Revision of Pylopagurus and Tomopagurus (Crustacea: Decapoda: Paguridae), with descriptions of new genera and species. Addendum and taxonomic summary

•The discovery of two new species, Enallopagurus provenzanoi, and EnallopaguropsLs williamsi, has required an addendum and concluding part to the revisionary study of the "Pylopagunis-Tomopagurus" group of genera. The diagnoses of Enallopagurus McLaughlin and Enallopaguropsis McLaughlin, are emended, and a corrected key to the genera is provided. The discovery of additional specimens of Phimochirus randalli (Provenzano) from the Gulf of Mexico has provided supplemental information on the morphology, color, and distribution of this uncommon species; it is rediagnosed and illustrated. Also provided is a summary of the original taxonomic assignments in part 1 (published in 1981) of this revisionary study, and the subsequent changes, corrections, deletions, and additions that have occurred since then. An updated list of the 13 genera, two subgenera, and 64 valid species currently recognized in the "Pylopagurus-Tomopagurus" group, is presented. McLaughlin (1981a) proposed the new genera Enallopagurus McLaughlin, 1981a and Enallopaguropsis McLaughlin, 1981a, for five relatively uncommon eastern Pacific hermit crab species that had been included by earlier authors in the genus Pylopagurus A. Milne-Edwards & Bouvier, 1891. To Enallopagurus, McLaughlin (1981a) assigned Pylopagurus affinis Faxon, 1893, P. coronatus (Benedict, 1892), and P. spinicarpus Glassell, 1938; Enallopaguropsis included P. guaternoci Glassell, 1937, and Pylopagurus hancocki Walton, 1954. One of the primary characters that McLaughlin (1981a) used to differentiate between Enallopagurus and Enallopaguropsis was the rows of scales on the propodal rasp of the fourth pereopod. In a subsequent, more detailed account of these two genera, McLaughlin (1982) placed Enallopaguropsis hancocki in synonymy with E. guatcinoci and added the new species E. janetae McLaughlin, 1982. In the present addendum and concluding part of the revision of the "Pylopagurus-Tornopagurus" group of genera, two new species, one each of Enallopagurus, and Enallopaguropsis, are described. Recent sampling off the Louisiana coast in the northern Gulf of Mexico has yielded a number of additional specimens of Phimochirus randalli (Provenzano, 1961), a species discussed by McLaughlin (1981b) and at the time known only from the Bahamas Islands and Caribbean region. Examination of these Gulf of Mexico specimens revealed a number of morphological details that complement previous definitions of P. randalli. Furthermore, Provenzano "s ( 1961:161 ) color description was incomplete as it was based on faded, preserved specimens, whereas for this study VOLUME 1 16, NUMBER 465 live specimens from the Gulf of Mexico were available and photographed (D. L. Felder, pers. comm.). Phimochints randalli is herein rediagnosed, and illustrated. Since the publication of part 1 (McLaughlin 198 La) of the six-part revisionary study of the "'Pylopagunis-Tomopagiinis" group of genera, several new species have been added that have necessitated emendations of generic diagnoses, and two new genera have been erected. Some of the taxonomic assignments made in part I have been changed in light of new evidence gleaned from direct examination of type materials not initially available. In some cases, new morphological discoveries, such as the existence of short sexual tubes in some species of Pylopagiinis sensu stricto (see McLaughlin & Lemaitre 2001), or the existence of short protrusions of the vas deferentia in the form of tubular papillae in Agaricochirus McLaughlin, 1981a, Enallopagunis. and Enallopaguiopsis (see McLaughlin 2003, this study), have required adjustments to our concept of these genera. In the new species of the latter two genera herein described, the morphology of the terminal margin of the telson has been foimd to deviate from the original generic diagnoses. As this maigin shape was used as a character in the updated generic key provided by McLaughlin & Lemaitre (2001:480), that key now requires corrections. Also, several taxa have been synonyniized since part 1 was published, whereas Pylopugurus Háchele Bai-nai^d, 1947, a taxon tentatively assigned in parts 1 and 2 (McLaughlin 1981a, 1981b) to Phimochirus McLaughlin, 1981a, was transferred by McLaughlin (1988) to Pagurus Fabricius, 1775, a genus not considered part of the ^'Pylopagunis-Toinopagiirus" group. McLaughlin's (J981a) genus Attstralerennis was reduced by de Saint Laurent & McLaughlin (2000) to a subgenus of Lophopagurus McLaughlin, 1981a. Thus, it was felt useful to have in a single convenient publication, a summai-y of the taxonomJc assignments, changes, corrections, and additions that have occurred during the 2Hyears that have elapsed since the appearance of part 1 of the revision of this group. There are cunently 13 genera, two subgenera, and 64 valid species known in this group (see Table 1). Materials used in this study are deposited in the National Museum of Natural History, Smithsonian Institution, Washington, D.C. (USNM), and University of Louisiana at Lafayette (ULL-Z). The specimens of the new species of Enullnpagurus and Enallopagurnpsis were collected by Dr. Anthony J. Provenzano, Jr. aboard the R/V John Elliott Pillsbury, of the Rosenstiel School of Marine and Atmospheric Science, University of Miami, Florida. The new material of Phimochirus randalli was collected by Dr Darryl L. Felder aboard the R/V Pelican, of the Louisiana Universities Marine Con.sortium, Cocodrie. The interpretation of the ocular acicle is used in accordance with Forest et a!, (2000), rather than that of Boyko & Harvey (1999). A single measurement, shield length (si), measured from the tip of the rostrum to the midpoint of the posterior margin of the shield provides an indication of animal size. Other abbreviations used are; ovig, ovigerous female; sta, station. Eiiallopagiinis McLaughlin, 1981a PylopagLirus: Walton, 1954:140 (in part).• Scanland & Hopkins, 1969:257 (in part) [Not Pytopagurus A. Milne-Edwards & Bouvier, 1891J. Enallopagurus MchsMghhn, 1981a:7; 1982: 843; 2003:122. Emended diagnosis.•Eleven pairs of biserial gills. Ocular acicle triangular or subovate, usually with strong submarginal spine. Sternite of third maxillipeds with spine on either side of midline. Third maxillipeds each with well-developed crista dentata, 1 accessory tooth; merus with or without dorsodistal spine. Maxillule with internal lobe moderately well produced, with 1 terminal bristle; external lobe weakly produced. 466 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON Table I.•List of valid names of "Pylopagiinis-Tomopagunis" group of laxa. Pylopagurus A. Mine-Edwards & Bouvier, 1891 sensu stricto P. discnicliilis (A. Milne-Edwards, 1880) P. holmcsi Schinitt, 1921 P. palliae Lemaiire & Campos, 1993 P, inacgeorgei McLaughlin & Lemaitre, 2001 P. (•orei McLaughlin & Lemaitre, 2001 Lophopagurus (Lophopagurus) McLaughlin, 1981a L. (L.) foresii McLaughlin & Gunn, 1992 L. (L.) laccriosus (Henderson, 1888) L. (L.) iKiiuis (Henderson, 1888) L. (L.) iiodiilosus McLaughlin & Gunn, 1992 L. (¿.) pumiliis de Saint Laurent & McLaughlin (2000) L. (L.) Ihompsoni (Filhol, 1885) Lophopagurus (Australelemus) McLaughlin, 19813 _ ., L. (A.) cookii (Filhol, 1883) L. (A.) eltaninae McLaughlin & Gunn, 1992 L. {A.) cristaius (H. Milne Edwards, 1836) L. (A.) kirkii (Filhol, 1883) L. (A.) laun-nlac (McLaughlin & Gunn, 1992) t. (A.) idi'vraMi'(Filhol, 1883) L. {A.) iriserrulus (Orlniann, 1892) Rhodochirus McLaughlin, 1981a R. rostireus (A. Milne-Edwards & Bouvier, 1893) R. ¡linimunus (Pa\on, 1893) Phimochirus McLaughlin, 1981a P. operculatus (Stimpson, 1859) P. californiensis (Benedict. 1892) ~ • P. vemistus (Bouvier, 1898) P. roseus (Benedict, 1892) ^ P. runilulli (Provenzano, 1961) P. Iiollhuisi (Provenzano, 1961) P. ocf/u.vw.v (Henderson, 1888) P. ¿t'li/'wa/pu.v McLaughlin, 1981b Haigia McLaughlin, 1981a i _ . H. diegensis (Scanland & Hopkins, 1969) Agancoc/i/n« McLaughlin, 1981a A. boletifer (A. Milne-Edwards & Bouvier, 1893) A. alexaiidri (A. Milne-Edwards & Bouvier, 1893) A. ctivimunus (Chace, 1939) A. erosHS (A. Milne-Edwaj'ds. 1880) A. gibbosimanus (A. Milne-Edwards, 1880) A. hispidas (Benedict, 1892) A. echiiwius McLaughlin, 1982 A. acaiuhinus McLaughlin, 1982 Anisopagurus Mchaoghlm, 1981a A. banleiH (A. Milne-Edwards, 1880) A. pygmoeus (Bouvier, 1918) A. vossi Lemaitre & McLaughlin, 1996 A. acliiwphorus Lemaitre & McLaughlin, 1996 A. hopkinsi Lemaitre & McLaughlin. 1996 VOLUMF. 1 16. NUMBER 2 Table I.•Cominuecl. 467 Eiuiltopatiiiriis McLaughlin. 1981a Ennllojniiiiiiopsis McLaughlin, lysia Miinucompliiims McLaughlin. 1981 a M. cen-iconii.s (Benedict. 