Records of Shallow-Water Marine Isopods from Bermuda with Descriptions of Four New Species Author(s): Brian Kensley Source: Journal of Biology, Vol. 14, No. 2, (May, 1994), pp. 319-336 Published by: The Crustacean Society Stable URL: http://www.jstor.org/stable/1548912 Accessed: 25/07/2008 14:24

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http://www.jstor.org JOURNAL OF CRUSTACEAN BIOLOGY, 14(2): 319-336, 1994

RECORDS OF SHALLOW-WATER MARINE ISOPODS FROM BERMUDA WITH DESCRIPTIONS OF FOUR NEW SPECIES

Brian Kensley

ABSTRACT An extensivecollection of marineisopods from the PeabodyMuseum, Yale University,taken from 114 collecting stations on Bermuda,is reported.The collection adds 10 new recordsto the list of known Bermudanisopods, includingtwo undescribedspecies, Mesanthurabermu- densisand Joeropsisnigricapitis. These two speciescan be recognizedinitially by theirdistinctive pigmentpatterns. Fresh material shows Anthomuda stenotelson Schultz to be a junior synonym of Antheluraaffinis Richardson. Also included in the report are the descriptionsof two new species collected from a cave, Munnogoniumsomersensis and Stenobermudailiffei. The latter, which exhibits protogynoushermaphroditism, is shown to be a member of the Gnathostene- troididae.

While several reportson the isopod fauna Of the marine forms, the Carpias of Bermudahave been published (Richard- appears to be ecologically the most wide- son, 1902; Schultz, 1986; Kensley and spread,occurring in 86 of the 114 collection Schotte, 1989), the collection of about 1,500 stations (see Table 1). One species, Carpias marine and littoral specimens reportedhere minutus (Richardson),is representedat 40 from the Peabody Museum, Yale Univer- of these stations. sity, includes 23 species and is probablythe Includedin this reportare descriptionsof most comprehensive collection yet made. two new species collected from Bermudan The isopods were collected from the inter- caves by Dr. Thomas Iliffe of Texas A&M tidal to about 10-m depth, from a variety University, and deposited in the National of habitatsincluding algal clumps and turfs, Museum of Natural History (USNM), turtle grass beds, sponges, rocks, coral rub- SmithsonianInstitution. Holotypes from the ble and coarse sediments, and mangrove Yale collection are depositedin the Peabody roots. Museum, Yale University (YPM), along The current list of 60 isopod species with the rest of the collection which is cat- known from Bermuda (Table 1, including alogued in a block commencing with cata- for completeness the terrestrialoniscideans logue number YPM 9551; where available, and the parasiticepicarideans) contains 10 paratypeswere deposited in the Smithson- new records,five of which have a Caribbean ian Institution and the BermudaBiological distribution. Of the remaining five, one Station (BBS). wood-boring species, Limnoria multipunc- tata Menzies, has a widespreadmarine dis- SYSTEMATICS four were tribution; species previously un- AntheluridaePoore and described.This list cannot be consideredas Family Lew Ton, 1988 complete, given that single specimens of an Anthomuda unidentifiable and affinis (Richardson, 1902) munnid, arcturid, gna- Fig. 1 thiid are present in the Peabody collection. Further, no records of interstitial Antheluraaffinis Richardson, 1902: 288, pi. 28, figs. isopods 29-32. are known,probably because no has attempt Ananthuraaffinis: Schultz, 1969: 101, fig. 137. been made to collect them. Overall, how- Anthomudaaffinis: Poore and Lew Ton, 1988: 500. ever, the Bermudan marine isopod fauna Anthomudastenotelson Schultz, 1979: 909, figs. 12- continues to be primarilytropical/subtrop- 21.-Poore and Lew Ton, 1988: 500.-Kensley ical Caribbeanin its affinities,while its 30% and Schotte, 1989: 23, 270, fig. 7A. endemism (14 of 46 marine species), pre- Material Examined.-Holotype of A. affinis, YPM since the last Pleisto- 3349, nonovigerous 9 total length (tl) 5.0 mm, Ber- sumably developed muda, collector A. E. Verrill, 1901.-Holotype of A. cene glaciation, emphasizes its relative iso- stenotelson,USNM 171263, nonovigerousQ tl 8.0 mm, lation. paratypeUSNM 171264, nonovigerous9 tl 7.9 mm, 319 320 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 14, NO. 2, 1994

