2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 233

Fig. 75. Albunea speciosa Dana, 1852: A, (, 9.0 mm cl, USNM 260868, neotype; B±J, (, 10.4 mm cl, USNM 287087. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible; mesial view. F. Left maxillule, lateral view. G. Right maxilla, lateral view. H. Left maxilliped I, lateral view. I. Right maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 1.6 mm (B, E, F), 2.2 mm (I), and 3.3 mm (A, C, D, G, H, J). gins. Segment II medially in¯ated in dorsal gose with long plumose setae; long plumose view, with plumose setae on dorsal and ven- setae on dorsal and ventral margins. tral margins, and scattered on ventrolateral Antenna (®g. 75D) with segment V ap- third of surface. Segment I wider than long, proximately three times longer than wide, unarmed; dorsal third of lateral surface ru- with long plumose setae on dorsal and ven- 234 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 tral margins; ¯agellum with ®ve or six arti- wide, with long simple setae on dorsal mar- cles, long plumose setae on dorsal, ventral, gin. Merus approximately three times longer and distal margins. Segment IV expanded than wide, margins parallel, with simple se- distally, with long plumose setae on dorsal, tae on ventrolateral margin and plumose se- ventral, and distal margins, and row of setae tae on dorsolateral margin. Basis-ischium in- on dorsolateral margin. Segment III with completely fused, with plumose setae on long plumose setae on ventral margin. Seg- margins. Exopod one-fourth longer than mer- ment II short, widening distally, with plu- us, ¯agellum with one article. mose setae on margins; antennal acicle long, Maxilliped III (®g. 75J) dactylus evenly thin, not exceeding distal margin of segment rounded; with long plumose setae on margins IV, with long plumose setae on dorsal mar- and lateral surface. Propodus with longitu- gin. Segment I rounded proximally, ¯attened dinal median row of plumose setae on lateral ventrolaterally, with long plumose setae on surface; margins with plumose setae. Carpus margins; lateral surface with acute spine dor- slightly produced onto propodus; lateral sur- sally, with low semicircular dorsolateral lobe face with row of plumose setae ventromedi- ventrodistal to spine; segment with ventro- ally; plumose setae on margins. Merus un- mesial antennal gland pore. armed, with plumose setae on margins. Ba- Mandible (®g. 75E) incisor process with sis-ischium incompletely fused, without cris- one tooth; cutting edge with one tooth. Palp ta dentata. Exopod two-segmented: proximal three-segmented, with plumose setae on mar- segment small; distal segment styliform, ta- gins and long, thick, simple setae arising pering, approximately one-third length of from bend in second segment. merus, with plumose setae on margins; with- Maxillule (®g. 75F) distal endite proxi- out ¯agellum. mally narrow, widening to in¯ated distal end, Pereopod I (®g. 76A) dactylus curved and with thick simple setae on distal margin. tapering; lateral and mesial surfaces smooth; Proximal endite with thick simple setae on dorsal margin with long plumose and short distal margin. Endopodal external lobe trun- simple setae; ventral margin with short sim- cate distally and curled under; internal lobe ple setae. Propodal lateral surface with nu- reduced with three thick setae at distolateral merous short, transverse rows of setose ru- margin. gae; dorsal margin unarmed; ventral margin Maxilla (®g. 75G) exopod evenly rounded, distally produced into acute spine; cutting with plumose setae along distal margin. Sca- edge lacking teeth, lined with long plumose phognathite bluntly angled on posterior lobe, setae; dorsal margin with long plumose setae, with plumose setae. ventral margin with short simple setae. Car- Maxilliped I (®g. 75H) epipod with plu- pus with dorsodistal angle produced into mose setae on distal margin and on distola- strong corneous-tipped spine, dorsal margin teral surface. Endite tapered distally and sub- with few large and small spines posteriorly equal to ®rst segment of exopod. Exopod along distal third; dorsal and distal margins with two segments: proximal segment nar- with long plumose setae; lateral surface with row, margins parallel, margins with plumose small distal rugose area, with few transverse setae; distal segment spatulate, approximate- setose ridges on distal half of surface; mesial ly as long as wide, broadest medially, mar- surface smooth with few scattered rows of gins with long plumose setae. Endopod ¯at- long plumose setae, margins with long plu- tened and elongate, reaching to distal end of mose setae. Merus unarmed; lateral surface proximal exopodal segment, with plumose with scattered transverse rows of long plu- setae on margins. mose setae, margins with long plumose se- Maxilliped II (®g. 75I) dactylus evenly tae; mesial surface with few short rows of rounded, length equal to width, with thick setae; proximal quarter of mesial surface simple setae distally. Propodus one-half wid- with decalci®ed window. Basis-ischium in- er than long, with plumose setae on dorsal completely fused, unarmed. Coxa unarmed. margin and long simple setae on distal mar- Pereopod II (®g. 76B) with dactylus gin. Carpus not strongly produced dorsodis- smooth; with base to heel straight, heel tally, approximately two times longer than smoothly rounded, heel to tip with rounded, 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 235

Fig. 76. Albunea speciosa Dana, 1852: A±E, (, 10.4 mm cl, USNM 287087; F, (, 9.0 mm cl, USNM 260868, neotype; G, &, 14 mm cl, USNM 287087. A. Right pereopod I, lateral view. B. Right pereopod II, lateral view. C. Right pereopod III, lateral view. D. Right pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 3.0 mm (F, G), 4.0 mm (A±D), and 4.2 mm (E). broad indent, tip acute, tip to base broadly face; distal and ventral margins with long convex; lateral surface smooth, with several plumose setae; dorsolateral surface as nar- small tufts of short setae in generally straight row, oblique, ¯attened shelf, with short setae line across medioproximal surface, several on dorsal margin and long plumose setae on widely spaced submarginal tufts of short se- ventral margin; mesial surface with elevated, tae dorsodistally; mesial surface smooth, curved, setose ridge from ventral junction ventral margin with long plumose setae, dor- with dactylus almost to ventral proximal sal margin with short simple setae and patch junction with carpus; decalci®ed region just of long plumose setae at base. Propodus with distal to junction with carpus. Carpus slightly dorsal surface smooth, ventral margin in¯at- produced dorsodistally; lateral surface nearly ed and rounded; oblique row of long plu- smooth, with irregular, interrupted row of ru- mose setae on distal margin of lateral sur- gae and submarginal elevated ridge ventrally, 236 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 rugae and ridge with long plumose setae; imally; lateral surface smooth, ventral mar- dorsodistal projection with mat of short setae gin with long plumose setae, dorsal margin on lateral surface; margins with long plu- with short simple setae; mesial surface with mose setae; mesial surface smooth, with long dorsal decalci®ed region, demarcated ven- plumose setae in scattered patches on surface trally by longitudinal elevated ridge with row and on margins. Merus with few scattered of short setae; with setose punctae ventral to setae on lateral surface and margins; mesial decalci®ed window. Propodus expanded dor- surface nearly smooth, with few setae and sally and ventrally; ventral expansion ex- decalci®ed area on proximal fourth near ceeds ventral margin of dactylus, margin junction with basis-ischium. Basis-ischium with long plumose setae; dorsal expansion incompletely fused and unarmed. Coxa with with row of long plumose setae medially; lat- one small spine on anterior margin. eral and mesial surfaces smooth. Carpus not Pereopod III (®g. 76C) dactylus with base produced dorsodistally; lateral and mesial to heel straight, heel broadly rounded and surfaces smooth; dorsal margin with short low, heel to tip with broad, evenly rounded simple and long plumose setae; ventral mar- indent, tip acute, tip to base smoothly convex gin with short simple setae. Merus with scat- to straight; lateral surface smooth, with sev- tered short transverse rows of setae on lateral eral small tufts of short setae in generally surface, dorsal and ventrodistal margins with straight line across medioproximal surface, long plumose setae; mesial surface with large dorsodistal margin with tufts of short setae; decalci®ed window proximoventrally. Basis- ventromesial margin with long plumose se- ischium incompletely fused and unarmed. tae, dorsal margin with short simple and plu- Coxa unarmed. mose setae; mesial surface smooth, with plu- Abdomen (®g. 76E) with somite I approx- mose setae proximally at junction with pro- imately as long as wide, widest posteriorly; podus. Propodus not in¯ated dorsoventrally; dorsal surface with anterior margin straight; lateral surface smooth, with long plumose se- posterior margin straight, with elevated sub- tae distally, simple setae on dorsal margin marginal row of short setae; small transverse and long plumose setae on ventral margin; decalci®ed windows laterad of segment me- dorsolateral surface narrow, oblique, ¯at- dian. Somite II dorsal surface with submar- tened; mesial surface with scattered long se- ginal transverse ridge anteriorly; with small tae on and near distal margin, with decalci- transverse decalci®ed windows laterad of ®ed window near junction with carpus. Car- segment median just anterior to submarginal pus produced