1892) M. sininilosiis (Holthuis. 1959) M. Iun};iiiuiiiii.\ (Fi[\on. 189.^) M. i'¡(/7íí/).s (Benedict. 1892) M. unniiliiius (Studer. 1883) /•-•. spiiiirarpiis (Gla.ssell. 1938) IC. iiffinis (Faxon. 1893) /;". coioiHiliix (Benedict. 1892) [Í. /inni'iizciiiiii. new species /•-'. I'ikilemoci (Glassell. 1937) !•:. Jimelae McLaughlin. 1982 H. williüiiísi. new species 'l'i>nii>p(ii;iinis A. Milne-Hdwards & Bouvier. 1893 7'. nihyopiiiicliiliis A. Milnc-Hdwai'ds & Bouvier. 1893 T. cokeri (Hay. 1917) 7'. nuicltnii^hliniic Haig. 1976 r. nihensis (Wass. 1963) 7'. uvM.v/McLaughlin. 1981a T. piirpiiniuis (Benedict. 1892) I'. iiii'iiiiiiiciilDSiis (Glassell. 1937) r. rluui'i (Wass. 1963) Prnliiiiiopdiiunis l.einailre & McLaughlin, 1996 P. hiojH'rciiluliix Lemaiire & McLaughlin, 1996 Py!í>l>íiíi¡in'i/iiiiii McLaughlin & Ix'mailie. 2001 P. niiirklniini McLaughlin & Lemaiire. 2001 Right cheliped with subovate or subrectangiilar chela. Left cheliped with chela triangulaf in cioss-section, but dorsal siifface not elevated into pi'ominent keel or cre.st. Dorsodistal spine ol' caipi of second and thifd pcfeopods lacking. Stei'nite oC thifd pereopods with subcifculai", siibt)vatc or subquadrate anterior lobe. Slernites of third to fifth pereopods often with capsúlate setae. Fourth pereopod with propodal rasp consisting of single row of corneous scales; dactyl with small prcungual process usually present. Males with paired gonopores, no distinct sexual tubes hut occasionally with vas deferens protruded to form short tubular papilla, usually on right; without paired pleopods, with 3 uniramous or weakly biramous left pleopods on somites 3-5. Females with paired gonopores. paired first pleopods modified as gonopods, 4 unpaired left pleopods, second to fourth with both rami well-developed, fifth usually uniramous. Uropods symmetrical or asymmetrical. Telson without indication of division into anterior and posterior portions; terminal margin entire or with inconspicuous median indentation, unarmed. Remarks.•Faxon (1893, 1895; pi. 12. fig. 2e) reported that his single male specimen of Pylopcii^iinis uffinis (= Eiiallopagiirus ajßnis) had the vas deferentia extruded as "small tlireads" from both gonopores. McLaughlin (1982) reexamined Faxon's (1893) specimen and ccjncluded that while there were slight protrusions, these were more probably an artifact of preservation 46S PROCEHDINGS OF THF. RIOI.ÜGICAl. SOCIETY OF WASHINGTON ralher than actual, very small sexual tubes. Since McLaughlin's (1982) study, our information on and understanding of sexual tubes in the Paguridae has increased substantially. Among the "Pylopcigurii.s-Tomopa^unis" group, species of Enullopaf>iirnpxis and Aiiuricochirus can also have the vas deferentia protruded as tubular papillae, albeit not as a distinct sexual tube. In other Paguridae, similar protrusions are known to occur in genera such as Discorsopcii;iirii.s McLaughlin, 1974, Pagurilla Melin, 1939, and Pagiiriis (see de Saint Laurent 1970, McLaughlin & Lemaitre 199.3, McLaughlin 2003). However, as reported by McLaughlin & Lemaitre (2001), short but distinct sexual tubes are present in some species of Pylopcigiirus sensu stricto. Recently, McLaughlin & Jensen (1996) found a very small sexual tube developed on the right coxa of males of Parapcii>iirock's liartac McLaughlin & Jensen, 1996, and Komai ( 1998) provisionally transferred two Japanese species formerly assigned to Pai^unis {P. gracilipe.s Stimpson, 18.58 and P. nipponensis Yokoj'a, 1933), to Parupagitrocli's McLaughlin & Haig, 1973, because of the observed small .sexual tubes in males of both species. McLaughlin (1981a, 1982) included Faxon's (1893) Pylopagunis ajfuiis in Enallopagurus. despite the fact that Faxon described the propodal rasp of the fourth pereopod as "multiserial". The type specimen has been reexamined, and Faxon's description and subsequent illustration (Faxon 189.5: pi 12, fig. 2d) found to be in error; the propodal rasp of the fourth pereopod actually has a single row of scales. This species is retained in Enallopagiinis. although this generic assignment is still provisional given that E. affinis is known only from the male hololype. Until the discovery of the new species of Enallopagurus described herein, the genus had been considered exclusively an eastern Pacific taxon. Although the right cheliped of this new species lacks the operculate-like