Table 1. Isopod recordsfrom Bermuda.! stygobiont. Table 1. Continued. + endemic form. * new record. SuborderOniscidea SuborderAnthuridea lentus Amakusanthura and Agabiformius (Budde-Lund,1885) magnifica(Menzies Armadillidiumvulgare (Latreille, 1840) Frankenberg,1966) Armadilloniscus + Anthelura ellipticus(Harger, 1878) affinis(Richardson, 1902) Halophilosciacouchi (Kinahan, 1858) + Apanthuraharringtoniensis Wigele, 1981 + Colanthuratenuis 1902 Leptotrichuspanzeri (Audouin, 1826) Richardson, baudinianaH. Milne Edwards,1840 +? Curassanthurabermudensis 1985 Ligia Wigele, Porcelliolaevis Latreille,1804 * Kupellonuraimswe (Kensley, 1982) * Porcelliolamellatus Budde-Lund, 1885 + Mesanthurabermudensis, new species Porcellioscaber 1804 Paranthura Latreille, infundibulataRichardson, Porcellionidespruinosus (Brandt, 1833) 1902 Porcellionides Pendanthura Menzies and sexfasciatus(Koch, 1847) tanaiformis Stenoniscuspleonalis Aubertand Dollfus, Glynn, 1968 1890 SuborderAsellota Tylos latreilleiAudouin, 1825 + Atlantaselluscavernicolus Sket, 1979 Tylos niveusBudde-Lund, 1885 + * Carpiasalgicola (Miller, 1941) Carpiasbermudensis Richardson, 1902 + Carpiasminutus (Richardson, 1902) * Carpiasserricaudus Menzies and Glynn, off Castle Roads, Bermuda,9.0 m.-Peabody locality 1968 123, North Rock, 9 m, 1 nonovigerousQ tl 5.0 mm. Joeropsisrathbunae Richardson, 1902 + * Joeropsisnigricapitis, new species + * Munnogoniumsomersensis, new species Remarks.-Poore and Lew Ton (1988) not- * Santia milleri(Menzies and Glynn, 1968) ed that Anthelura affinis and Anthomuda Stenetriumstebbingi Richardson, 1902 stenotelsonappeared to be very similar, and + Stenobermudaacutirostrata Schultz, 1979 were Examinationof + * Stenobermudailiffei, new species perhapssynonymous. the type material of both species confirms SuborderEpicaridea this synonymy. The pereiopodsand uropo- Bopyrissawolffi Markham, 1978 dal rami, previously poorly illustrated,are Cancriceponchoprae (Nierstrasz and Bren- der a Brandis, 1925) figuredhere, as is the most recentlycollected Leidya bimini Pearse, 1951 nonovigerous female, to show the dorsal Parathelgespiriformis Markham, 1972 pigmentation. Parathelgestumidipes Markham, 1972 Pseudioneaffinis (Sars, 1882) Family AnthuridaeLeach, 1814 Probopyrinellalatreuticola (Gissler, 1882) Mesanthura bermudensis, new species + Stegias clibanariiRichardson, 1904 3 Synsynellachoprae (Pearse, 1932) Figs. 2, Synsynelladeformans Hay, 1917 MaterialExamined. -Holotype, YPM 9553, 6 6.8 mm, SuborderFlabellifera Peabodylocality 20, Shelly Bay, in subtidalalgae, col- lectorM. F. 3 YPM Alcironakrebsi 1890 Gable, Jun 1985.-Paratype, 9554, Hansen, 6 5.6 Whalebone + Arubolanaaruboides (Bowman and Iliffe, mm, Peabodylocality 45, Bay, rocky intertidal,collector M. F. Gable, 24 1987.-Para- 1983) May 1902 type, YPM 9555, nonovigerous Q 6.2 mm, Peabody Colopisthusparvus Richardson, off oestrum locality 47, promontory Shelly Bay, algal clumps, Cymothoa (Linnaeus,1793) collectorM. F. 28 1987. YPM perforata(Moore, 1901) Gable, May -Paratype, Dynamenella 9556, manca 2.6 mm, locality 73, cove west personataKensley, 1987 Peabody Eurydice of Bermuda Station, Reach, turtleand * Excirolana mayana (Ives, 1891) Biological Ferry manateegrass and sediment,collector A. J. Baldinger, Excorallanaquadricornus (Hansen, 1890) 6 * 1957 29 May 1987.-Paratype, YPM 9557, 4.6 mm, Pea- Limnoria multipunctataMenzies, Evans from Limnoriatuberculata Sowinsky, 1884 body locality 74, Bay, algal scrapings pier, collectorE. A. Lazo-Wasem,30 Lironecareniformis Menzies and Franken- May 1987.-Paratype, YPM 9558, manca2.7 mm, Peabodylocality 105, North berg, 1966 Schioedteand Mei- Rock,red algae, collector A. J. Baldinger,2 Jun 1987.- Nerocilaacuminata YPM 3.2 manca 1881 Paratypes, 9559, nonovigerousQ mm, nert, 2.33 196, cove west of Devon- Paracerceiscaudata (Say, 1818) mm, Peabodylocality shireBay, washings from rocks, midtide level, collector SuborderValvifera E. A. Lazo-Wasem, 15 Jun 1988.-Paratype, BBS, S Idotea balthica(Pallas, 1772) 4.4 mm, Peabody locality 61, cove west of Bermuda Idotea metallicaBosc, 1802 BiologicalStation, Ferry Reach, bed of Thalassia,col- lectorA. J. Baldinger,29 May 1987.-Paratype, USNM 243585, 8 5.4 mm, Peabody locality 99, SharkHole, KENSLEY: MARINE ISOPODS FROM BERMUDA 321

G

Fig. 1. Anthomuda affinis: A, habitus nonovigerous 9 PMY locality 123. Paratype of A. stenotelson, USNM 171264: B, maxilliped; C, pereiopod 1; D, pereiopod 2; E, pereiopod 4; F, uropodal exopod; G, uropodal protopod and endopod; H, pereiopod 7.