dorsodistally, exceeding prox- ridge; with tuft of setae at posterolateral an- imal margin of propodus by approximately gle, extending onto pleuron posteromesially; one-fourth length of propodus, pointed but posterior margin with indistinct, punctate, not acute; dorsolateral margin unarmed; lat- submarginal groove laterally; pleura expand- eral surface slightly rugose dorsodistally, ed and directed slightly anteriorly; lateral with mat of short setae and two longer rows margins rounded, anterior and lateral mar- of setae ventrally; mesial surface smooth, gins with long plumose setae, posterior mar- with long plumose setae on margins and scat- gin with short setae. Somite III similar to so- tered on surface. Merus smooth; dorsal and mite II, but narrower, shorter, and lacking an- ventral margins unarmed, with long plumose terior submarginal ridge; small tuft of short setae; laterodistal margin with long plumose thick setae on posterolateral angle; pleura setae; mesial surface smooth, with decalci- thinner and shorter than on somite II, direct- ®ed window at junction with basis-ischium. ed anterolaterally, with setae as in somite II; Basis-ischium incompletely fused and un- anterolateral angle acute; dorsal surface armed. Coxa unarmed. Female with large obliquely ¯attened anterolaterally. Somite IV gonopore on anterior mesial surface of coxa, similar to somite III, but thinner and shorter; surrounded with short plumose setae; male dorsal surface with thick setae posterolater- with small pore. ally; pleura thinner and shorter than on so- Pereopod IV (®g. 76D) dactylus with base mite III, directed posterolaterally; dorsal sur- to tip convex to concave, tip acute, tip to face obliquely ¯attened anterolaterally; mar- base straight distally, becoming convex prox- gin with long plumose setae. Somite V wider 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 237 than somite IV; lateral margins with plumose quently, Boyko (2000a) reported the discov- setae; pleura absent. Somite VI subequal to ery of additional specimens of A. speciosa somite V in width but longer; dorsal surface from the Marquises Islands with identical with short transverse rows of setae laterad of banding patterns as seen in specimens of A. midline anteriorly; lateral margins with long madagascariensis. In addition, it is herein re- plumose setae; pleura absent. ported that A. Milne Edwards' (1862) spec- Females with uniramous, paired pleopods imen of ``Albunea symnista'' [sic] from ReÂ- on somites II±V; males without pleopods. union is actually the oldest record of A. spe- Telson of male (®g. 76F) spatulate, length ciosa from outside Hawaii (this record was subequal to width, produced into short repeated by Hoffman, 1874). Unfortunately, rounded tip distally; weakly calci®ed except Thomassin's (1973) type specimens have for large triangular anterior plate; median never been deposited in any museum, and all longitudinal groove long, extending to distal attempts to obtain them have proved unsuc- end of calci®ed plate, lined with long thin, cessful. In spite of this, it is clear from the simple setae; calci®ed plate slightly elevated morphological similarities between the two medially but without ridge. Telson of female taxa (discussed in detail by Boyko, 1999), (®g. 76G) ovate, longer than wide, broadly the discovery of similar banding patterns on triangular, dorsal surface smooth, with me- individuals from widely separated localities, dian longitudinal groove anteriorly; with row and the proximity of ReÂunion to Madagascar, of setose punctae lateral to midline from me- that Thomassin's (1973) Albunea madagas- dian of longitudinal groove to distal end of cariensis is a junior synonym of A. speciosa. groove; margins with long plumose setae. Little is known about the biology of this DISTRIBUTION: From Madagascar eastward species other than the few records of ovig- to the Marquises Islands and Hawaii, in 3.0± erous females as herein reported from April, 34 m depth. September, and October. The Japanese male MAXIMUM SIZE: Males: 10.4 mm cl; fe- has a large sperm ribbon between the gono- males, 14.0 mm cl. pores of the ®fth pereopods. Several A. spe- TYPE SPECIMENS: USNM 260868 (neotype ciosa specimens (QM W22285) were col- of A. speciosa, selected by Boyko, 1999); ho- lected together with the holotype of Albunea lotype and 18 paratypes of A. madagascar- danai, but it is not known if the two species iensis in the private collection of B. Tho- are typically sympatric in Hawaii. The col- massin and not deposited in MNHN as stated oration of this species is off-white with whit- by Thomassin (1973). ish setae, both in life (from color transpar- TYPE LOCALITIES: ``Kirk,'' Oahu, Hawaii, encies) and in preservative. USA (A. speciosa); Grand ReÂcif de Tulear, The fusion of CG1 is a character that Madagascar (A. madagascariensis). unites this species with two fossil taxa: A. REMARKS: This species was redescribed cuisiana Beschin and De Angeli and A. hah- and all known aspects of its biology dis- nae Blow and Manning. This character is cussed by Boyko (1999). Discovery of spec- otherwise unknown in the genus Albunea, imens of A. speciosa in the southern Paci®c but fused CG1 elements are found in other islands supports a southwest to northeast dis- albuneid genera of both subfamilies (e.g., persal pattern for this species from its pre- Lepidopa, Stemonopa). It appears that A. sumed western Indo-Paci®c origin towards speciosa, A. cuisiana, and A. hahnae form a Hawaii, the type locality and easternmost clade that is sister to all the other species of part of its range (see Boyko, 1999). Albunea. Because two of the three taxa in Because specimens of Albunea speciosa this clade are fossil, and therefore known were identi®ed from several localities outside from only limited characters, and because A. of Hawaii, Boyko (1999) suggested that the speciosa shares so many other characters similar A. madagascariensis might be a ju- with all other Albunea species, it is inadvis- nior synonym. This synonymy was actually able to erect a genus at this time based on ®rst suggested, but also not formally pro- these three species alone. Albunea speciosa posed, by SereÁne (1977) in a publication that can easily be separated from all of the other escaped the notice of later authors. Subse- Recent species of Albunea by the concave 238 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

as small acute tooth. Ocular sinus smoothly concave and unarmed. Frontal region smooth; setal ®eld narrow anteriorly and widening posteriorly; posterior lateral ele- ments reduced to narrow bands of setae. CG1 parallel to anterior margin of carapace, sinuous, strongly crenulate, medial fragment and curved posterior lateral elements united. Mesogastric region smooth; CG2 absent; CG3 broken into four short elements be- tween posterior lateral elements of CG1; CG4 with two short medial elements be- tween longer supralateral elements of CG4. Hepatic region smooth, with oblique setose groove at median of lateral margin. Epi- branchial region generally triangular, smooth; posterolateral margin with three short rows of setae. Metagastric region smooth; CG5 present as two triangular ele- Fig. 77. Albunea hahnae Blow and Manning, ments anteriorly displaced and overlapping 1996: A, 16.6 mm cl, USNM 484530, holotype. CG4. CG6 strongly crenulate, strongly an- A. Carapace, dorsal view. Scale ϭ 5.3 mm. teriorly concave medially and sloping out to anteriorly convex lateral thirds. CG7 oblique, united with lateral margins of me- shape of the lateral margins of the distal pe- dian segment of CG6. Cardiac region duncular segments. smooth; CG8 present as one long element. CG9 present as two short lateral grooves Albunea hahnae Blow and Manning, 1996 with gap at midline. CG10 present as two Figure 77 long lateral elements, with gap between Albunea hahnae Blow and Manning, 1996: 4, pl. fragments. CG11 present as short medial el- 1, ®g. 2*. ement. Post-CG11 element present. DISTRIBUTION: Known only the type local- MATERIAL EXAMINED: USA: South Caro- ity. lina: USGS 26882, Santee Limestone, M. M. TYPE SPECIMEN: USNM 484530 (holo- Berkeley Quarry, Berkeley Co., coll. un- type). known: 1 carapace, 16.6 mm cl, holotype TYPE LOCALITY: USGS 26882, Santee (USNM 484530). Limestone, M. M. Berkeley Quarry, Berke- DIAGNOSIS: Carapace longer than wide, ley Co., South Carolina, USA. covered with crenulate grooves. Anterior REMARKS: Blow and Manning (1996) cor- margin with six or seven spines on either rectly noted that this taxon is most similar to side of ocular sinus. Setal ®eld with thick A. cuisiana. Of the extant taxa, it is most lateral elements and concave anterior margin. similar to A. speciosa, but differs in a num- CG1 with fused posterior lateral elements; ber of characters including length of disto- CG4 with two short medial elements be- lateral submarginal groove and fusion of tween longer supralateral elements of CG4; CG6 and CG7. CG5 present as two triangular elements; CG6 and CG7 fused; CG8 complete; CG11 pre- Albunea cuisiana Beschin and De Angeli, sent. Rostrum present. 1984 DESCRIPTION: Carapace (®g. 77A) slightly longer than wide. Anterior margin slightly Figure 78 concave on either side of ocular sinus, be- Albunea cuisiana Beschin and De Angeli, 1984: coming convex laterally, with several spines 97±99, pl. 1, ®gs. 1, 1a, pl. 2, ®gs. 1, 1b*. ± De (six or seven visible) along length. Rostrum Angeli, 1998: 17±18, ®g. 1(2). 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 239

Fig. 78. Albunea cuisiana Beschin and De Angeli, 1984: A, 21.1 mm cl, MCSNV 10439, holotype; B, MCSNV, paratype. A. Carapace, dorsal view. B. Right pereopod I dactylus, propodus, and carpus, lateral view. Scale ϭ 8.6 mm.

