Off Greenland. 1978); N. orientalis (Hansen, 1919); N. Etymology.—Derived from the Greek semissis (Burkenroad, 1940); N. tantillus ‘eu’ meaning true or original, plus the (Burkenroad, 1940). root generic epithet ‘sergestes’. Species.—E. arcticus (Krøyer, 1855); E. Parasergestes, new genus similis (Hansen, 1903). Fig. 1E Diagnosis.—Cuticle semitransparent; Neosergestes, new genus photophores absent. Carapace with he- Fig. 1D patic and supraorbital spines present. Diagnosis.—Cuticle semitransparent; Organs of Pesta present, antero- and photophores absent. Carapace with he- posterolateral organs spheroid. Ocular patic and supraorbital spines present. tubercle present or absent. Antennular Organs of Pesta present, antero- and peduncle with first and third articles posterolateral organs spheroid. Ocular subequal in length; stylocerite absent. tubercle absent. Antennular peduncle with Maxillule with palp. Maxilliped 1 with first and third articles subequal in length; palp. Maxilliped 3 much longer than stylocerite fixed. Maxillule with palp. pereopod 3; dactylus consisting of 4 Maxilliped 1 with palp. Maxilliped 3 much articles. Ischium of pereopods 1 and 2 longer than pereopod 3; dactylus consist- bearing spine. Fingers of chela of pereo- ing of 6 articles and 2 terminal spines. pod 2 subequal. Pereopod 3, coxa having Ischium of pereopods 1 and 2 bearing mesial tooth; posterior arthrobranchia VOLUME 121, NUMBER 1 77 above pereopod 3 lamellar. Pereopods 4 broad; processus ventralis apically acute. and 5 each having 6 podomeres. Branchi- Appendix masculina of pleopod 2 with ae present above pereopod 4. Two distal spines absent, or with spines gradually podomeres of pereopod 5 setose only on decreasing in size apically. Outer margin posterior margin. Petasma with lobes and of lateral uropodal ramus setose for distal processes short, often rounded, especially 30% of length, with tooth present. lobus connectens and lobus armatus; pars Type species.—By monotypy, Sergestes media broad; processus ventralis apically atlanticus H. Milne-Edwards, 1830. acute. Appendix masculina of pleopod 2 Type locality.—North Atlantic Ocean with spines gradually decreasing in size near Azores. apically. Outer margin of lateral uropodal Species.—S. atlanticus H. Milne-Ed- ramus setose for distal 60–80% of length, wards, 1830; S. cornutus Krøyer, 1855. lacking tooth. Type species.—By present designation, Sergia Stimpson, 1860 Sergestes armatus Krøyer, 1855. Fig. 2A–F Type locality.—North Atlantic in re- Diagnosis.—Cuticle opaque-red; lens- gion of 7u379N, 22.5uW. bearing or lensless photophores present Etymology.—Derived from the Greek or absent. Carapace lacking hepatic and ‘para’ meaning beside or near, plus the supraorbital spines. Organs of Pesta root generic epithet ‘sergestes’. absent. Ocular tubercle present or absent. Species.—P. armatus (Krøyer, 1855); P. Antennular peduncle with first article diapontius (Bate, 1881); P. extensus (Ha- much longer than third; stylocerite mo- namura, 1983); P. halia (Faxon, 1893); P. bile. Maxillule with palp. Maxilliped 1 stimulator (Burkenroad, 1940); P. vigilax with palp. Maxilliped 3 subequal in length (Stimpson, 1860). to pereopod 3; dactylus consisting of 4, 6, or 7 articles. Ischium of pereopods 1 and Sergestes H. Milne-Edwards, 1830 2 lacking spine. Fingers of chela of Fig. 1F pereopod 2 subequal. Pereopod 3, coxa Diagnosis.—Cuticle semitransparent; bearing mesial tooth; posterior arthro- photophores lacking. Carapace with he- branchia above pereopod 3 well devel- patic and supraorbital spines present. oped. Pereopods 4 and 5 each having 6 Organs of Pesta present, antero- and podomeres. Branchiae present above pe- posterolateral organs spheroid. Ocular reopod 4. Two distal podomeres of tubercle absent. Antennular peduncle with pereopod 5 setose along both margins. first and third articles subequal; stylocerite Petasma, except in S. lucens species group fixed. Maxillule with palp. Maxilliped 1 (Vereshchaka 2000), with pars media with palp. Maxilliped 3 subequal in length elongate; lobus inermis usually apically to pereopod 3; dactylus consisting of 7 or acute, occasionally rounded; lobus arma- 8 