Harrington Sound, subtidal encrusted rocks, collector Description. -Nonovigerous female: Body E. A. Lazo-Wasem, 1 Jun 1987.-Paratype, USNM proportions C<1<2=3=4=5=6 243586, nonovigerous e 4.9 mm, Peabody locality 46, C < 1 Harrington Sound, attached wall community, collector > = P Cephalon th moderate rostral M. F. Gable, 25 May 1987. point; anterolateral eyes strongly pigment- 322 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 14, NO. 2, 1994 ed. Telson posteriorly rounded, having of aesthetascs.Pereiopod 1, carpuspostero- transparentmarginal flange, with 6 pairs of distally rounded; propodus with rounded apical setae. transparentlobe at aboutmidlength of palm, Antennular peduncle with basal article submarginalband of stiff setae on mesial broaderand slightly longerthan 2 distal ar- surface;unguis slightly less than length of ticles together; flagellum of 3 articles, ter- rest of dactylus. Pleopod 2, endopod with minal article bearing 4 aesthetascs. Anten- 5 distal plumose setae, copulatorystylet ar- nal flagellumconsisting of 4 setose articles. ticulatingin proximalhalfofmesial margin, Mandibular palp, article 3 with 6 distal reaching well beyond distal margin of ra- spines, subapicalspine longest;incisor with mus;exopod with transversesuture at about 3 sclerotized cusps; lamina dentata having midlength, bearing 8 distal plumose setae. 5 teeth; molar truncate, barely sclerotized. Color pattern:peduncles of antennulesand Maxilla having 6 distal spines. Maxilliped antennae pigmented; entire dorsal surface with basalarticle of palp slightlyshorter than ofcephalon pigmented;pereionites 1-7 with article 2; terminal article with mesial mar- ovate patch of pigment; posterior half of gin truncate, bearing 3 fringed setae. Pe- pleon with 5 rough transverse bands, me- reiopod 1, carpus posterodistallyrounded; diallyjoined, and with anteromedialexten- propodal palm with transparent rounded sion; uropods and telson stronglypigment- lobe at about midpoint; unguis about equal ed, that of latter falling well short of apex. in to rest with small length ofdactylus, spine -The name derives at base. Etymology. specific Pereiopod 2, carpus triangular, from the the island of Ber- free anterior with type locality, lacking margin;propodus muda. strong posterodistal spine, fringed scales along posterior margin; unguis half length Remarks.-Mesanthura bermudensisclose- of rest ofdactylus. Pereiopod 7, carpuswith ly resembles the widespreadM. paucidens short spine on posterior margin, latter lon- Menzies and Glynn, 1968, in overall pro- ger than freeanterior margin; propodus with portions and armature of the appendages, posterodistalfringed scales and strongspine; and in the general distribution of pigment unguis less than half length of rest of dac- (the latter being well-nigh species-specific tylus. Pleopod 1, endopod parallel-sided, within the genus). The following differences about 6 times longer than wide, with 10 can be detected, however, in a comparison distalplumose setae;exopod about 3.5 times with material from Puerto Rico (Menzies wider than endopod, widest at midlength. and Glynn, 1968), Belize (Kensley, 1982), Uropodal exopod almost twice longer than and Martinique(Miiller, 1991a). (1) Distri- wide, apically acute, with broad subapical bution of pigment: the males of M. pauci- notch, bearing numerous marginal setae; dens do not have pigmentedantennular and endopod broadly rounded, one-fifth longer antennalpeduncles. The pleon of the earlier than basal width, bearing numerous mar- species lacks both the short anterior ginal setae. Color pattern: cephalon with "threads"of pigment as well as an antero- broad pigment patch between eyes and medial extension seen in both males and reachingposteriorly three-fourths length of females from Bermuda. (2) The Bermudan segment;pereionites 1-7 each with roughly male has two more articlesin the antennular rectangulardorsal patch of pigment falling flagellumthan do the specimens from Mar- well short of posterior margin, those of pe- tinique. (3) The endopod of pleopod 1 has reionites 2-5 with clear oval area on mid- 10 distal plumose setae in the Bermudan line; pleon with pigment patch anteriorly form, three or four in those from the rest of lobed, and with short anterolateralpigment the Caribbean. (4) The telson of the Ber- lines; telson with rectangularpigment patch mudan form has four pairs of long simple fallingwell short of posteriormargin; small setae, two pairs of short simple setae, and patch of pigment on uropodalendopod and one pair of featheredsetae on the posterior exopod. margin. The Caribbeanform has two pairs Male: Body proportions,C < 1 < 2 > 3 of long setae and one pair of featheredsetae. =4 < 5 > 6 > 7