MATERIAL EXAMINED: Italy: Middle Eo- long elements between posterior lateral ele- cene, Valle del Chiampo, Eastern Lessini, ments of CG1; CG4 with two or three short, coll. unknown: 1 carapace, 21.1 mm cl, ho- anteriorly displaced, medial elements be- lotype (calco-mold of MCSNV 10439). tween longer supralateral elements. Hepatic DIAGNOSIS: Carapace as long as wide, cov- region smooth, with oblique setose groove at ered with crenulate grooves. Anterior margin median of lateral margin. Epibranchial re- with few spines on either side of ocular si- gion generally triangular, smooth. Metagas- nus. Setal ®eld with thin lateral elements and tric region smooth; CG5 present as two me- produced anterior margin. CG1 with fused dial elements. CG6 strongly crenulate, posterior lateral elements; CG4 with three or strongly anteriorly concave medially and four short, anteriorly displaced, medial ele- sloping out to anteriorly convex lateral ments between longer supralateral elements; thirds. CG7 oblique, united with lateral mar- CG5 present as two medial elements; CG6 gins of median segment of CG6. Cardiac re- and CG7 fused; CG8 broken; CG11 present. gion smooth; CG8 present as one long and Rostrum present. Pereopod I dactylus dorsal two short elements; short elements displaced margin smooth; carpus dorsal margin slightly anteriorly. CG9 present as ®ve short smooth. elements alternating between anterior and DESCRIPTION: Carapace (®g. 78A) approx- posterior displacement. CG10 present as two imately as wide as long. Anterior margin long lateral elements, with two short ele- slightly concave on either side of ocular si- ments between. CG11 present as two short nus, becoming convex laterally with large elements. Post-CG11 element(s) present. spines along length. Rostrum as small acute Branchial region with numerous short, trans- tooth. Ocular sinus smoothly concave and verse rows of setae. Posterior margin deeply unarmed. Frontal region smooth; setal ®eld and evenly convex. narrow anteriorly and widening posteriorly; Pereopod I (®g. 78B) subchelate. Dactylus posterior lateral elements reduced to narrow curved and tapering; lateral and mesial sur- bands of punctations. CG1 parallel to ante- faces smooth. Propodal lateral surface with rior margin of carapace, sinuous, strongly numerous short, transverse rows of setose ru- crenulate, divided into medial fragment and gae; dorsal margin unarmed; ventral margin curved, posteriorly displaced, lateral ele- distally produced into acute spine (broken); ments. Mesogastric region smooth; CG2 pre- cutting edge lacking teeth. Carpus with dor- sent as one or two short medial elements; sodistal angle apparently not produced into CG3 broken into one to three short and two strong corneous-tipped spine; lateral surface 240 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 with transverse ridges on distal two-thirds of Jan. 15, 1964, coll. A. G. Smith and J. surface. DeRoy: 1 megalopa, 5.2 mm cl (CASIZ DISTRIBUTION: Known from the type local- 109337); South Seymour Island, 10 fms (ϭ ity as well as Cava ``Lovara'' and Cava 18.3 m), March 18, 1937, coll. W. A. Sea- ``Boschetto'' di Chiampo in Italy (De Angeli, holm and F. E. Lewis on R/V ``Stranger'': 1 1998: 18). (, 14.3 mm cl, holotype (USNM 267793); TYPE SPECIMENS: MCSNV 10439 (holo- South Seymour Island, 00Њ26ЈS, 90Њ19ЈW, 1 0 type), De Angeli Collection (paratype), pri- fms (ϭ 18.3 m), March 18, 1937, coll. W. I. vate collection (paratype). Seaholm on R/V ``Stranger'': 2 (, 7.3±8.3 TYPE LOCALITY: Middle Eocene, Valle del mm cl, 1 &, 15.0 mm cl, allotype and para- Chiampo, Eastern Lessini, Italy. types (USNM 267796). REMARKS: Because accurate generic place- Mexico: Baja California Sur: Isla EspõÂ- ment of a species depends on a number of ritu Santo, Baja California Sur, 6±20 fms (ϭ characters obtained from many morphologi- 11.0±36.6 m), March 31, 1939, coll. F. E. cal structures, it is always dif®cult to place Lewis on R/V ``Stranger'': 1 (, 18.4 mm cl, fossil species, especially when they are 1 &, 15.0 mm cl, paratypes (USNM 267795); known from fragmentary material. In the Sta. 37, La Vantana Bay, 24Њ08Ј15ЉN, case of Albunea cuisiana, it is fortunate that 109Њ52ЈW, 13±15 fms (ϭ 23.8±27.4 m), the carapace is well preserved and that ad- April 20, 1939, coll. F. E. Lewis on R/V ditional parts of the animal (part of pereopod ``Stranger'': 1 (, 15.5 mm cl, 3 &, 16.8±19.3 I, part of abdomen) are known. I was able to mm cl, paratypes (USNM 267797); Guer- directly examine a ``calco-mold'' of the ho- rero: Sta. 3, Zihuatanejo, 18±26 fms (ϭ lotype specimen carapace, but could not ex- 32.9±47.6 m), Jan. 30, 1939, coll. F. E. Lew- amine the pereopod I, abdomen, or other is on R/V ``Stranger'': 2 &, unmeasurable parts illustrated by Beschin and De Angeli (USNM 267794). (1984). Based on all available evidence this DIAGNOSIS: Carapace wider than long, cov- species appears to be closest to the fossil tax- ered with strongly setose grooves. Anterior on A. hahnae Blow and Manning. It shares margin with 8±14 spines on either side of with that species a similar arrangement of ocular sinus. Setal ®eld with thick lateral el- CG6 and CG7 (fused) and numerous short ements and concave anterior margin. CG1 grooves on the posterior hepatic region, many more than are typical of other species with separate posterior lateral elements; CG4 of Albunea. Both A. cuisiana and A. hahnae medial elements semicontiguous with supra- possess fused anterior and posterior elements lateral elements; CG5 present as two convex, of CG1, and a contiguous setal ®eld paral- nearly united, elements; CG6 and CG7 sep- leling this groove. This is marked contrast to arate; CG8 broken; CG11 present. Rostrum all other species of Albunea (except A. spe- present, not reaching posterior margin of oc- ciosa) where CG1 is separated into distinct ular plate. Ocular plate subquadrate. Distal anterior and posterior elements. peduncular segments dorsoventrally ¯attened and subtriangular in shape, tapering at tip, Albunea galapagensis, new species separated along mesial margins, lateral mar- Figures 79, 80 gins convex except slightly concave at tip, mesial margins convex. Cornea at tip. Dac- Albunea lucasia: Ramos and Rios, 1995: 103, ®g. tylus of pereopod II with heel produced and 5 (not Albunea lucasia de Saussure, 1853). rounded. Dactylus of pereopod III with heel MATERIAL EXAMINED: Ecuador: GalaÂpa- slightly produced and subquadrate. Dactylus gos Islands: Hard coral mud, off north Bar- of pereopod IV evenly sinuous from base to ranca, Academy Bay, Isla Santa Cruz, 5±10 tip, with shallow indent. Telson of male sub- fms (ϭ 9.1±18.3 m), Feb. 21, 1964, coll. A. triangular, elongated and tapering, length DeRoy, J. DeRoy and A. G. Smith: 1 &, 7.2 greater than width, distal tip narrow and sub- mm cl, paratype (CASIZ 109248); off the acute; thickly calci®ed and raised into ridge Barranca (Angemeyer's Houses), Academy dorsomedially, median of ridge lined with Bay, Isla Santa Cruz, 9.1±18.3 m (5±10 fms), long thin setae; lateral margins decalci®ed. 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 241

Telson of female similar to male, but with branous, with numerous irregular fragments less tapered tip. and sparsely covered with long plumose se- DESCRIPTION: Carapace (®g. 79A) wider tae. than long. Anterior margin concave on either Ocular plate (®g. 79B) triangular with side of ocular sinus, becoming convex lat- broad median indentation; median peduncu- erally, with 8±14 large spines (n ϭ 6) on lar segments present as small ovate calci®ed each side along length. Rostrum as small areas lateral to ocular plate. Distal peduncu- acute tooth, not reaching proximal margin of lar segments elongate, teardrop-shaped, with ocular plate. Ocular sinus smoothly concave. broadly convex lateral and weakly convex Frontal region smooth; setal ®eld narrow an- mesial margins, cornea covering lateral por- teriorly, broad posteriorly; posterior lateral tion of distal tip; lateral margins without elements reduced to narrow bands of setae. notch; mesial margins separated at base; me- CG1 parallel to anterior margin of carapace, sial and distolateral margins with long plu- convex, sinuous, strongly crenulate, divided mose setae; tuft of plumose setae at proxi- into medial fragment and curved, posteriorly molateral ventral angles and ventrolateral displaced lateral elements. Mesogastric re- oblique row of plumose setae extending from gion smooth; CG2 present as one or two tuft to three-fourths length of peduncle. short medial elements; CG3 present as one Antennule (®g. 79C) with segment III nar- semicontiguous, long, medial element be- row proximally, expanding distally to three tween posterior lateral elements of CG1 times proximal width; plumose setae on dor- (rarely broken into four shorter elements); sal and ventral margins and scattered on lat- CG4 medial elements semicontiguous with eral surface; dorsal exopodal ¯agellum with supralateral elements (rarely broken into four 74±94 articles (n ϭ 6), long plumose setae shorter elements). Hepatic region smooth, on dorsal and ventral margins; ventral en- with oblique setose groove at median of lat- dopodal ¯agellum with two or three articles eral margin. Epibranchial region generally (n ϭ 6), plumose setae on dorsal and ventral triangular, smooth; posterolateral margin margins. Segment II medially in¯ated in dor- with three oblique rows of short setae. Me- sal view, with long plumose setae on dorsal tagastric region smooth; CG5 present as two and ventral margins and scattered on lateral convex, nearly united, elements (rarely bro- surface. Segment I as wide as long, unarmed; ken into four shorter elements). CG6 strongly dorsal third of lateral surface rugose with crenulate, strongly anteriorly concave medi- long plumose setae; long plumose setae on ally and sloping out to anteriorly convex lat- dorsal and ventral margins. eral thirds. CG7 almost transverse, not reach- Antenna (®g. 79D) with segment V ap- ing lateral margins of median segment of proximately 2.5 times longer than wide, with CG6. Cardiac region smooth; CG8 present as long plumose setae on dorsal margin and two medial elements and two longer lateral scattered on distal half of lateral surface; ¯a- elements. CG9 present as two short lateral gellum with seven articles (n ϭ 6), long plu- grooves with narrow gap at midline. CG10 mose setae on dorsal, ventral, and distal mar- present as two long lateral elements. CG11 gins. Segment IV expanded distally, with present as one or two short elements. Post- long plumose setae on dorsal, ventral, and CG11 element present as one or two short distal margins, and two rows of short setae lateral elements. Branchial region with nu- on dorsolateral and ventrolateral surface. merous short and long transverse rows of se- Segment III with long plumose setae on ven- tae in anterior three-fourths. Posterior margin tral margin; short simple setae on dorsal mar- deeply and evenly convex, with submarginal gin and scattered on surface. Segment II groove reaching halfway up margin of pos- short, widening distally, rugose, with long terior concavity. Branchiostegite with long plumose setae on dorsal and distoventral anterior, submarginal spine; anterior region margins and short simple setae scattered on with scattered short, transverse lines ventral lateral surface; antennal acicle long, thin, and to linea anomurica; with many short rows of reaching distal margin of segment IV, with setae and sparsely covered with long plu- long plumose setae on dorsal margin. Seg- mose setae ventrally; posterior region mem- ment I rounded proximally, ¯attened ventro- 242 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Fig. 79. Albunea galapagensis, n. sp.: A±J, &, 19.3 mm cl, USNM 267797, paratype. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 1.7 mm (B), 2.1 mm (F), 3.3 mm (E, I), 4.4 mm (C, D, H, J), 5.9 mm (A), and 6.7 mm (G). 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 243 laterally, with long plumose setae on disto- with simple and plumose setae on margins. ventral margin, short plumose setae on dis- Basis-ischium incompletely fused, with plu- todorsal margin, and short simple setae in mose setae on margins. Exopod one-half lon- two short rows on surface rugae ventral to ger than merus, ¯agellum with one elongate spine; lateral surface with distal, subdorsal, article, shorter than carpus. acute spine; low semicircular dorsolateral Maxilliped III (®g. 79J) dactylus oblong, lobe ventrodistal to spine, margin of lobe with rounded tip; long plumose setae on mar- with long plumose setae; segment with ven- gins and in transverse row on distolateral sur- tromesial antennal gland pore. face. Propodus dorsodistally in¯ated, with Mandible (®g. 79E) incisor process with longitudinal median row of plumose setae on two teeth; cutting edge with one tooth. Palp lateral surface; margins with plumose setae. three-segmented, with plumose setae on mar- Carpus produced onto propodus approxi- gins and long, thick, simple setae arising mately one-fourth length of propodus; lateral from bend in second segment and on distal surface with two transverse rows of long plu- margin of terminal segment. mose setae; long plumose setae on dorsal Maxillule (®g. 79F) distal endite proxi- margin. Merus in¯ated, unarmed, with plu- mally narrow, widening to in¯ated distal end, mose setae on dorsal margin and few scat- with thick simple setae on distal margin and tered small areas on proximoventral half of thin simple setae on dorsal margin. Proximal lateral surface. Basis-ischium incompletely endite with thick simple setae on distal mar- fused, with weak crista dentata of two or gin. Endopodal external lobe truncate distally three teeth. Exopod two-segmented: proxi- and curled under; internal lobe reduced with mal segment small; distal segment styliform, three thick setae at distolateral margin. tapering, approximately one-fourth length of Maxilla (®g. 79G) exopod evenly rounded, merus; with plumose setae on margins; with- with plumose setae along distal margin. Sca- out ¯agellum. phognathite bluntly angled on posterior lobe, Pereopod I (®g. 80A) dactylus curved and with plumose setae. tapering; lateral and mesial surfaces smooth; Maxilliped I (®g. 79H) epipod with plu- dorsal margin with long plumose setae; ven- mose setae on margins, distolateral surface tral margin with short simple setae. Propodal and mesial surface. Endite tapered distally lateral surface with numerous short, trans- and subequal to ®rst segment of exopod. Ex- verse rows of setose rugae; dorsal margin un- opod with two segments; proximal segment armed; ventral margin distally produced into narrow, margins parallel with plumose setae; acute spine; cutting edge lacking teeth, lined distal segment spatulate, longer than wide, with long plumose setae; dorsal margin with broadest medially, margins and mesiodorsal long plumose setae, ventral margin with surface with long plumose setae. Endopod short simple setae; mesial surface with few ¯attened and elongate, reaching two-thirds to scattered rows of short plumose setae. Car- distal end of proximal exopodal segment; pus with dorsodistal angle produced into plumose setae on margins and median of lat- strong corneous-tipped spine; dorsal margin eral surface. with short transverse grooves behind spine; Maxilliped II (®g. 79I) dactylus evenly dorsal, distal, and distoventral margins with rounded, length subequal to width, with thick long plumose setae; lateral surface with simple setae distally and in transverse row small distal rugose area, few transverse se- on distolateral surface. Propodus two times tose ridges on distal half of surface; mesial wider than long, slightly produced at dorso- surface smooth, with medial and subdorsal distal angle, with plumose setae on dorsal interrupted transverse rows of setae, margins margin and patch of long simple setae on with long plumose setae. Merus unarmed; dorsodistal and ventrodistal angles. Carpus lateral surface with scattered transverse rows not produced dorsodistally, approximately of long plumose setae, distal margin with two times longer than wide; long simple se- long plumose setae; mesial surface with few tae on dorsal margin and in few patches on scattered short setae; fully calci®ed. Basis- lateral surface. Merus approximately three ischium incompletely fused, unarmed. Coxa times longer than wide, margins parallel; unarmed. 244 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Fig. 80. Albunea galapagensis, n. sp.: A±E, H, &, 19.3 mm cl, USNM 267797, paratype; F, G, (, 15.5 mm cl, USNM 267797, paratype. A. Left pereopod I, lateral view. B. Left pereopod II, lateral view. C. Left pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of (, lateral view. H. Telson of &, dorsal view. Scale ϭ 3.3 mm (F±H) and 4.4 mm (A±E). 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 245

Pereopod II (®g. 80B) dactylus smooth; smooth, with long plumose setae in oblique base to heel slightly convex, heel low and medial row, simple setae on dorsal margin, rounded, heel to tip with narrow, acute in- plumose setae on ventral margin; dorsolateral dent, tip subacute, tip to base broadly con- surface narrow, oblique, ¯attened; mesial vex; lateral surface smooth, with several surface smooth. Carpus produced dorsodis- small tufts of short setae in generally straight tally, exceeding proximal margin of propo- line across medioproximal surface, several dus by one-®fth length of propodus; tip widely spaced submarginal tufts of short se- rounded, dorsolateral margin unarmed; lat- tae dorsodistally; mesial surface smooth, eral surface slightly rugose in dorsodistal ventral margin with long plumose setae, dor- half, with mat of short setae and two inter- sal margin with short simple setae, patch of rupted rows of long plumose setae ventrally long plumose setae at base. Propodal dorsal in proximal half; mesial surface smooth, with surface smooth, with ventral margin in¯ated long plumose setae on distal margin. Merus and rounded; oblique rows of long plumose smooth, with large decalci®ed window cov- setae on distal margin of lateral surface and ering nearly all of lateral surface medially; in midline; distal and ventral margins with dorsal and ventral margins unarmed, dorso- long plumose setae; dorsolateral surface as distal and ventrodistal margins with long plu- narrow, oblique, ¯attened shelf, with short mose setae, few long plumose setae on lateral setae on dorsal margin and long plumose se- surface in proximal third; mesial surface tae on ventral margin; mesial surface with smooth. Basis-ischium incompletely fused elevated, curved setose ridge from ventral and unarmed. Coxa unarmed. Female with junction with dactylus almost to ventral large gonopore on anterior mesial margin of proximal junction with carpus. Carpus coxa, directly opposing other pore and sur- strongly produced and subacute dorsodistal- rounded with short plumose setae; male with ly, dorsal margin smooth; lateral surface minute pore. smooth, dorsodistal region smooth, irregular, Pereopod IV (®g. 80D) dactylus with base interrupted row of rugae and submarginal el- to tip slightly convex proximally, with indis- evated ridge ventrally, rugae and ridge with tinct heel and faintly concave indent, almost long plumose setae; dorsal and distoventral straight from indent to tip, tip subacute, tip margins with long plumose setae; mesial sur- to base convex; lateral surface smooth, ven- face smooth, with row of long plumose setae tral margin with long plumose setae, dorsal distally and subventrally and interrupted row margin with short simple setae; mesial sur- of long plumose setae subdorsally. Merus face with dorsal decalci®ed region, demar- with large median decalci®ed window cov- cated ventrally by longitudinal elevated ering nearly all of lateral surface, long plu- ridge, with row of short setae; setose punc- mose setae on dorsodistal and ventral mar- tations ventral to decalci®ed window. Pro- gins; mesial surface nearly smooth, with two podus expanded dorsally and ventrally; ven- long rows of long plumose setae. Basis-is- tral expansion exceeding ventral margin of chium incompletely fused and unarmed. dactylus, ventral margin with long plumose Coxa unarmed. setae; dorsal expansion with row of long plu- Pereopod III (®g. 80C) dactylus with base mose setae dorsally, oblique area with mat of to heel slightly convex, heel low and round- short simple setae; lateral surface smooth, ed, heel to tip with broadly concave indent, mesial surface smooth, with few patches of tip subacute, tip to base smoothly convex; long plumose setae on distoventral area. Car- lateral surface smooth, with several small pus slightly produced dorsodistally; ventral tufts of short setae in generally straight line four-®fths of lateral surface and mesial sur- across medioproximal surface, dorsodistal face smooth, dorsodistal one-®fth of lateral margin with tufts of short setae; ventral mar- surface with mat of short setae; dorsal mar- gin with long plumose setae, dorsal margin gin with short simple