conformation seen in the other three species assigned to the genus, the unarmed carpi of the second and third pereopods, the single row of scales on the propodal rasp of the fourth pereopod, and the presence in females of paired first gonopods, support its placement in the genus. The terminal margin of the telson in this new species of £";?allopagunis has a minute tnedian indentation and is not clearly entire as described in the original generic diagnosis (McLaughlin 1981a. 1982); thus, the need for the emendation. An updated key to the species of Enullopagitnis is provided following the description of this new species. Enallopagurus provenzanoi, new species Figs. 1, 2 Holotypc.•Ovig. ? (si = 2.40 mm), R/V John Elliot! Pillshiiry, sta 581, 2l°05'N, 86''23'W, Arrowsmith Bank, 146265 m, 22 May 1967, USNM 1007525. Paraiype.•1 â in poor condition (si = 1.80 mm), same data as holotype. USNM 1007526. Descripiion.•Shield (Fig. la) longer than broad; anterior margin between rostrum and lateral projections slightly concave; anterolateral margins sloping; posterior margin truncate, dorsal surface glabrous. Rostrum triangular, terminating acutely. Lateral projections rounded, unarmed. Ocular peduncle approximately 0.65 length of shield, moderately stout, with cornea slightly dilated; ocular acicles subtriangular, each terminating subacutely, with moderately well-developed submarginal spine (not always visible in dorsal view); separated basally by approximately basal width of 1 acicle. Antennular peduncle overreaching ocular peduncle by 0.35-0.50 length of ultimate segment; ultimate and penultimate segments unarmed; basal segment with small spine on lateral face. Antennal peduncle slightly shorter to equaling length of ocular peduncle. Fifth and fourth segments with scattered short seVOLUME 116. NUMBER 2 469 Fig. I.•Eiiallapiiaitni.s i>r(>\'í'n-.iiru>i. new species, hololypc ovig 9 (si = 2.40 mm), norihvvestern C;ii'ibbe;in, R/V .IDIIII Ellioti I'illshiiry. sl;i .'»SI. USNM ll)07.')2.'i. a. shiekl and cephalic appendages, dorsal; b. righi eheliped. doi'sal; c. chela of same, mesial; d. led eheliped. dorsal; e. chela of same, mesial; I', telson, dorsal. Scales equal (.).?>() mm (a). 1.00 mm (b. c). and {).?• mm (d-f). 470 PROCHK13INC;S Ol' THü BIOI.OGICAI. SOClirCY OlWASHINGTON tae. Third segment with acute spine at ventrodistai angle. Second segment with dorsolateral distal angle produced, terminating in small spine; dorsomesial distal angle with small spine. First segment produced ventrally and with acute spine on ventrolateral margin. Anlcnnal acicle moderately short, reaching only to distal margin of fourth peduncular segment. Flagellum long, with 1 or 2 short setae every several articles. Sternite of third maxillipeds with strong spine on cither side of midline. Right cheliped (Figs. lb. c) elongate, slender, overreaching left by length of dactyl. Dactyl approximately 0.75 length of palm, with row of small spines on proximal half of dorsomesial margin, separated from similar short row near cutting edge by shallow sulcus; cutting edge with 1 prominent double tooth in proximal half and single prominent tooth in distal half; mesial and ventral surfaces with lew short setae. Palm exceeding carpus by approximately 0.20 own length; dorsal surface convex, unarmed, but with sparse tufts of setae, dorsomesial and dorsolateral margins not delimited; mesial, lateral and ventral surfaces also with sparse tufts of setae; hxed finger with row of low spines on dorsolateral margin and second row adjacent to cutting edge; latter with 2 or .1 small and 1 large broad calcareous teeth. Carpus slightly shorter than merus, subtriangular; dorsomesial and dorsolateral margins not delimited, surfaces unarmed but with scattered setae, I prominent spine on distomesial margin. Merus triangular, margins and surfaces unarmed, but with scattered short setae. Ischium unarmed. Left cheliped (Figs. Id, e) with dactyl approximately as long as palm; surfaces of dactyl, (ixed finger and palm all unarmed but with sparse tufts of setae. Carpus only slightly shorter than merus; surfaces of both segments and ischium unarmed but with scattered sht)rl to moderately long setae. Second and third pereopods (Figs. 2a, b) long and slender. Dactyls longer than propodi; dorsal surfaces with few short setae; ventral margins each with row of 10-12 corneous spines and few