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3 3 3 E

r'I, h,ji9 PB S^l

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Fig. 2. Mesanthura bermudensis, new species: A, habitus 6, dorsal view; B, habitus Q,dorsal view; C, antennule 2; D, antennule 8, aesthetascs omitted; E, antenna; F, mandible; G, maxilla; H, maxilliped; I, pleotelson. Fig. 3. Mesanthura bermudensis, new species: A, pereiopod 1 6; B, pereiopod 1 i; C, pereiopod 2; D, pereiopod 7; E, pleopod 1; F, pleopod 2 8; G, uropodal exopod; H, uropodal protopod and endopod. KENSLEY: MARINE ISOPODS FROM BERMUDA 325

M. paucidens, perhaps even derived from mesiodistalhook; mesial ramusslightly lon- this species, but also reflect the isolation of ger than lateral. the The is Bermudan population. latter Etymology.-The specificepithet, which re- therefore a regardedas separatespecies. fers to the dorsally pigmented head of the ,is derivedfrom the Latinniger, black Family JoeropsidaeNordenstam, 1933 or dark, and capitus, head. Joeropsisnigricapitis, new species Remarks.--The only species of Joeropsis Fig. 4 previously recorded from Bermuda is J. MaterialExamined. -Holotype, YPM 9551, 2.9 mm, rathbunaeRichardson, 1902. The two male paratypes,YPM 9552, 37 d8 2.2-2.9 mm, 18 ovigerous syntypes of this species were examined 99 2.0-2.2 mm, 164 immature and juvenile.-Para- (USNM 24870, YPM 3251) and severaldif- types, USNM 243584, 10 8, 10 ovigerous99, Peabody ferences noted when with the locality 196, cove west of Devonshire Bay, washings compared from rocks,0.3 m, collectorE. A. Lazo-Wasem,15 Jun present species. These differences include 1988.-Paratypes, BBS, Peabody locality 211, 3 66, 1 the color pattern(faintly reticulateover en- ovigerous9, 6 juvenile, Bailey'sBay, 1 m, collectorE. tire body, strongeston cephalon, in J. rath- A. Lazo-Wasem,17 Jun 1988.-Peabody locality 3, 1 bunae, dark brown on most 8, 1 9, cove on small island in Tobacco Bay, collector strong pigment M. F. Gable, 2 Jun 1985.-Peabody locality 19, 1 8, of cephalon and pereionite 4, fainter retic- off Barryand GrenadireRoads, St. George's,in algal ulation on pereionites 2,3,6,7, and anterior clumps, 2-3 m, collector M. F. Gable, 2 Jun 1985.- pleon, pereionites 1 and 5 unpigmented in Peabody locality 47, 1 9, promontoryoff Shelly Bay, J. nigricapitis),the rostral shape (apically from algal clumps and shells, collector M. F. Gable, in 28 May 1987.-Peabody locality 48, 1 8, north side of broadlyrounded J. rathbunae,very faint- Shelly Bay, shallow subtidal algal scrapings,collector ly emarginatein J. nigricapitis),the pleonal E. A. Lazo-Wasem,28 June 1987. serrations(4 in the earlier species, 5 in the Description. -Body glabrous. Cephalon present species), and the relative lengths of lackinglateral serrations; rostrum wider than the terminalantennular article, and the uro- median length,with faint indentationin an- podal rami. teriormargin, latter bearing border of trans- parent scales. Pleon with 5 lateral serra- Family ParamunnidaeVanhoffen, 1914 tions. Color pattern: dorsal surface of Munnogoniumsomersensis, new species cephalon dark red-brown;pereionite 1 un- Figs. 5, 6 pigmented;pereionites 2-4 with dorsal re- MaterialExamined. -Holotype, USNM 243587, 0.93 ticulation, that of 4 strongest;pereionite 5 mm, paratypes, USNM 243588, 8 0.71 mm, 4 oviger- unpigmented;pereionites 6 and 7 with faint ous i$0.98 mm, 1juvenile 2 0.57 mm, Red Bay Cave, reticulation; anterior pleon with irregular collectorT. Iliffe, 23 Oct 1986. reticulation. Description. -Male: Body pear-shaped, Basal antennular article with fringe of twice longerthan wide, widest at pereionite transparentscales; terminal article shorter 3. Frontal marginof cephalon broadly con- than subterminal, bearing 2 aesthetascs. vex, flattened at midline; ovate black pig- Fourthantennal peduncle article with fringe ment of eyes on short lateralprojections. In of transparentscales; flagellum consisting of dorsal view, coxal plates clearly visible on largebasal and 6 tiny distal articles.Mouth- pereionites5-7. Pleotelson consistingof an- parts typical of genus. Mandibularpalp, ar- terior pleonite plus subcircularpleon, latter ticle 2 with 3 fringed spines on laterodistal with 8 lateralteeth, posteriormargin round- margin, terminal article with 7 distal setae. ed. Maxillipedal palp with article 2 broadest, Antennularpeduncle of 2 subequalelon- with rounded lobe at distomesial margin; gate articles, several jointed setae distally articles 4 and 5 slender. Pereiopods typical on article 2; flagellum of 4 articles, distal- of genus, pereiopod 1, dactylus with 2 un- most bearingsingle elongateaesthetasc. An- gui; pereiopods 2-7 with 3 dactylar ungui tennal peduncle with article 2 short, set on each. Male pleopods typical of genus, pleo- anterior margin of article 3; article 4 set at pod 5 consisting of single lamellar ramus. right angle to article 3, penultimate article Operculum(pleopod 2) of female pentago- slightly shorterthan last article, latter with nal, marginsdistally faintly concave, setose, severaljointed setae distally; flagellumof 7 apical anglejust less than 90?.Uropod with articles.Mandible lacking palp; incisor hav- 326 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 14, NO. 2, 1994