articles. Ischium of pereopods 1 and 2 tus strong, curved or straight, rarely bearing spine. Fingers of chela of pereo- small, occasionally with small lobus pod 2 subequal. Pereopod 3, coxa lacking accessorius at its base; processus ventralis mesial tooth; posterior arthrobranchia usually apically acute, rarely rounded or above pereopod 3 lamellar. Pereopods 4 truncate, with hooks. Petasma in S. lucens and 5 each having 6 podomeres. Branchi- with processus ventralis enlarged and ae present above pereopod 4. Two distal armed with hook(s), lobus connectens podomeres of pereopod 5 setose only on and lobus terminalis inferior in size to posterior margin. Petasma with lobes and processus ventralis, and lobus inermis processes, especially lobus connectens and reduced or absent. Appendix masculina lobus armatus, short, rounded; pars media of pleopod 2 with proximalmost spines of 78 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 2. Petasmata of selected species of Sergia.A,S. creber;B,S. grandis;C,S. inequalis;D,S. japonica; E, S. fulgens;F,S. lucens. c - capitulum; la - lobus armatus; lc - lobus connectens; li - lobus inermis; lt - lobus terminalis; pu - processus uncifer; pv - processus ventralis. VOLUME 121, NUMBER 1 79 medial margin much longer than those shorter than or subequal in length to immediately following. Outer margin of pereopod 3. Pereopods 1 and 2 lacking lateral uropodal ramus setose for distal ischial spine. Fingers of pereopod 2 chela 30% of length, with tooth present. equal. Pereopod 3 coxa with at least one Type species.—By monotypy, Sergia tooth in most species. Pereopods 4 and 5 remipes Stimpson, 1860 [based on late lacking, pereopod 5 represented by pair of mastigopus larva]. genital protuberances in male. Single Type locality.—Pacific Ocean at arthrobranchia present above where pe- 27.5uN, 138.5uE. reopod 4 would have been. Petasma with Species.—S. bigemmea (Burkenroad, pars media relatively elongate, with num- 1940); S. bisulcata (Wood-Mason, 1891); ber of distal processes usually reduced to S. burukovskii Vereshchaka, 2000; S. processus ventralis and capitulum. Ap- challengeri (Hansen, 1903); S. crosnieri pendix masculina of pleopod 2 bearing Vereshchaka, 2000; S. erythraeensis few marginal hooks. Outer margin of Iwaski & Couwelaar, 2001; S. extenuata lateral uropodal ramus with setose por- (Burkenroad, 1940); S. filicta (Burken- tion subequal to, or shorter than non- road, 1940); S. fulgens (Hansen, 1919); S. setose portion; tooth present or absent on gardineri (Kemp, 1913); S. grandis (Sund, lateral margin. 1920); S. hansjacobi Vereshchaka, 1994; Type species.—By original designation, S. inequalis (Burkenroad, 1940); S. inoa Acetes indicus H. Milne-Edwards, 1830. (Faxon, 1893); S. japonica (Bate, 1881); S. Type locality.—Ganges River. jeppesensi Vereshchaka, 2000; S. kensleyi Species.—A. americanus americanus Vereshchaka, 2000; S. laminata (Burken- Ortmann, 1893; A. americanus carolinae road, 1940); S. lucens (Hansen, 1922); S. Hansen, 1933; A. binghami Burkenroad, maxima (Burkenroad, 1940); S. oksanae 1934; A. chinensis Hansen, 1919; A. Vereshchaka, 2000; S. phorca (Faxon, erythraeus Nobili, 1905; A. indicus H. 1893); S. plumea (Illig, 1927); S. prehensi- Milne-Edwards, 1830; A. intermedius lis (Bate, 1881); S. regalis (Gordon, 1939); Omori, 1975; A. japonicus Kishinouye, S. robusta (Smith, 1882); S. scintillans 1905; A. johni Nataraj, 1947; A. marinus (Burkenroad, 1940); S. splendens (Sund, Omori, 1975; A. natalensis Barnard, 1955; 1920); S. stellata (Burkenroad, 1940); S. A. paraquavensis Hansen, 1919; A. serru- talismani (Barnard, 1947); S. tenuiremis latus (Krøyer, 1855); A. sibogae australis (Krøyer, 1855); S. umitakae Hashizume Colefax, 1940; A. sibogae sibogae Hansen, & Omori, 1995; S. vityazi Vereshchaka, 1919; A. sibogae sibogalis Achuthankutty 2000; S. wolffi Vereshchaka, 1994. & George, 1973; A. vulgaris Hansen, 1919. Other Sergestid Genera Peisos Burkenroad, 1945 Acetes H. Milne-Edwards, 1830 Fig. 3B Fig. 3A Diagnosis.—Cuticle lacking photo- Diagnosis.