and long plumose se- with short simple and plumose setae; mesial tae; ventral margin with few short simple se- surface smooth, with plumose setae proxi- tae; mesial surface decalci®ed medially. Mer- mally at junction with propodus. Propodus us with scattered short transverse and oblique not in¯ated dorsoventrally; lateral surface rows of setae on lateral surface, dorsal and 246 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 ventrodistal margins with long plumose se- one-half length, row of long simple setae of tae; proximoventral third of mesial surface either side of median groove beginning at with large decalci®ed window. Basis-ischium median and continuing to distal margin of incompletely fused and unarmed. Coxa un- calci®ed area, converging distally; proximo- armed. lateral angles with short simple setae; mar- Abdomen (®g. 80E) somite I wider than gins with long simple setae. Telson of female long, widest posteriorly; dorsal surface with (®g. 80H) similar to male, but with less ta- anterior margin straight; posterior margin pered distal tip and less pronounced lateral curved, with elevated submarginal row of expansion, dorsal surface evenly calci®ed; short setae; few scattered short, simple setae median groove similar to male, but with distolateral to submarginal row; small trans- shorter setae; proximolateral angle with few verse, decalci®ed windows laterad of seg- short simple setae, margins with long simple ment median. Somite II dorsal surface with setae. irregular submarginal, transverse ridge ante- DISTRIBUTION: Known from the GalaÂpagos riorly; with small transverse, decalci®ed win- Islands, Baja California Sur (Gulf side), and dows laterad of segment median just anterior Guerrero, Mexico, in 9.1±47.6 m depth. Also to submarginal ridge; pleura expanded and from Colombia (Ramos and Rios, 1995). directed anterolaterally; anterolateral margins MAXIMUM SIZE: Males: 18.4 mm cl; fe- angled, anterior and lateral margins with long males: 19.3 mm cl. plumose setae, posterolateral angle rounded, TYPE SPECIMENS: USNM 267793 (holo- posterior margin with short setae; postero- type), CASIZ 109248 (paratype), USNM mesial angle with mat of short simple setae 267796 (allotype, 2 paratypes), USNM extending onto somite. Somite III similar to 267795 (paratype), USNM 267797 (3 para- somite II, but narrower, shorter, anterior sub- types). marginal windows present; pleura thinner TYPE LOCALITY: South Seymour Island, and shorter than on somite II, directed pos- GalaÂpagos Islands, Ecuador, 18.3 m. terolaterally proximally and anterolaterally ETYMOLOGY: This taxon is named after the distally, with setae as in somite II; anterolat- type locality islands which served as Dar- eral angle subacute; dorsal surface obliquely win's inspiration and have yielded so many ¯attened anterolaterally, with submarginal important discoveries in the biological sci- row of short setae. Somite IV similar to so- ences. mite III, but thinner and shorter, anterior sub- REMARKS: The discovery of this species in marginal windows present; pleura thinner both the GalaÂpagos Islands and the Gulf of and shorter than on somite III, directed lat- California demonstrates how much we have erally; dorsal surface obliquely ¯attened an- yet to learn about these highly diverse areas, terolaterally; lateral and posterior margins in spite of many decades of intensive col- with long plumose setae, anterior margin lecting. This species shares its distributional with short simple setae. Somite V wider than pattern with numerous other species of Ga- somite IV, anterior submarginal windows laÂpagos anomurans that are also known from present; lateral margins with plumose setae; the Panamic province, such as the porcelain pleura absent. Somite VI slightly broader crabs Petrolisthes haigae Chace and P. ton- than somite V, anterior submarginal windows sorius Haig, and possibly the sand crab, Lep- present; dorsal surface with two short trans- idopa wollebaeki, if that species and L. mex- verse rows of setae laterad of midline and on icana are conspeci®c (see under L. wollebae- posterior margin; pleura absent. ki and L. mexicana). Females with uniramous, paired pleopods Albunea galapagensis belongs to the on somites II±V; males without pleopods. group of Albunea containing A. microps and Telson of male (®g. 80F, G) subtriangular, A. elioti. elongated, and tapering, length greater than width, distal tip narrow and subacute; thickly Albunea microps Miers, 1878 calci®ed medially, in¯ated dorsomedially; Figures 81, 82 distal two-thirds with lateral decalci®ed re- Albunea microps White, 1847: 129* (nomen nu- gion; median longitudinal groove extending dum). ± Miers, 1878: 328±329, pl. 5, ®gs. 12, 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 247

13*. ± Henderson, 1888: 40*. ± Ortmann, 1896: on ``Pele'': 1 (, 9.1 mm cl, 1 &, 10.1 mm 224±225 (list). ± Ortmann, 1901: pl. 72, ®g. 4, cl (WAM 10409); south lagoon, Sibutu, pl. 79, ®g. 3. ± Borradaile, 1904: 751*. ± Gor- 04Њ31ЈN, 119Њ22ЈE, 9±13 fms (ϭ 16.5±23.8 don, 1938: 187, ®g. 3c*. ± SereÁne and Umali, m), Feb. 25±26, 1964, coll. B. R. Wilson on 1965: 95±97, pl. 4, ®gs. 1±6, text-®g. 12c. ± ``Pele'': 1 (, 8.6 mm cl (WAM 10416); west Thomassin, 1969: 140±143 (part), text-®gs. 2, 3b. ± Miyake, 1978: 154±155, ®g. 60b. ± CoeÃl- of Pearl Bank, 9±12 fms (ϭ 16.5±21.9 m), ho and Calado, 1987: 43, table 1. ± Calado, Feb. 21, 1964, coll. B. R. Wilson on ``Pele'': 1997a: 17, 21±22. ± Markham and Boyko, 1 (, 4.9 mm cl (WAM 10412), 1 (, 10.2 1999: 5, 7*. ± Boyko and Harvey, 1999: 383± mm cl (ZRC 1970.11.3.1); 2.5 mi and 358Њ 386, 400 (list), 402 (key), ®gs. 1, 4*. ± Boyko, from Malanipa Island, Basilan Straits, 15 fms 1999: 145 (list), ®g. 1*. ± Clark and Presswell, (ϭ 27.4 m), Feb. 12, 1964, coll. B. R. Wilson 2001: 154 (list). on ``Pele'': 2 (, 8.2±10.8 mm cl, 1 oviger, Albunea [sp.] Gordon, 1938: 189±190, ®g. 1d*. 10.0 mm cl (WAM 10417). Albunea symnista [sic]: SereÁne, 1977: 47. ± Ca- India: Andaman Islands: Port Blair, coll. lado, 1995: 71±73, pl. 4, ®g. i, pl. 5, ®g. h, pl. unknown: 2 (, 6.2±6.7 mm cl (BMNH 21, ®gs. a, b, pl. 22, ®gs. a±f* (not Albunea symmysta (Linnaeus, 1758)). 1956.1.14.20). not Albunea microps: Thomassin, 1969: 140±143 Indonesia: Corindon II, Sta. DR293, (part), pl. 2, ®gs. 1±9. ± Calado, 1995: 46±49, 02Њ37.7ЈS, 117Њ49.4ЈE, Makassar, 45.0 m, pl. 4, ®g. e, pl. 5, ®g. d, pl. 12, ®gs. a±f (ϭ Nov. 10, 1980, coll. R/V ``Coriolis'': 1 ovi- Albunea elioti Benedict, 1904). ger, 12.1 mm cl (MNHN-Hi 201). New Guinea: Sta. 549, east coast of Rouw MATERIAL EXAMINED: Zanzibar: Dredged Island, Aoeri Group, West Geelrinck Bay, in muddy sand, around Bawi and Change Is- Feb. 23, 1956, coll. A. F. Ostheimer on ``Glo- lands, off Zanzibar City, 6±16 fms (ϭ 11.0± ria Maris'': 1 (, 11.2 mm cl (RMNH 23641). 29.3 m), Feb. 27, 1957, coll. A. J. Ostheimer Australia: Queensland: Michaelmas Cay, III: 1 &, 10.9 mm cl (ANSP CA4646); Sta. Great Barrier Reef, north of Cairns, Sept. 14, 650, dredged in shell and sponge, approxi- 1963, coll. W. Goode: 1 (, 11.6 mm cl mately 1.5 mi east of Puopu Island, 7±9 fms (WAM 23385); Cape Sandwich, north of (ϭ 12.8±16.5 m), Feb. 20, 1957, coll. A. J. Townsville, 10 fms (ϭ 18.3 m), Nov. 1963, Ostheimer III: 1 &, 12.8 mm cl (ANSP coll. W. Goode: 1 unsexable specimen, 10.4 CA4647). mm cl (WAM 23388); Cairns Reef near Oman: Muscat, 10±15 fms (ϭ 18.3±27.4 Cooktown, Oct. 1, 1963, coll. W. Goode: 1 m), coll. unknown: 1 (, 7.8 mm cl (BMNH (, 11.4 mm cl (WAM 23400); Rudder Reef 1901.4.20.10). off Mossman, 16Њ11ЈS, 145Њ40ЈE, 30 m, Oct. Seychelles: MaheÂ, July±Aug. 1972, coll. 21, 1973, coll. R. J. Plant: 2 (, 6.6±11.0 mm Mission Zoologique MRAC-ULB: 2 &, 6.0± cl, 4 &, 6.2±13.0 mm cl (MOV J14553), 1 10.2 mm cl (MRAC 53.604). &, 15.7 mm cl (MOV J40128 ex MOV Maldives: Mahlosmadulu Atoll, 20 fms J14553); Rudder Reef, 30 mi northeast of (ϭ 36.6 m), pre±1900, coll. J. S. Gardiner: 2 Mossman, 16Њ11ЈS, 145Њ40ЈE, 3.1 m, Oct. &, 4.6±4.7 mm cl (UMZC). 21, 1973, coll. R. J. Plant: 1 (, 11.0 mm cl Philippines: Sulu Archipelago: ``Sooloo (MOV J44734); Rudder Reef, 30 mi north- Islands, Eastern Seas,'' coll. unknown: 1 (, east of Mossman, 16Њ11ЈS, 145Њ40ЈE, 3 m, 11.3 mm cl, holotype (BMNH 1937.6.7.3); Oct. 1973, coll. R. J. Plant: 2 (, 5.9±6.4 mm Sta. 212, 06Њ54ЈN, 122Њ18ЈE, 10 fms (ϭ 18.3 cl (MOV J44735); Michaelmas Reef, m), Jan. 30, 1875, coll. R/V ``Challenger'': 16Њ35ЈS, 146Њ00ЈE, Jan. 1964, coll. D. Beck- 1 (, 9.5 mm cl (ZMUC 2715 ex BMNH), 1 worth: 1 &, 12.9 mm cl (MOV J44727); unsexable, unmeasurable specimen (MNHN- north edge of reef around coral patches, 1.25 Hi 11 ex BMNH); south lagoon, Sibutu, mi from east end of cay, Michaelmas Reef, 04Њ31ЈN, 119Њ22ЈE, 12 fms (ϭ 21.9 m), Feb. 15 ft (ϭ 4.5 m), Oct. 14, 1965, coll. Austra- 25±26, 1964, coll. B. R. Wilson on ``Pele'': lian Museum Party: 2 (, 10.4±10.6 mm cl, 1 &, 12.0 mm cl (WAM 10408); south la- 1 &, 10.2 mm cl (AM P19646); Lizard Is- goon, Sibutu, 04Њ31ЈN, 119Њ22ЈE, 13 fms (ϭ land, near Cooktown, 4 m, May 26±June 4, 23.8 m), Feb. 25, 1964, coll. B. R. Wilson 1975, coll. G. Anderson: 1 &, 12.3 mm cl 248 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