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Fig. 4. Joeropsisrathbunae, PMY 3251: A, rostrum;B, antennule.Joeropsis nigricapitis, new species:C, habitus, dorsal view; D, rostrum; E, antennule; F, antenna; G, mandible; H, maxilla 1; I, maxilla 2; J, maxilliped; K, pleopod 1 B;a;L, pleopod 2 6;a; M, pleopod 3; N, pleopod 4; 0, pleopod 5; P, uropod.

Fig. 5. Munnogonium somersensis, new species: A, habitus, dorsal view; B, pereiopod 1; C, antennule; D, antenna; E, pereiopod 2; F, pereiopod 7. KENSLEY: MARINE ISOPODS FROM BERMUDA 327

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C D 328 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 14, NO. 2, 1994

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Fig. 6. Munnogonium somersensis, new species: A, left and right mandible, with surface of molar; B, maxilla 1; C, maxilla 2; D, maxilliped; E, uropod; F, pleopod 1 6; G, operculum 9. KENSLEY: MARINE ISOPODS FROM BERMUDA 329 ing severalacute cusps;spine row consisting eye is larger in M. somersensis. The uro- of lacinia mobilis having 3 cusps plus 3 podal exopod is relatively more slender in spines, or of 4 spines; molar distally trun- the new species, while the first pleonite and cate, with few small marginalteeth. Maxilla anteriorpleon are longer than in the some- 1 with 4 setae on inner ramus, 9 setae on what squat M. wilsoni. outer. Maxilla 2, 2 outer rami each with 4 Gnathostenetroididae distal setae, inner ramus with setae and set- Family ules of thickness. Maxillipedal en- Kussakin, 1967 varying Stenobermuda new dite 2 hooks, distal margin iliffei, species having coupling 10 with few slender setae; palp of 5 articles, Figs. 7, 8, 9, article 3 widest, distal article about half MaterialExamined. -Holotype, USNM 243589, ovi- length of article 4. Pereiopod 1 basis sub- gerous2 2.9 mm, paratypes,USNM 243590, 6 d! 2.1 in to merus car- mm, 2.3 mm, 2.3 mm, 2.3 mm, 2.6 mm, 3.0 mm, 7 equal length plus ischium; juvenile, WalsinghamCave, Bermuda,collector T. I1- pus with 2 strong sensory spines on poste- iffe, 11 Sep 1986.-Paratypes, USNM 243591, ovi- rior margin;propodus ovate, with 5 slender gerous Q2.9 mm, 6 d! 2.1 mm, 2.3 mm, 2.4 mm, 2.5 setae on convex posterior margin; unguis mm, 3.0 mm, 3.2 mm, 4 juvenile, WalsinghamCave, subequalin lengthto rest ofdactylus, strong Bermuda,collector T. Iliffe, 10 Oct 1986. accessory spine present. Pereiopods 2-7 Description.-Body length/widthratio 3.5- similar, elongate-slender,carpi and propodi 3.6. Cephalon having triangular, apically each having 2 slendersensory spines on pos- subacute rostrum; anterolateral process terior margin; ungui subequal in length of reachinganteriorly just beyond distal mar- rest of slender dactyli. Pleopod 1 broadly gin of first antennal article; eye consisting sagittate,median fused rami longerthan free of 4 close-set ommatidia. Pleon consisting distal part. Uropodal endopod about five- of 2 short anterior pleonites plus shield- eighths length of, and slightly wider than, shaped pleotelson; latter with single pos- exopod, with 2 distal setae; exopod with terolateraltooth with short stout seta just cluster of severaljointed setae distally. posteriorto notch;posterior margin broadly Female: slightly broader than male; rounded. mouthparts and pereiopods as in male. Antennularflagellum with 3 articles,sub- Pleonal operculum distinctly longer than terminalbearing single aesthetasc,terminal greatestwidth, lateralmargins bearing about with 2 aesthetascs.Antenna about 1.25 times 10 setae, posterior margin produced, nar- body length; peduncle article 3 with short rowly rounded. scale;flagellum with about 50 articles.Man- dibular with article2 -The derives palp 3 articles, bearing Etymology. specific epithet 2 and 4 shorter setae from Somers a formername for Ber- stout setae intervening Island, on distolateral article3 with muda. margin, falcate, row of 14 fringedspines increasingin length Remarks.-The genus Munnogonium cur- distally to stout terminalspine; incisor hav- rently contains six species, namely, M. ing 4 sclerotized cusps; spine row with 9 waldronenseGeorge and Str6mberg, 1968, stout dentate spines, or with lacinia mobilis from the San Juan Archipelago, Washing- plus 5 (?6) dentate spines; molar apically ton State; M. maltinii (Schiecke and Fresi, broadly truncate and sclerotized, having 1972), from the Bay of Naples, Mediterra- marginaldenticles. Maxilla 1, inner ramus nean Sea; M. subtilis Kensley, 1976, from with 4 distal setae, outer ramus with 9 den- Amsterdam Island, Indian Ocean; M. wil- tate spines. Maxilla 2, inner ramus with 4 soni Hooker, 1985, from the FloridaMiddle distal fringed setae plus 7 setae on mesial Grounds; M. polynesiensis Miiller, 1989, margin;both lobes of outer ramus bearing from the Society Islands; M. adenensis 4 distal fringed setae. Maxillipedal epipod Miiller, 1991b, from the Gulf of Aden. The acutely tapering, subequal in length to en- only known western Atlantic representa- dite; latter bearing 3 coupling hooks, distal tive, M. wilsoniHooker, 1985, from the Gulf marginbearing 3 broadly expanded fringed of Mexico, falls within the same size range spines, single dentate spine, and short sim- as, and is very similar to, the present Ber- ple mesiodistal spine; palp with 5 articles mudan species, but differs in a few subtle bearing several simple setae mesiodistally, features. The pigmented ovate part of the 3 basal articlesbroad, 2 distal articles slen- 330 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 14, NO. 2, 1994