short setae. Propodi approximately twice as long as carpi; dorsal surfaces with few setae; ventral margins each with 3 or 4 corneous spines, longest on third pereopod. Carpi less than half length of meri, unarmed, but with scattered setae dorsally and ventrally. Meri and ischia unarmed, but with dorsal and ventral sparse tufts of setae. Foiuth pereopod (Fig. 2d) with small preungual process on dactyl. Sternite of third pereopods with reduced, slender anterior lobe. Sternites of fourth and fifth pereopods with capsúlate setae (Fig. 2c). Telson (Fig. If) without indications of division into anterior and posterior portions; terminal margin broadly rounded, with inconspicuous median indentation, unarmed. Color.•Overall body and appendage color very pale yellow. Ocular peduncles with white and yellow chromatophores and dark red longitudinal stripe on ventromesial and ventrolateral surfaces forming an elongate U-shaped pattern; corneas black with "frosted" overtone. Chelipeds each with fevii scattered orange spots. Dactyls of second and third pereopods with 2 extremely faint yellow bands, most apparent on .second. (From A. J. Provenzano, Jr., field notes). Affinities.•This species, known from only two specimens, bears certain habitat and morphological similarities to Fylopagiini.s paltiae Lemaitre & Campos, 1993. Both have been found living in membranous tubes and both have similarly elongate and weakly armed chelipeds. However, in addition to differences in generic characters, E. provenzcinoi is easily distinguished from P. pattiae by the armature of the dactyl and fixed finger of the right chela, greater number oí spines on the ventral margins of the dactyls of the second and third pereopods, unarmed ambulatory carpi, and structure of the telson. Habitai.•The holotype was found living in a semi-transparent polychaete tube. VOLUME 1 16. NUMBIiR 2 471 Fiy. 2.•E¡uiUo¡Ki¡>u¡-iis pri>Ví'¡i:üiu>i, new specics, lioiotype oviy 9 (sl = 2.40 iniii). nonliwcslcni CiiiibbcMii. R/V Jithu Ellioii Pill.\hiiiy. sui .'581. USNM l()()7.'i2.'i. ;i. Icli second pcrcopiul. Uilenil; b. led lliial percopoü. laienil; e. coxae and sternile oC (il'lli percopod.s vviih c;ipsiil;ilc .sctao (siippled). and pari ol' abdomen (lower) showing (h'sl pleopods: d. pi'opodiis and daelyl ol' lel'l Ibuith pcreopod with preungual process (arrow), laleral. .Scales equal 1.00 mm (a. b). 0.50 mm (c), and 0.10 mm (d). DisirH)iiiioii.•Known at present only IVom the type locality, olT Arrowsmith Bank, in the northwestern Caribbean; 14626.'S m. Elyniology.•This species is named for Dr. Anthony J. Proven/ano. Jr., whose pioneering work on western Atlantic hermit crabs proviiJed the basis for the study of the "Pvlopaiiuru.s-TomDpa^urus" group of genera. Remarks.•As previously indicated, Enullopui^uni.s proveuz.iinoi, new species, is the first recogni/ed representative of the genus in the western Atlantic. Unlike some species OÏ Phiinochinis (another of the "Pylopagiinis-Tomopngiirus'' group of genera), which is represented in both oceans by geminate pairs (cf. McLaughJin 1981b), E. prnvenzunoi appears to be related to the two eastern Pacilic species only through major generic characters. Key to Species of Eiuillopaiiiinis 1. DorsopiDximal margin of right chela armed with strong spines; eastern Pacilic E. roroiialus Dorsoproximal margin of right chela unaniied 2 2. Dorsal siul'ace of palm ol' right chela with relatively closely-spaced small spines or spinulcs; eastern Pacilic E. affini.'i Dorsal surface of palm of light chela unarmed or rarely slightly tiiberculatc ... 3 ,^. Carpus of right eheliped with two or three spines on dorsomesial margin separated by concavity IVom prominent tlorsodislai spine; eastern Pacific E. .spinicai¡>u.'i 472 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OH WASHINGTON Carpus of lighi cheliped with unarmed dorsomesial margin; Caribbean Sea . . . E. provenzunoi. new species Enallopagiiropsis McLaughlin, 1981a Pylopcifiiinis: Walton. 1954:140 (in part).• Scanland & Hopkins, 1969:257 (in part) I Not Pylopiifiunis A. Milne-Edwards & Bouvier, 18911. Eiiullopciiiiiiüpsis McLaughlin 1981a:7; 1982:848; 2003:12L Eiucijiled í//íí,í>;io,v/.v.