—Cuticle transparent, lack- phores. Carapace with supraorbital and ing photophores. Carapace with supraor- hepatic spines. Organs of Pesta absent. bital and hepatic spines present. Organs Tiny ocular tubercle present. Antennular of Pesta absent. Tiny ocular tubercle peduncle with third article subequal to present. Antennular peduncle with third first in male, shorter than first article in article shorter or subequal to first in female; stylocerite fixed. Maxillule with female, much longer than first in male; palp reduced to low conical tubercle. stylocerite fixed. Maxillule lacking palp. Maxilliped 1 with palp. Maxilliped 3 Maxilliped 1 lacking palp. Maxilliped 3 shorter than pereopod 3. Pereopods 1 80 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Petasmata of selected species of other sergestid genera. A, Acetes indicus;B,Peisos petrunkevitchi; C, Petalidium foliaceum;D,Sicyonella maldivensis. c - capitulum; la - lobus armatus; lc - lobus connectens; li - lobus inermis; lt - lobus terminalis; pu - processus uncifer; pv - processus ventralis. VOLUME 121, NUMBER 1 81 and 2 lacking ischial spines. Fingers of setose for about distal fifth, bearing tiny chela of pereopod 2 subequal. Pereopod tooth distally. 3, coxa lacking tooth; single arthrobran- Type species.—By monotypy, Petali- chia present. Pereopod 4 consisting of 5 dium foliaceum Bate, 1881. podomeres; arthrobranchia present. Pe- Type locality.—Off Marion Island, reopod 5 represented by pair of vestigial 46u469S, 45u319E, 2516 m.; South of conical tubercles in male, consisting of 3 Australia, 47u259S, 130u229E, 3935 m. podomeres in female. Petasma with pars Species.—P. foliaceum Bate, 1881; P. media relatively short but not broad; obesum (Krøyer, 1859); P. suspiriosum bearing short capitulum and elongate Burkenroad, 1937. processus ventralis. Appendix masculina bearing pair of marginal hooks. Outer Sicyonella Borradaile, 1910 margin of lateral uropodal ramus setose Fig. 3D for less than two-fifths of length, bearing Diagnosis.—Cuticular photophores tooth. lacking. Carapace having supraorbital Type species.—By original designation, and hepatic spines. Organs of Pesta Peisos petrunkevitchi Burkenroad, 1945. lacking. Ocular tubercle lacking. Anten- Type locality.—Montevideo, Uruguay. nular peduncle with third article much Species.—P. petrunkevitchi Burken- shorter than first; stylocerite fixed. Max- road, 1945. illule and maxilliped 1 each with palp. Maxilliped 3 longer and more robust than Petalidium Bate, 1881 pereopod 3; dactylus consisting of 4 Fig. 3C articles. Pereopods 1 and 2 lacking ischial Diagnosis.—Cuticle lacking photo- spines. Pereopod 2, fingers of chela sub- phores. Carapace having minute hepatic equal. Pereopod 3, coxa lacking tooth. spine, lacking supraorbital spine. Organs Two branchiae present above pereopod 4. of Pesta absent. Ocular tubercle present. Pereopods 4 and 5 each of 7 podomeres. Antennular peduncle with third article Last 2 podomeres of pereopod 5 setose on subequal to or slightly shorter than first; posterior margin only. Petasma with pars stylocerite fixed. Maxillule and maxilliped media and processes relatively elongate; 1 each with well-developed palp. Maxilli- lobus armatus short, straight; processus ped 3 not longer or more robust than ventralis consisting of 2 or 3 strong lobes. pereopod 3, propodus and dactylus not Appendix masculina of pleopod 2 short, subdivided. Pereopod 2, fingers of chela ovate, bearing few distal marginal setae. subequal. Pereopod 3, coxa 3 lacking Outer margin of lateral uropodal ramus tooth; posterior arthrobranchia lamellar. setose for distal 20%, bearing tooth. Branchiae present or absent above pereo- Type species.—By monotypy, Sicyo- pod 4. Pereopods 4 and 5 each of 6 nella maldivensis Borradaile, 1910. podomeres, two distal podomeres setose Type locality.—Maldive Islands, Indian only on posterior margin. Petasma with Ocean. pars media elongate; processes, especially Species.—S. antennata Hansen, 1919; S. lobus terminalis and distally bifurcate elegans Calman, 1913; S. inermis (Paul’- processus ventralis relatively elongate; son, 1875): S. maldivensis Borradaile, 1910. lobus armatus usually bipartite; lobus connectens and lobus inermis short. Appendix masculina of pleopod 2 rela- Key to the Genera of the tively elongate, 3–4 times longer than Family Sergestidae wide, bearing few distal marginal hooks. 1. Pereopod 4 present, pereopod 5 Outer margin of lateral uropodal ramus present or absent ...... 2 82 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