F KENSLEY: MARINE ISOPODS FROM BERMUDA 331

A

H

G

~\ F / Fig. 8. Stenobermudailiffei, new species:A, maxilla 1; B, apex of outer ramusenlarged; C, maxilla 2; D, apex of outer ramus enlarged;E, inner ramus enlarged;F, lower lip; G, maxilliped;H, apex of maxillipedalendite enlarged.

Fig. 7. Stenobermudailiffei, new species;A, 8 2.5 mm; B, left mandible;C, left mandible,spine row enlarged; D, right mandible;E, right mandible, spine row enlarged;F, pleotelson of male in ventralview; G, antennule; H, uropod;I, terminalarticle of mandibularpalp. 332 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 14, NO. 2, 1994

B KENSLEY: MARINE ISOPODS FROM BERMUDA 333

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7~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~i

Fig. 10. Stenobermudailiffei, new species:A, 2 operculum,with precursorsof 6 pleopod 1; B, S pleopod 1; C, S pleopod 2; D, pleopod 3; E, pleopod 4; F, pleopod 5. der. Pereiopod 1 in male: subchelate; basis width, posterior margin with 4 stout setae, subequal in length to carpus, merus, and distal margin armed with 6 distally expand- ischium together; carpus with 5 fairly stout ed and denticulate spines, largest at pos- setae on posterior margin; propodus wid- terodistal comer; dactylus curved, with row ening distally, about twice longer than distal of 6 slender denticulate spines on inner mar-