(AM P20648); Little Trunk Reef, 18Њ20ЈS, dial elements; CG3 broken into one to ®ve 146Њ46ЈE, 9.1±12.2 m, Nov. 5, 1990, coll. K. short and one or two long elements between Lamprell: 1 ( 10.0 mm cl, 2 &, 10.2±11.7 posterior lateral elements of CG1; CG4 with mm cl (QM W17460). one short, anteriorly displaced, medial ele- New Caledonia: Sta. DW 1390, ment and two longer lateral elements spaced 18Њ27.5ЈS, 163Њ08.7ЈE, 38 m, May 11, 1999, approximately equally between longer supra- coll. ``Alis'' Campagne SURPRISES (B. lateral elements of CG4. Hepatic region Richer De Forges): 1 (, 9.7 mm cl (MNHN- smooth with oblique setose groove at median Hi 261). of lateral margin. Epibranchial region gen- DIAGNOSIS: Carapace longer than wide, erally triangular, smooth; posterolateral mar- covered with strongly setose grooves. Ante- gin with three short rows of setae. Metagas- rior margin with 8±11 spines on either side tric region smooth; CG5 entire. CG6 strongly of ocular sinus. Setal ®eld with thick lateral crenulate, strongly anteriorly concave medi- elements and straight anterior margin. CG1 ally and sloping out to anteriorly convex lat- with separate posterior lateral elements; CG4 eral thirds. CG7 oblique, almost reaching lat- fragmented, median element displaced ante- eral margins of median segment of CG6. riorly; CG5 entire; CG6 and CG7 united; Cardiac region smooth; CG8 present as three CG11 present. Rostrum present, exceeding long elements. CG9 present as two short lat- posterior margin of ocular plate by approxi- eral grooves with gap at midline. CG10 pre- mately one-half length of ocular plate. Ocu- sent as two long lateral fragments, with gap lar plate subquadrate. Distal peduncular seg- between fragments, short line anterior to ments dorsoventrally ¯attened and oblong in CG11 may be posteriorly displaced medial shape, rounded at tip, approximated along fragment of CG10. CG11 present as one or distal two-thirds of mesial margins, lateral two short elements. Branchial region with margins convex except at concave tip, mesial numerous short, transverse rows of setae. margins convex. Cornea on lateral margin at Posterior margin deeply and evenly convex, tip. Dactylus of pereopod II with heel pro- with submarginal groove reaching less than duced and rounded. Dactylus of pereopod III onee-third up margin of posterior concavity. with heel produced and rounded. Dactylus of Branchiostegite with short anterior submar- pereopod IV evenly sinuous from base to tip, ginal spine; anterior region with scattered with shallow indent. Telson of male broadly short, transverse lines ventral to linea ano- triangular, in¯ated dorsally, broadly rounded murica; with many short rows of setae and at tip, strongly calci®ed proximally, large de- sparsely covered with long plumose setae calci®ed area on either side of thin medial ventrally; posterior region membranous, with calci®ed strip, long thin setae medially and numerous irregular fragments and sparsely along anterior margin of windows. Telson of covered with long plumose setae. female ¯attened, ovate, longitudinal row of Ocular plate (®g. 81B, C) subquadrate, short, thin setae medially. with shallow median indentation; median pe- DESCRIPTION: Carapace (®g. 81A) slightly duncular segments present as small ovate, wider than long. Anterior margin slightly calci®ed areas lateral to ocular plate. Distal concave on either side of ocular sinus, be- peduncular segments irregularly oblong, with coming convex laterally, with 8±11 large convex lateral and mesial margins, cornea lo- spines (n ϭ 5) along length. Rostrum as cated laterally on produced tip; mesial mar- small acute tooth, extending halfway across gins approximated along distal two-thirds; the ocular plate. Ocular sinus smoothly con- mesial and lateral margins with long plumose cave and unarmed. Frontal region smooth; setae; tuft of plumose setae at proximolateral setal ®eld broad anteriorly and posteriorly; ventral angle and medial row of plumose se- posterior lateral elements lacking. CG1 par- tae extending from tuft to base of cornea. allel to anterior margin of carapace, faintly Antennule (®g. 81D) with segment III nar- sinuous, strongly crenulate, divided into me- row proximally, expanding distally to two dial fragment and curved, posteriorly dis- times proximal width; with plumose setae on placed, lateral elements. Mesogastric region dorsal and ventral margins and sparsely scat- smooth; CG2 present as one or two short me- tered on lateral surface; dorsal exopodal ¯a- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 249

Fig. 81. Albunea microps Miers, 1878: A, C, oviger, 12.1 mm cl, MNHN Hi-201; B, D±K, (, 11.0 mm cl, MOV J44734. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular pe- duncles, dorsal view. C. Ocular peduncles, dorsal view. D. Left antennule, lateral view. E. Left antenna, lateral view. F. Right mandible, mesial view. G. Right maxillule, lateral view. H. Right maxilla, lateral view. I. Right maxilliped I, lateral view. J. Left maxilliped II, lateral view. K. Right maxilliped III, lateral view. Scale ϭ 1.6 mm (C, G), 2.2 mm (B, F, J), 3.3 mm (D, E, H, I, K), and 4.4 mm (A). gellum with 55±75 articles (n ϭ 5), long plu- ¯ated from dorsal view, with plumose setae mose setae on dorsal and ventral margins; on dorsal and ventral margins and scattered ventral endopodal ¯agellum short, with two on ventrolateral third of surface. Segment I articles (n ϭ 5) and plumose setae on dorsal wider than long, unarmed; dorsal third of lat- and ventral margins. Segment II medially in- eral surface rugose, with long plumose setae; 250 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 long plumose setae on dorsal and ventral two-thirds of distance to distal end of prox- margins. imal exopodal segment; plumose setae on Antenna (®g. 81E) with segment V ap- margins and median of lateral surface. proximately 2.5 times longer than wide, with Maxilliped II (®g. 81J) dactylus evenly long plumose setae on dorsal margin and rounded, length equal to width, with thick scattered on lateral surface; ¯agellum with simple setae distally and on distolateral sur- ®ve or six articles (nϭ5), long plumose setae face. Propodus two times wider than long, on dorsal, ventral, and distal margins. Seg- slightly produced at dorsodistal angle, with ment IV expanded distally, with long plu- plumose setae on dorsal margin and patch of mose setae on dorsal, ventral and distal mar- long simple setae on lateral surface and ven- gins, and row of setae on dorsolateral sur- trolateral angle. Carpus not produced dorso- face. Segment III with long plumose setae on distally, approximately two times longer than dorsal and ventral margin and in short row wide; long simple setae on dorsal and distal on surface. Segment II short, widening dis- margins. Merus approximately three times tally, rugose, with plumose setae on margins longer than wide, margins parallel; with sim- and scattered on lateral surface; antennal aci- ple and plumose setae on margins and scat- cle long, thin, and just exceeding distal mar- tered on surface. Basis-ischium incompletely gin of segment IV, with long plumose setae fused with plumose setae on margins. Exo- on dorsal margin. Segment I rounded proxi- pod one-third longer than merus, ¯agellum mally, ¯attened ventrolaterally, with long with one elongate article. plumose setae on margins and scattered on Maxilliped III (®g. 81K) dactylus with surface rugae behind spine; lateral surface rounded tip; long plumose setae on margins with acute spine dorsally, low semicircular and lateral surface. Propodus slightly in¯ated dorsolateral lobe ventrodistal to spine; seg- medially, with longitudinal median row of ment with ventromesial antennal gland pore. plumose setae on lateral surface; margins Mandible (®g. 81F) incisor process with with plumose setae. Carpus produced onto two teeth; cutting edge with one tooth. Palp propodus almost one-half length of propo- three-segmented, with plumose setae on mar- dus; lateral surface with two rows of plumose gins and long, thick, simple setae arising setae on surface; plumose setae on margins. from bend in second segment and on distal Merus in¯ated, unarmed, with plumose setae margin of terminal segment. on margins and scattered on lateral surface. Maxillule (®g. 81G) distal endite proxi- Basis-ischium incompletely fused, with weak mally narrow, widening to in¯ated distal end, crista dentata of two or three teeth. Exopod with thick simple setae on distal margin and two-segmented: proximal segment small; thin simple setae on dorsal margin. Proximal distal segment styliform, tapering, approxi- endite with thick simple setae on distal mar- mately one-half length of merus; with plu- gin. Endopodal external lobe truncate distally mose setae on margins; without ¯agellum. and curled under; internal lobe reduced, with Pereopod I (®g. 82A) dactylus curved and two thick setae at distolateral margin. tapering; lateral and mesial surfaces smooth; Maxilla (®g. 81H) exopod evenly rounded, dorsal margin proximally rugose, with long with plumose setae along distal margin. Sca- plumose and short simple setae; ventral mar- phognathite bluntly angled on posterior lobe, gin with short simple setae. Propodal lateral with plumose setae. surface with numerous short, transverse rows Maxilliped I (®g. 81I) epipod with plu- of setose rugae; dorsal margin unarmed; ven- mose setae on margins, distolateral surface, tral margin distally produced into acute and mesial surface. Endite tapered distally spine; cutting edge lacking teeth, lined with and subequal to ®rst segment of exopod. Ex- long plumose setae; dorsal margin with long opod with two segments: proximal segment plumose setae, ventral margin with short narrow, margins parallel, margins with plu- simple setae. Carpus with dorsodistal angle mose setae; distal segment spatulate, longer produced into strong corneous-tipped spine; than wide, broadest medially, margins and dorsal margin with short transverse grooves mesioventral surface with long plumose se- behind spine; dorsal and distal margins with tae. Endopod ¯attened and elongate, reaching long plumose setae; lateral surface with 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 251

Fig. 82. Albunea microps Miers, 1878: A±D, (, 11.0 mm cl, WAM J44734; E, (, 10.4 mm, AM P19646; F, (, 11.3 mm cl, BMNH 1937.6.7.3, holotype; G, oviger, 12.1 mm cl, MNHN Hi-201. A. Left pereopod I, lateral view. B. Left pereopod II, lateral view. C. Left pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 2.2 mm (F), 3.3 mm (E, G), and 4.4 mm (A±D). small distal rugose area, with few transverse broadly convex; lateral surface smooth, with setose ridges on distal half of surface; mesial several small tufts of short setae in generally surface smooth, with medial transverse row straight line across medioproximal surface, of setae, margins with long plumose setae. several widely spaced submarginal tufts of Merus unarmed; lateral surface with scat- short setae dorsodistally; mesial surface tered transverse rows of long plumose setae, smooth, ventral margin with long plumose margins with long plumose setae; mesial side setae, dorsal margin with short simple setae, with few scattered setae; fully calci®ed. Ba- with patch of long plumose setae at base. sis-ischium incompletely fused, unarmed. Propodal dorsal surface smooth, with ventral Coxa unarmed. margin in¯ated and rounded; oblique row of Pereopod II (®g. 82B) dactylus smooth; long plumose setae on distal margin of lateral base to heel slightly concave, heel smoothly surface; distal and ventral margin with long rounded, slightly produced, heel to tip with plumose setae; dorsolateral surface as nar- wide, acute indent, tip acute, tip to base row, oblique, ¯attened shelf, with short setae 252 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 on dorsal margin and long plumose setae on ed with short plumose setae; male without ventral