•Eleven pairs of biserial gills. Ocular acicle acutely triangular, with moderate to well-developed submarginal spine. Sterniie of third maxillipeds with prominent spine on either side of midline. Third maxilliped with well-developed crista dentata and 1 accessory tooth. Maxillule with internal lobe of endopod moderately well-developed, and with I stiff bristle terminally; posterior lobe somewhat produced, not recurved. Right cheliped with suboperculate chela. Left chela subtriangular in cross-section, but dorsal surface not produced into prominent keel or crest. Dorsodistal spine of carpi of second and third pereopods lacking. Anterior lobe of sternite of third pereopods represented by 1 or 2 capsúlate selae. Sternites of fourth and fifth pereopods often with 1 to several large capsúlate .setae. Fourth pereopod with propodal rasp consisting of multiple rows of corneous scales. Males with paired gonopores, no distinct sexual tubes but occasionally with vas deferencia produced to form short tubular papillae on one or both coxae; no paired pleopods; 3 unpaired left pleopods with endopods reduced or absent. Females with paired gonopores, paired first pleopods modified as gonopods, 4 unpaired left pleopods, second to fourth with both rami well-developed, fifth with endopod reduced or absent. Abdomen straight or slightly fiexed, usually moderately long; uropods symmetrical or asymmetrical. Telson without indication of division into anterior and posterior portions; terminal margin convex, entire or with shallow median concavity, unarmed. Remarks.•The original diagnosis of Enaliopcii^iiropsis indicated that the terminal margin of the telson was entire in all the species. In the new species described herein, however, the terminal margin is divided into nearly symmetrical lobes by a shallow median concavity, and the diagnosis of the genus has been corrected accordingly. With the addition of the new species of Enallopa(>itr()psis described herein, the genus is now represented by two eastern Pacific and one western Atlantic species. An updated key to the species is provided following the description of this new species. Enallupaguropsis williamsi, new species Figs. 3, 4 Holotype.•â (si = 1.70 mm), R/V John Elliott Pillshtiry. sta 421, ()9°32.rN, 78°33.5'W, Golfo de San Blas, Panamá, 5358 m, 19 Jul 1966, USNM ] 007523. Description.•Shield (Fig. 3a) longer than broad; anterior margin between rostrum and lateral projections slightly concave; anterolateral margins sloping; posterior margin truncate, dorsal surface with few setae. Rostrum triangular, with terminal spinule. Lateral projections triangular, each with terminal spine or spinule. Ocular peduncle approximately 0.85 length of shield, moderately stout, with cornea slightly dilated. Ocular acicles subtriangular, each terminating subacuiely, with well-developed submarginal spine; separated basally by slightly more than basal width of I acicle. Antcnnular peduncle overreaching ocular peduncle by approximately 0.50 length of ultimate segment; ultimate and penultimate segments unarmed; basal segment unarmed. Antennal peduncle slightly shorter to equaling length of ocular peduncle. Fifth and fourth segments with scattered setae. Third segment with acute spine at ventrodistal angle. Second segment with dorsolateral distal angle prominently produced, terVOLUME I 16. NUMBER 2 473 Fig. 3.•Enall(>piii;iiropsis williamsi. new species, holotype S (si = 1.70 mm), soiillivvcstcrn C;irJbbe;in. R/V John lillioll Pillshuiy. sla 421. Golfo de S;\ii Bhis, PLinamá. U.SNM I()ü7.i23. ;i. shield ¡nul ecphiiiic appendages (riglit antennal llagellum missing), dorsal; h. right chclipcd. dorsal; e. chela of same, mesial; d. same, lateral; c. left cheliped. dorsal; I', chela of same, mesial; I', lelson. dorsal. Scales equal O.-^iO mm (a-d. g) anil 0.25 mm (e. f). juinating in small .spine; dorsomesial distal inargin of fourth peduncular segment. Flaangle with small spine. First segment progellum with short setae s; I article in length duced ventrally and with acute spine on in inost articles, and moderately long setae ventrolateral margin distall}'. Antennal aci2 or 3 articles in length every 2 articles. ' ele moderately short, reaching only to distal Right cheliped (Figs. 3b-d) with subop474 PROCEEDINCIS OH THK BIOLOGICAI. SOCIHTY OF WASHING TON erculate chela. Dactyl approximately Ü.80 length of palm, approximately equal to fixed finger in width, with row of spines on distal 0.7.'S of elevated dorsomesial margin, dorsal sin-face with few small tubercles in distal half; culling edge with row of