Fig. 9. Stenobermuda iliffei, new species: A, a pereiopod 1; B, 6 pereiopod 1, apex of propodus and dactylus enlarged; C, Q pereiopod 1; D, Q pereiopod 1, apex of propodus and dactylus enlarged; E, pereiopod 2; F, pereiopod 2, dactylus enlarged; G, pereiopod 7; H, pereiopod 7, dactylus enlarged. 334 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 14, NO. 2, 1994 gin. Pereiopod 1 in female: carpus with 4 roditism explains why the holotype and al- stout setae on posterior margin; propodus lotype of S. acutirostrata Schultz, both of with single stout seta on posterior margin, which possess a marsupium, also possess distal marginwith 3 distally expandedden- first and second male pleopods. Schultz ticulate spines; dactylus with 4 denticulate (1979), in describingthis species, missed the spines on posterior margin. Pereiopods 2- marsupiumin the holotype, but referredto 7 similar, becoming slightly more elongate the allotype as a hermaphrodite, thereby posteriorly;merus short, about half length hinting at the occurrence of protogyny in of carpus, with sensory anterodistal spine; the species. carpusslender, elongate, only slightlyshort- Etymology.-The species is named for Dr. er than propodus, with 3 sensory spines on Thomas Iliffe of Texas A&M University, posterior margin, propodus with 4 sensory who collected this material. spines on posterior biun- margin;dactylus Remarks.-Members of the Gnathostene- guiculate. troididaeare marine forms known from the Pleopodal operculum of female, length about 1.25 times widest at Mediterranean,the western Atlantic, and greatest width, The about with 2 pairs of short lat- the eastern and western Pacific. five midlength, of the Stenober- eral setae, plus pair of submedian distal se- genera family, including muda for the first can be differentiated tae; precursorsof male second pleopods vis- time, on six characters.Since the re- ible beneath integument when stained and phylogenetic of the "lower"asellotes have cleared. Pleopod 1 of male covering second lationships yet to be these charactersare seen pleopods, but shorter and narrower than elucidated, as a convenient means of differen- broadly opercular exopods of pleopod 3. simply 2 of male with blunt- tiation, and are not regardedas necessarily Pleopod exopod short, of ly rounded,bearing slender distal spine;en- being phylogenetic significance. Gnathostenetroides Amar, 1957. Eyes dopod flexed, proximal part only slightly Hornlike of mandibular shorterthan distal, latter with complex dis- present. process tal structures short tube and body present. Mandibular molar distally including sperm truncate.Rostrum bifid. Antero- fringedplatelike lamella. Pleopods 3-5 sim- anteriorly ilar in male and female. 3 lateral processes of cephalon weak. Body Pleopod endopod ratio 3.7. about one-thirdwidth of exopod,just reach- length/width suture of with 4 distal Caecostenetroides Fresi and Schiecke, ing beyond exopod, 1968. absent. Mandible horn- plumose setae; exopod broadly ovate, with Eyes lacking transversesuture at about of me- like process. Mandibular molar distally midlength slender. Rostrum Anterolateral sial margin. Pleopod 4 endopod elliptical, lacking. about three-fourths of latter processes of cephalon weak. Body length/ length exopod; width ratio 7.0-8.8. with transversesuture, distal margin bear- 6 setae. 5 StenobermudaSchultz, 1979. Eyes pres- ing plumose Pleopod consisting ent. Mandible hornlike of single ovate ramus bearing 1 simple and lacking process. 1 seta Mandibular molar distally truncate. Ros- plumose distally. Uropodal protopod trum Anterolateral about two-thirds length of endopod, latter acutely triangular. pro- with several distal about half cesses of cephalon well produced. Body setae; exopod ratio 3.5-3.9. length and half width of endopod, with sev- length/width eral distal setae. MaresiellaFresi and Scipione, 1980. Eyes present.Mandible lacking hornlike process. Biology.-The 22 availablespecimens of this Mandibular molar distally truncate. Ros- species can be groupedaccording to size and trum anteriorly bifid or rounded. Antero- reproductivestructures: 8 individuals, 1.2- lateral processes of cephalon weak. Body 1.6 mm, females with pleopodal operculum length/width ratio 4.1-5.4. containing male pleopod 1 precursors;12 NeostenetroidesCarpenter and Magniez, individuals, 2.1-3.2 mm, primary males 1982. Eyes absent. Mandible lacking horn- with male pleopods;2 individuals, 2.9 mm, like process. Mandibular molar distally secondarymales with emptymarsupium and slender. Rostrum rounded. Anterolateral male pleopods. processes of cephalon weak. Body length/ This sequence of protogynoushermaph- width ratio 2.5. KENSLEY: MARINE ISOPODS FROM BERMUDA 335