margin; mesial surface with elevated, pore. curved, setose ridge from ventral junction Pereopod IV (®g. 82D) dactylus with base with dactylus almost to ventral proximal to tip convex to concave, tip acute, tip to junction with carpus. Carpus slightly pro- base convex; lateral surface smooth, ventral duced and gently rounded dorsodistally, dor- margin with long plumose setae, dorsal mar- sal margin unarmed; lateral surface smooth, gin with short simple setae; mesial surface with setose mat on produced area and irreg- with dorsal decalci®ed region, demarcated ular, interrupted row of rugae and submar- ventrally by longitudinal elevated ridge with ginal elevated ridge ventrally, rugae and row of short setae; with setose punctations ridge with long plumose setae; margins with ventral to decalci®ed window. Propodus ex- long plumose setae; mesial surface smooth, panded dorsally and ventrally; ventral expan- with row of long plumose setae subdorsally. sion reaching ventral margin of dactylus, Merus with large median decalci®ed window margin with long plumose setae; dorsal ex- covering nearly all of lateral surface, with pansion with row of long plumose setae dor- few scattered setae on surface and margins; sally, oblique area with mat of short simple mesial surface nearly smooth, with two long setae; lateral and mesial surfaces smooth. rows of setae. Basis-ischium incompletely Carpus slightly produced dorsodistally; ven- fused and unarmed. Coxa unarmed. tral three-fourths of lateral surface and mesial Pereopod III (®g. 82C) dactylus with base surface smooth, dorsodistal quarter of lateral to heel straight, heel broadly rounded and surface with mat of short setae; dorsal mar- low, heel to tip with broadly concave indent, gin with short simple and long plumose se- tip acute, tip to base smoothly convex; lateral tae; ventral margin with short simple setae; surface smooth, with several small tufts of mesial surface partially decalci®ed medially. short setae in generally straight line across Merus lateral surface with scattered short medioproximal surface, dorsodistal margin transverse rows of setae, dorsal and ventrod- with tufts of short setae; ventral margin with istal margins with long plumose setae; prox- long plumose setae, dorsal margin with short imoventral third of mesial surface with large simple and plumose setae; mesial surface decalci®ed window. Basis-ischium incom- smooth with plumose setae proximally at pletely fused and unarmed. Coxa unarmed. junction with propodus. Propodus not in¯at- Abdomen (®g. 82E) with somite I wider ed dorsoventrally; lateral surface smooth, than long, widest posteriorly; dorsal surface with long plumose setae in oblique row, sim- with anterior margin straight; posterior mar- ple setae on dorsal margin; dorsolateral sur- gin curved, with elevated submarginal row face narrow, oblique, ¯attened, with setose of short setae; small transverse decalci®ed mat; mesial surface with scattered long setae windows laterad of segment median. Somite on and near distal margin, and in oblique row II dorsal surface with submarginal transverse on surface. Carpus produced dorsodistally, ridge anteriorly; with small transverse decal- only slightly exceeding proximal margin of ci®ed windows laterad of segment median propodus; dorsolateral margin unarmed; lat- just anterior to submarginal ridge; pleura ex- eral surface slightly rugose dorsodistally, panded and directed anterolaterally; lateral with mat of short setae and row of setae ven- margins rounded, anterior and lateral mar- trally; mesial surface smooth, with long plu- gins with long plumose setae, posterior mar- mose setae on margins. Merus smooth, with gin with short setae; posteromesial angle large decalci®ed window covering nearly with mat of short simple setae. Somite III half of lateral surface medially; dorsal and similar to somite II, but narrower, shorter; ventral margins unarmed, with long plumose pleura thinner and shorter than on somite II, setae; distolateral margin with long plumose directed posterolaterally, with setae as in so- setae; mesial surface smooth. Basis-ischium mite II; anterolateral angle acute; dorsal sur- incompletely fused and unarmed. Coxa of face obliquely ¯attened anterolaterally. So- male with small anteromesial spine; coxa of mite IV similar to somite III, but thinner and female unarmed. Female with large gonopore shorter; dorsal surface with few short setae on anterior mesial margin of coxa, surround- anterolaterally; pleura thinner and shorter 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 253 than on somite III, directed posterolaterally; institution, and they cannot be examined. dorsal surface obliquely ¯attened anterolat- Thomassin's (1969: text-®g. 12) distribution erally; margins with long plumose setae. So- map is correct for the localities marked, but mite V wider than somite IV; lateral margins the species spans a wide geographic range with plumose setae; pleura absent. Somite VI between Madagascar and Samoa. The Japa- slightly broader than somite V; dorsal surface nese name for this species is ``Togenashi-Ku- with short transverse rows of setae laterad of dahigegani'' (Asakura, personal commun.). midline and posteriorly; pleura absent. It is obvious from the illustrations of Ca- Females with uniramous, paired pleopods lado (1995) that the specimens she examined on somites II±V; males without pleopods. and cited as A.``symnista'' are A. microps. Telson of male (®g. 82F) broadly trian- This was con®rmed by direct examination of gular, longer than wide, with broadly round- those specimens. As discussed further under ed tip; proximal third thickly calci®ed, in- A. symmysta, this misidenti®cation is proba- ¯ated dorsally; distal two-thirds with large bly the basis for Calado's (1997a) subsequent decalci®ed windows laterad of thin medial description of A. edsoni, which is a synonym calci®ed strip, margins of windows with long of A. symmysta. Calado (1995: pl. 21, ®g. a) simple setae; median longitudinal groove ex- also omitted the setal ®eld in the carapace tending one-half length; proximolateral an- drawing. gles with patch of short simple setae; mar- As with most albuneids, little is known of gins with long simple setae. Telson of female the biology of A. microps. Ovigerous females (®g. 82G) ¯attened, ovate, and evenly calci- are known from February and November. ®ed; median groove similar to male but This species is fairly common in the Philip- ¯anked along distal half by long thin simple pines, as shown by the abundant material cit- setae; proximolateral angle with patch of se- ed above and by the 65 specimens listed by tae, margins with long simple setae. SereÁne and Umali (1965). MOV J40128 is DISTRIBUTION: From Zanzibar and Mada- the type and only known host for the bopyrid gascar north to Oman and eastward to the isopod Albunione australiana Markham and Philippines and New Caledonia, in 3±45 m Boyko, 1999. depth. Also reported from Japan (Miyake, This species was considered the senior 1978). synonym of A. elioti until Boyko and Harvey MAXIMUM SIZE: Males: 11.6 mm cl; fe- males: 15.7 mm cl. (1999) clearly showed the distinctiveness of both species. The shape of both the telson of TYPE SPECIMEN: BMNH 1937.6.7.3 (holo- type). the male and the distal peduncular segments can be used to tell the two species apart. The TYPE LOCALITY: ``Sooloo Island'' (ϭ Sulu Archipelago, Philippines). telson of the male of A. microps is heavily REMARKS: Although it is surprising that calci®ed and somewhat in¯ated proximally, two of three specimens of the type series of but partially decalci®ed distally and narrow- A. microps reported by Henderson (1888) ing to a produced tip. An oblique row of long were transferred from BMNH to other mu- setae is present just proximal to the demar- seums, this was apparently common practice cation line between the calci®ed and decal- with much of the ``Challenger'' material ci®ed regions on each side of the median (Lingwood, 1981). line. In A. elioti, the telson of the male is Thomassin's (1969) material of ``Albunea narrowly triangular, fully calci®ed, with short microps'' from Madagascar was a mixture of thick setae on a strong medial ridge, and that species and A. elioti. This is clearly lacks the oblique row of setae. The mesial shown by his illustrations, as ®gures 2 and margins of the distal peduncular segments 3b are A. microps, while all of the ®gures on are strongly convex in A. elioti but only plate 2 are A. elioti (see also Boyko and Har- slightly so in A. microps. In addition, the cor- vey, 1999). How many specimens of each nea of A. elioti is more posterolaterally dis- taxon were present in Thomassin's (1969) placed from the tip of the distal peduncular samples is unknown, as he did not deposit segment. Albunea microps is the sister spe- any specimens in the MNHN or any other cies to A. elioti and typi®es the ``microps- 254 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 group'' of Albunea, which also includes A. almost united but separate; CG11 present. elioti and A. galapagensis. Rostrum present, exceeding posterior margin of ocular plate by approximately one-half Albunea elioti Benedict, 1904 length of ocular plate. Ocular plate subquad- Figures 83, 84 rate. Distal peduncular segments dorsoven- Albunea elioti Benedict, 1904: 623, ®g. 2*. ± Gor- trally ¯attened and oblong in shape, rounded don, 1938: 187 (list). ± CoeÃlho and Calado, at tip, separated along mesial margins, lateral 1987: table 1. ± Boyko and Harvey, 1999: 386, margins sinuous, mesial margins convex. 400 (list), 402 (key)*. ± Boyko, 1999: 145 Cornea on lateral margin at tip. Dactylus of (list). pereopod II with heel produced and rounded. Albunea microps: Thomassin, 1969: 140±143 Dactylus of pereopod III with heel produced (part), pl. 2, ®gs. 1±9. ± Calado, 1995: 46±49, and rounded. Dactylus of pereopod IV even- pl. 4, ®g. e, pl. 5, ®g. d, pl. 12, ®gs. a±f (not ly sinuous from base to tip, with shallow in- Albunea microps Miers, 1878). dent. Telson of male elongate triangular with MATERIAL EXAMINED: Madagascar: Sta. rounded tip; median area thickly calci®ed, 15/15, Tulear, 1976, coll. P. Galenon: 1 (, marginal area weakly calci®ed, medial row 10.4 mm cl (MNHN-Hi 88). of short thick simple setae. Telson of female Seychelles: Sta. 4, 32 m, Sept. 2, 1980, similar to male but broader, ¯attened, evenly coll. ORSTOM 1980±Reves 2: 1 &, 14.4 calci®ed, with medial row of thin setae. mm cl (MNHN-Hi 195). DESCRIPTION: Carapace (®g. 83A) wider Australia: Western Australia: Barrow Is- than long. Anterior margin slightly concave land, 3 ft (ϭ 0.9 m), June 6, 1964, coll. W. on either side of ocular sinus, becoming con- H. Butler: 1 (, 12.0 mm cl (WAM 23394). vex laterally, with seven to nine large spines Caroline Islands: Sta. 137, Kapinga- (n ϭ 5) along length. Rostrum as small acute marangi Atoll, Polim Reef, ¯at region next tooth, extending halfway across ocular plate. to Tipongowakaram Pass, 01Њ02Ј00ЉN, Ocular sinus smoothly concave and unarmed. 