moderatel}' large teeth; mesial, lateral and ventral surfaces with scattered short setae. Palm slightly exceeding length of carpus; dorsal surface weakly convex, armed with only 1 small spine slightly mesiad of midline in proximal half, elevated dorsomesial margin forming spinose ci-est, dorsolateral margin also elevated as low ridge proximally becoming more pronounced and weakly serrate distally and on fixed linger; cutting edge of lixed linger with row of distinct teeth; mesial, lateral and ventral surfaces with sparse tufts of setae. Carpus approximately equal to length of merus, subIrape/.oidal; dorsomesial margin with row of 4 prominent spines, dor.solateral margin not delimited, dorsal surface with longitudinal row of 2 spinose protuberances proximally, otherwise surfaces unarmed, but with few scattered setae. Merus triangular, margins anti surfaces unarmed but with scattered, short setae. Ischium unarmed. Left chcliped (Figs, ."^e, f) with dactyl approximately as long as palm; surfaces of dactyl and fixed ñnger unarmed but with .sparse ti[fts of setae, dorsal surface of palm with 2 small spine.s in midline proximally. Carpus only slightly shorter than merus; dorsal suiface with longitudinal row of 2 spines medially, and 2 additional spines on distal margin; mesial, lateral and ventral surfaces unarmed but with few scattered .setae. Merus and ischium with only scattered short to moderately long sclae on all surfaces. Second and third pereopods (Figs. 4a, b) moderately long and slender. Dactyls equal to or slightly longer than propodi; dorsal surfaces with few short setae; ventral margins each with row of 6 or 7 long corneous spines and few short setae. Propodi approximately \ .50 to nearly twice as long as carpi; dorsal surfaces with few setae; with I or 2 ventrodistal spinilbrm bristles, and I or 2 additional slightly shorter spiniform bristles on each ventral margin distally. Carpi 0.45-0.75 length of meri, unarmed, but wilh scattered setae dorsally and ventrally. Meri and ischia unarmed, but with dorsal and ventral sparse tufts of setae. Fourth pereopod (Fig. 4d) with propodal rasps consisting of 3 rows of corneous scales. Dactj'l lacking preimgual process. Sternites of third, fourth, and fifth pereopods (Fig. 4c) each with 3, 4, and 2 capsúlale setae, respectively. Coxae of fifth pereopods with vas deferentia slightly produced forming short tubular papillae on right and left sides (Fig. 4c). Uropods nearly symmetrical. Telson (Fig. 3g) witln)ut indications of division into anterior and posterior portions; terminal margin with broad median concavity, unarmed. Color.•Shield weakly mottled orange with large darker orange spot on each side laterally and few smaller spots and patches in distal half. Ocular peduncles mottled light orange with scattered darker orange specks. Antennal peduncles with band of light orange on second segments superimposed with few darker orange specks; antennal acicles wilh few patches of light orange. Chelipeds each generally light orange with few scattered darker orange patches; dactyls each with proximal darker orange band. Dactyls of second and third pereopods each with orange band in proximal half; propodi with scattered orange spots and oi'ange band al mid-length; carpi very light (Hange, that of third wilh at least scattered darker orange specks; meri faint orange, darker proximally and with numerous darker orange specks. (From notes by B. Stolen, artist for A. J. Proven/.ano. Jr.). Etynioloi>y.•This species is dedicated to the memory of Dr. Austin B. Williams (1919-1999, see Lemaitre & Collette 2Ü()Ü), esteemed colleague who during many years of work al the Systematics Laboratoiy. National Marine Fisheries Service, contributed greatly to our knowledge of western Atlantic decapod crustaceans. VOLUME 116, NUMBER 2 475 ¡'¡g. 4.•li>it¡lli>pii!^iiropsis n-illi{ii>isi. new species, holotype ó" (si = 1.70 min), soiithweslcrn Cnribheiin, R/V John Ellioti PWsbiiry. sta 421. Golfo de San Blas. Panamá, U.SNM 1007.52.^. a. left second pcreopod, lateral; h. left (liii'd pereopod. lateral; c. coxae and slernitcs ol' first to (iilh pereopods with capsiilaie setae {stippled), and protruded vas del'erens (vd), ventral; d, propodus and dactyl of left fourth pereopotl, lateral. .Scale equals ()..5() mm (a-c) anil 0.15 mm (d). Habitat.•Hou.sing unknown. Bottom whei'e found consisted of "cofalline plates and broken shells" (Voss 1966). Distribution.