The Walsingham Cave material differs Hooker, A. 1985. New species of from the from the of the Stenobermuda Florida Middlegrounds(Crustacea: ).- type genus, Proceedingsof the BiologicalSociety of Washington acutirostrata Schultz, 1979, known from 98: 255-280. Castle Roads, Bermuda,and the Turks and Kensley, B. 1976. Isopodan and tanaidaceanCrus- Caicos Islands, in a number of features:the tacea from the St. Paul and Amsterdam Islands, earlier species appearsto be larger(oviger- southern Indian Ocean.-Annals of the South Af- ous 9 4.0-4.8 mm versus 2.9 the ros- rican Museum 69: 261-323. mm), 1982. Anthuridea (Crustacea:Isopoda) of trum is more narrowly acute, and the an- CarrieBow Cay, Belize.-In: K. Riitzler and I. G. terolateral processes of the cephalon are Macintyre,eds., The Atlanticbarrier reef ecosystem more produced, while the antennule con- at Carrie Bow Cay, Belize, I: structureand com- sists of 7 (versus 6) articles; further, the munities. Pp. 321-353.-Smithsonian Contribu- to the lateral tooth is tions to MarineSciences 12: 1-539. pleotelson posterior , and M. Schotte. 1989. Guide to the marine relatively narrowerin the earlier species. isopod crustaceansof the Caribbean.-Smithsonian It is probable that several species de- Institution Press, Washington,D.C., and London. scribed under Stenetrium are actually rep- Pp. 1-308. resentativesof Stenobermuda.Examples of Kussakin,O. G. 1967. Fauna of Isopoda and Tan- these include Stenetrium acutirostrum aidacea in the coastal zones of the Antarctic and Subantarcticwaters.-Biological Reportsof the So- Muller, 1991b, from the Gulf of Aden; S. viet AntarcticExpedition (1955-1958) 3: 220-389. bourboniMuller, 1991c, from Reunion Is- [Englishtranslation by the Israel Programfor Sci- land; and S. temae Muller, 199 Id, from the entific Translations,Jerusalem, 1968.] Society Islands. Leach, W. E. 1814. Crustaceology.--In:Brewster's EdinburghEncyclopedia 7: 383-439. ACKNOWLEDGEMENTS Menzies,R. J., and P. W. Glynn. 1968. The common marineisopod Crustaceaof PuertoRico: a handbook The bulk of the materialreported in this paperwas for marinebiologists.-Studies on the Faunaof Cu- collected by E. A. Lazo-Wasemand M. F. Gable of racao and other CaribbeanIslands 27(104): 1-133. the PeabodyMuseum, Yale University, assisted by S. Muller,H.-G. 1989. Munnogoniumpolynesiensis n. Lasch, A. J. Baldinger,R. Denette, J. Mennone, J. sp. fromcoral reefs at BoraBora and Moorea,Society Damon, C. Booth,and J. Hamilton;E. A. Lazo-Wasem Islands(Isopoda: : Paramunnidae).-Bulle- loaned type material for comparative purposes; the tin Zoologisch Museum, Universiteit van Amster- materialfrom WalsinghamCave was collected by T. dam 12: 57-62. Iliffe of Texas A&M University; to all of these, my . 199la. MarineAnthuridea from Martinique, sincere thanks for making the material available for FrenchAntilles, with redescriptionsof some species study. I am gratefulto George H. F. Wilson of the (Crustacea:Isopoda).-Revue Suissede Zoologie98: AustralianMuseum, Sydney, for sharinghis valuable 739-768. comments on early notes and figures,and for his per- 199 b. Four new of shallow-water observationson asellote species ceptive systematics. Asellota from the Gulf of Aden.-Senckenbergiana Maritima21: 205-214. LITERATURECITED .1991c. Stenetriidaefrom coral reefs at Re- union Island,southern Indian Ocean. Description of Amar, R. 1957. Gnathostenetroideslaodicense nov. three new nov. nouveau d'Asellotaet classifica- species (Crustacea:Isopoda: Asellota).-- gen. sp. Type SenckenbergianaBiologia 71: 303-318. tion des Isopodes Asellotes.-Bulletin de l'Institut 1100: 1-10. 19 'd. The marine isopod family Stenetri- Oceanographique idae from ,oral reefs at Bora Bora and So- Carpenter,J. H., and G. J. Magniez. 1982. Deux Moorea, asellotes des Indes Occidentales:Neoste- ciety Islands, with descriptionsof four new species stygobies Suisse de 98: 51-76. netroidesstocki n. gen., n. sp., et Stenetriumsp.- (Crustacea).-Revue Zoologie Bijdragentot de Dierkunde52: 200-206. Nordenstam,A. 1933. MarineIsopoda ofthe families Fresi, E., and U. Schiecke. 1968. Caecostenetroides Serolidae,Idotheidae, Pseudidotheidae, Arcturidae, ischitanum(Isopoda: Parastenetriidae). A new genus Parasellidae,and Stenetriidaemainly from the South and species from the Bay of Naples.- Pubblicazioni Atlantic.-Further ZoologicalResults of the Swedish della Stazione Zoologicadi Napoli 36: 427-436. AntarcticExpedition 1901-1903 3(1): 1-284. , and M. B. Scipione. 1980. Maresiellanomen Poore,G. C. B., and H. LewTon. 1988. Antheluridae, novum pro MaresiaFresi, 1973, a gnathostenetroid a new familyof Crustacea(Isopoda: Anthuridea) with asellote isopod from GalapagosIslands. Pp. 1-2 in new species from Australia.-Journal of Natural Museo Zoologico dell'Universita di Firenze, Edit. History 22: 489-506. Galapagos,Studi e Ricerche,Spedizione "L. Mares- Richardson,H. 1902. The marineand terrestrialiso- G.R.S.T.S." pods of the Bermudas,with descriptionsof new gen- George,R. Y., and J.-O. Stromberg. 1968. Some new era and species.-Transactions of the Connecticut speciesand new recordsof marineisopods from San Academy of Sciences 11: 277-310. Juan Archipelago, Washington, U.S.A.--Crusta- Schiecke,U., and E. Fresi. 1972. Record of the asel- ceana 14: 225-254. lote isopod AustrosignumHodgson from the Bay of 336 JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 14, NO. 2, 1994

Naples:Austrosignum maltinii n. sp. (Paraselloidea, Vanh6ffen, E. 1914. Die Isopoden der Deutschen Munnidae).-Crustaceana,supplement 3: 31-38. Siidpolar-Expedition1901-1903.-Deutsche Siid- Schultz,G. A. 1969. How to know the marineisopod polar-Expedition1901-1903, 25 (Zoologie)7: 447- .-Wm. C. Brown Co., Dubuque, Iowa. 598. Pp. i-vii, 1-359. 1979. A new asellote (Stenetriidae)and two, RECEIVED:25 April 1993. one new, Anthuridea(Anthuridae) from Bermuda ACCEPTED: 18 June 1993. (Crustacea,Isopoda).- Proceedingsof the Biological Society of Washington91: 904-911. Address:Department of InvertebrateZoology, Na- 1986. Isopoda.-In: W. Sterrer,ed., Marine tional Museum of Natural History, Smithsonian In- fauna and flora of Bermuda.A systematicguide to stitution, Washington,D.C. 20560, U.S.A. the identificationof marineorganisms. Pp. 366-372. John Wiley and Sons, New York.

ANNOUNCEMENT

At its annual meeting in Los Angeles, California, in December 1993, The Crustacean Society bestowed the first TCS Service Awards on Denton Belk, Treasurer, and Arthur G. Humes, Editor, for their contributions over many years to the society.