— Pereopods 4 and 5 absent . . . . Acetes Acknowledgments 2. Pereopod 4 of fewer than 7 podo- meres ...... 3 In the original unfinished draft of this — Pereopod 4 consisting of 7 podomer- paper, Brian Kensley acknowledged Drs. es ...... Sicyonella A. Crosnier, I. Perez Farfante, A. B. 3. Pereopods 4 and 5 consisting of 6 Williams,F.A.Chace,M.Omori,andT. podomeres ...... 4 Kikuchi for their many useful comments — Pereopod 4 consisting of 5 podo- on this paper. David Judkins thanks meres; pereopod 5 vestigial in male, Marilyn Schotte of the Department of of 3 podomeres in female . . . . Peisos Invertebrate Zoology, Smithsonian Insti- 4. Processus ventralis of petasma not tution for assistance in preparing the final bifurcate; outer margin of lateral draft of the manuscript and its illustrations. uropodal ramus setose for 30% or moreoflength ...... 5 — Processus ventralis of petasma distal- Literature Cited ly bifurcate; outer margin of lateral uropodal ramus setose for 20% or Achutankutty, C. T., & M. J. George. 1973. Acetes lessoflength ...... Petalidium sibogalis sp. nov. (Crustacea: Decapoda, 5. Organs of Pesta present; body semi- Sergestidae) from Cochin back-waters with a note on its impregnation.—Indian Journal of transparent ...... 6 Marine Science 2:139–144. — Organs of Pesta absent; body opa- Barnard, K. H. 1947. Descriptions of new species of que ...... Sergia South African decapod Crustacea, with notes 6. Maxilliped 3 much longer than pe- on synonymy and new records.—Annals and reopod3 ...... 7 Magazine of Natural History (11)13:361–392. — Maxilliped 3 subequal in length to ———. 1955. Additions to the fauna-list of South pereopod3 ...... 9 African Crustacea and Pycnogonida.—Annal 7. Last 2 podomeres of pereopod 5 of the South African Museum 43(1):1–107. setose only on posterior margin . . . 8 Bate, C. S. 1881. On the Penaeidea.—Annals and Magazine of Natural History (5)8:169–196. — Last 2 podomeres of pereopod 5 Borradaile, L. A. 1910. The Percy Sladen Trust setose on both margins . . Allosergestes Expedition to the Indian Ocean in 1905, under 8. Outer margin of lateral uropodal the leadership of Mr. J. Stanley Gardiner. ramus setose for entire length; dacty- Volume 2. No. 10. Penaeidea, Stenopidea, lus of maxilliped 3 consisting of 6 and Reptantia from the Western Indian articles ...... Neosergestes Ocean.—Transactions of the Linnean Society — Outer margin of lateral uropodal of London, Series 2, Zoology 13(2):257–264. ramus not setose for entire length; Burkenroad, M. D. 1934. The Penaeidea of Louisi- dactylus of maxilliped 3 consisting of ana with a discussion of their world relation- 4 articles ...... Parasergestes ships.—Bulletin of the American Museum of Natural History 68(2):61–143. 9. Two distal podomeres of pereopod 5 ———. 1937. The Templeton Crocker Expedition. setose only on posterior margin; XII. Sergestidae (Crustacea Decapoda) from outer margin of lateral uropodal the Lower Californian region, with descrip- ramus setose for distal 1/3, having tions of two new species and some remarks on tooth ...... 10 the Organs of Pesta in Sergestes.—Zoologica, — Two distal podomeres of pereopod 5 New York 22(4):315–329. setose on both margins; outer margin ———. 1940. Preliminary descriptions of twenty- of lateral uropodal ramus setose for one new species of pelagic Penaeidea (Crus- distal 2/3, lacking tooth ...... tacea Decapoda) from the Danish oceano- ...... Deosergestes graphical expeditions.—Annals and Maga- zine of Natural History (11)6:35–54. 10. Third article of antennular pedun- ———. 1945. A new sergestid shrimp (Peisos cle subequal to or longer than petrunkevitchi, n. gen., n. sp.), with remarks first ...... Sergestes on its relationships.—Transactions of the — Third article of antennular peduncle Connecticut Academy of Arts and Sciences much shorter than first . . . Eusergestes 36:553–593. VOLUME 121, NUMBER 1 83

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