•Known only írom the type locality, GoU'o de San Bias, on the Caribbean coast of Panama. Remarles.•Because Enullopaguropsis willidin.si, new species, is known fVom only the male holotype, this species can only provisionally be assigned to Eiiallopui'iirop.sis. The tclson of E. williumsi bears some resemblance to species of Agaricochiru.s. However, the morphology of the right cheliped and propodal rasp suggests that this new species belongs in Enallopaguropsi.s. Key to Specie.s ol' EtuiHopugiiropsis 1. Palm of left chela with row of spines on dorsolateral martiin 2 476 PROCKKDINGS OF THK BIOLOGICAL SOCIBTY OlWASHINGTON Palm of left chela with unarmed dorsolateral margin; Caribbean Sea E. williiiinsi, new species 2. Doi'sal siirlace ol' right chela covered with closely-spaced corneous-tipped spinules or granules; eastern Pacihc . . . E. ¡•¡Hilcmoci Dorsal surface of lighi chela with scattered tubercles; eastern Pacific E. juneldc Phimochints nindalli (Proven/.ano, 1961) Figs. 5-7 Pylopagunis raiiclalli Proven/ano, 1961; 159, fig. 2 (type locality: ridge 5 miles süLitheasl of Lameshur Bay, St. .lohn. Virgin l.slands). FhiDiocliirus ranclalU: McLaughlin, 198 i a: 5; 19Slb:34(). (igs. 4b, 5b. 7b.•Williams el al.. 1989:32.•McLaughlin et al., 2003:125. Holoiype.^â (si = 6.20 mm), SE Lameshur Bay, St. John, Virgin Islands, 20 m, 5 Feb 1960, colls. J. Randall and T. Chess, USNM 106353. Material examined.•Northern Gulf of Mexico, R/V Pelican, U.S. DepaitmeiU of Energy project; 1 <5 (s! = 6.70 mm) + exuvia, 27°48.717'N, 93°02.880'W, dredged at night, calcareous rubble, 16 Aug 1998, USNM 276157 (photo voucher).• 1 9 ovig (.si = 5.9 mm), dredge sta 10-3, 28°05.999'N, 91°02.289'W, 57 m, 27 May 2000, ULLZ 4678 (photo voucher).• ! S (si = 2.80 mm), dredge sta 7-2, 91°02.48I'W, 57-60 m, 1 JuÍ2001, ULLZ 4551 (photo voucher). Diagnosis.•Shield (Fig. 5a) longer than broad; dorsal surface with several short transverse or oblique rows of bristle-like setae; rostrum proininent. acutely triangular; with minute terminal spine; lateral projections obsolete or broadly triangular, usually with minute terminal spine. Ocular peduncles stout, about 0.60 times length of .shield, with corneas strongly dilated. Ocular acides elongate, acutely triangular; dorsal surfaces concave distaliy; each usually with small, submarginal spine terminally; separately basally by basal width of 1 acide. Antcnnular peduncle exceeding distal margin oi' cornea by about 0.30 length of ullimale segment. Antennal peduncle exceeding distal margin of cornea by about 0.30 length of fifth segment; second .segment with dorsolateral distal angle produced, terminating in strong spine, dorsomesial distal angle with small spine; first segment unarmed; acide elongate, weakly arcuate, reaching to about midpoint of fifth antennal segment, terminating in spine, unarmed. Antennal flagella inodcratcly long; vvilh short setae every 3 or 4 articles, each seta about I Hagellar article in length. Right cheliped (Fig. 6) sparsely setose. Chela operculate. dorsoventrally compressed (more markedly .so laterally and mesially); dorsal surfaces t)f palm and fingers covered with low. mushroom-like tubercles replaced medially on palm by 2 longitudinal, rectangular or ovate granular patches separated by narrow tuberculate area; patches sjightly depressed, with granules arranged in irregular, transverse rows. Fixed finger broad basally. with up to 6 blunt spines dorsally more prominent than surrounding tubercles. Dactyl about as long or slightly shorter than length of palm; mesial margin with blunt spines; dorsal surface with 2 (1 distal, 1 proximal) blunt spines standing out from sunounding tubercles. Carpus with granules or low tubercles on dorsal, lateral and mesial surfaces, dorsoniesial margin with row of irregular, corneous-tipped spines; mesial surface concave distaliy. Merus with row of small spines or spinulose tubercles on ventromesial and ventrolateral margins; dorsodistal margin with row of 4-6 corneous-tipped spines. Left cheliped (Fig. 5b) with surfaces of carpus and chela usually covered with small, flattened tubercles or granules. Palrn with longitudinal, often ridge-like elevation in midline armed with row of spines or spinules, and flanked on each side by subrectangular or ovate area nearly smooth or with VOLUME I 16, NUMBF.R 2 477