154Њ45Ј14ЉE, Aug. 12, 1954, coll. George Frontal region smooth; setal ®eld broad an- Vanderbilt Foundation: 1 &, unmeasurable teriorly and posteriorly; posterior lateral el- (USNM 104746). ements broad and adjoining posterior lateral Fiji: Viti [Levu] Island, coll. unknown: 1 elements of CG1. CG1 parallel to anterior &, 17.0 mm cl (ZMH K±5134 ex Museum margin of carapace, faintly sinuous, strongly Goddefroy 941). crenulate, divided into medial fragment and Japan: Iko, Kuroshima Island, Yaeyama curved posteriorly displaced lateral elements. Islands, intertidal, April 28, 1998, coll. K. Mesogastric region smooth; CG2 present as Nomura: 1 (, 16.65 mm cl (CBM-ZC 5334). one or two elements; CG3 broken into one Tonga: South of Ohonua Harbor, Eua Is- to four long and short elements approximate- land, Tongatabu Group, 21Њ20Ј15ЉS, ly equally spaced between posterior lateral 174Њ58Ј14ЉW, 0 ± 5 f t ( ϭ 0±1.5 m), Nov. 2, elements of CG1; CG4 with one or two short 1993, coll. J. T. Williams et al.: 1 (, 16.5 medial elements and two longer lateral ele- mm cl (USNM 281472). ments spaced approximately equally between Samoa: ``Samoa'', coll. unknown: 1 &, longer supralateral elements of CG4. Hepatic 16.2 mm cl, holotype (USNM 26169). region smooth, with oblique setose groove at DIAGNOSIS: Carapace wider than long, cov- median of lateral margin. Epibranchial re- ered with strongly setose grooves. Anterior gion generally triangular, smooth; posterolat- margin with seven to nine spines on either eral margin with three short rows of setae. side of ocular sinus. Setal ®eld with thick Metagastric region smooth; CG5 ranging lateral elements and diffuse anterior margin. from entire to four short elements. CG6 CG1 with separate posterior lateral elements; strongly crenulate, strongly anteriorly con- CG4 with one or two short medial elements cave medially and sloping out to anteriorly and two longer lateral elements spaced ap- convex lateral thirds. CG7 oblique, almost proximately equally between longer supra- reaching lateral margins of median segment lateral elements of CG4; CG5 ranging from of CG6. Cardiac region smooth; CG8 present entire to four short elements; CG6 and CG7 as three long elements. CG9 present as two 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 255

Fig. 83. Albunea elioti Benedict, 1904: A, B, &, 16.2 mm cl, USNM 26169, holotype; C±K, (, 16.5 mm cl, USNM 281472. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Ocular peduncles, dorsal view. D. Left antennule, lateral view. E. Left antenna, lateral view. F. Left mandible, mesial view. G. Left maxillule, lateral view. H. Left maxilla, lateral view. I. Left maxilliped I, lateral view. J. Left maxilliped II, lateral view. K. Left maxilliped III, lateral view. Scale ϭ 1.6 mm (C), 2.2 mm (G), 3.3 mm (B, F, J), 4.4 mm (D, E, H, I, K), and 5.9 mm (A). 256 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 short lateral grooves with gap at midline. IV expanded distally, with long plumose se- CG10 present as two long lateral fragments, tae on dorsal, ventral, and distal margins, and with short gap between fragments. CG11 row of setae on dorsolateral surface. Segment present. Branchial region with numerous III with long plumose setae on dorsal and short, transverse rows of setae. Posterior ventral margin and in short row on surface. margin deeply and evenly convex, with sub- Segment II short, widening distally, with plu- marginal groove reaching less than one-third mose setae on margins and scattered on lat- up margin of posterior concavity. Branchios- eral surface; antennal acicle long, thin, and tegite with short anterior submarginal spine; just exceeding distal margin of segment IV, anterior region with scattered short trans- with long plumose setae on dorsal margin. verse lines ventral to linea anomurica; with Segment I rounded proximally, ¯attened ven- many short rows of setae and sparsely cov- trolaterally, with long plumose setae on mar- ered with long plumose setae ventrally; pos- gins; lateral surface with rounded spine dor- terior region membranous with numerous ir- sally, low semicircular dorsolateral lobe ven- regular fragments and sparsely covered with trodistal to spine; segment with ventromesial long plumose setae. antennal gland pore. Ocular plate (®g. 83B, C) subrectangular Mandible (®g. 83F) incisor process with with shallow median indentation; median pe- one tooth; cutting edge with one tooth. Palp duncular segments bluntly diamond-shaped three-segmented, with plumose setae on mar- and located ventral to ocular plate and distal gins and long, thick, simple setae arising peduncular segments; not visible in dorsal from bend in second segment and on distal view. Distal peduncular segments irregularly margin of terminal segment. oblong, with convex lateral and mesial mar- Maxillule (®g. 83G) distal endite proxi- gins, two lateral notches present with oblong mally narrow, widening to in¯ated distal end, cornea located in distal notch; mesial mar- with thick simple setae on distal margin. gins widely separated along all of length; Proximal endite with thick simple setae on mesial and lateral margins with long plumose distal margin and thin plumose setae on dor- setae; tuft of plumose setae at proximolateral sal margin. Endopodal external lobe truncate ventral angle and short row of plumose setae distally and curled under; internal lobe re- proximoventral to cornea notch. duced with six thick setae at distolateral mar- Antennule (®g. 83D) with segment III nar- gin. row proximally, expanding distally to two Maxilla (®g. 83H) exopod evenly rounded, times proximal width; with plumose setae on with plumose setae along distal margin. Sca- dorsal and ventral margins and sparsely scat- phognathite bluntly angled on posterior lobe, tered on lateral surface; dorsal exopodal ¯a- with plumose setae. gellum with 56±64 articles (n ϭ 3), long plu- Maxilliped I (®g. 83I) epipod with plu- mose setae on dorsal and ventral margins; mose setae on margins, distolateral surface ventral endopodal ¯agellum short with one and mesial surface (epipod shown curled in segment (n ϭ 4) and plumose setae on dorsal ®g. 83I). Endite tapered distally and subequal and ventral margins. Segment II medially in- to ®rst segment of exopod. Exopod with two ¯ated in dorsal view, with plumose setae on segments; proximal segment narrow, margins dorsal and ventral margins and scattered on parallel, with plumose setae; distal segment ventrolateral third of surface. Segment I lon- spatulate, longer than wide, broadest medi- ger than wide, unarmed; dorsal third of lat- ally, margins and mesioventral surface with eral surface rugose with long plumose setae; long plumose setae. Endopod ¯attened and long plumose setae on dorsal and ventral elongate, reaching nearly to distal end of margins. proximal exopodal segment; plumose setae Antenna (®g. 83E) with segment V ap- on margins and median of lateral surface. proximately 2.5 times longer than wide, with Maxilliped II (®g. 83J) dactylus evenly long plumose setae on dorsal margin and rounded, length equal to width, with thick scattered on lateral surface; ¯agellum with simple setae distally and on distolateral sur- ®ve articles (n ϭ 4), long plumose setae on face. Propodus two times wider than long, dorsal, ventral, and distal margins. Segment slightly produced at dorsodistal angle, with 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 257 plumose setae on dorsal margin and patch of few short rows of setae; fully calci®ed. Ba- long simple setae on lateral surface and ven- sis-ischium incompletely fused, unarmed. trolateral angle. Carpus not produced dorso- Coxa unarmed. distally, approximately two times longer than Pereopod II (®g. 84B) dactylus smooth; wide; long simple setae on dorsal and distal base to heel slightly concave, heel smoothly margins and in few small patches on surface. rounded, heel to tip with wide, acute indent, Merus approximately two times longer than tip acute, tip to base broadly convex distally wide, margins parallel; with simple and plu- and slightly concave proximally; lateral sur- mose setae on margins and scattered on sur- face smooth, with several small tufts of short face. Basis-ischium incompletely fused with setae in generally straight line across medi- plumose setae on margins. Exopod one-third oproximal surface, several widely spaced longer than merus, ¯agellum with one elon- submarginal tufts of short setae dorsodistally; gate segment. mesial surface smooth, ventral margin with Maxilliped III (®g. 83K) dactylus with long plumose setae, dorsal margin with short rounded tip; long plumose setae on margins simple setae, with patch of long plumose se- and lateral surface. Propodus slightly in¯ated tae at base. Propodal dorsal surface smooth, medially, with longitudinal median row of with ventral margin in¯ated and rounded; plumose setae on lateral surface; margins oblique row of long plumose setae on distal with plumose setae. Carpus produced onto margin of lateral surface; distal and ventral propodus almost one-half length of propo- margin with long plumose setae; dorsolateral dus; lateral surface with two rows of plumose surface as narrow, oblique, ¯attened shelf, setae on surface; plumose setae on margins. with short setae on dorsal margin and long Merus in¯ated, unarmed, with plumose setae plumose setae on ventral margin; mesial sur- on margins and scattered on lateral surface. face with elevated, curved, setose ridge from Basis-ischium incompletely fused, with weak ventral junction with dactylus almost to ven- crista dentata of one or two teeth. Exopod tral proximal junction with carpus; small de- two-segmented: proximal segment small; calci®ed region just distal to junction with distal segment styliform, tapering, approxi- carpus. Carpus slightly produced and gently mately one-half length of merus; with plu- rounded dorsodistally, dorsal margin un- mose setae on margins; without ¯agellum. armed; lateral surface smooth, with setose Pereopod I (®g. 84A) dactylus curved and mat on distodorsal quarter and irregular, in- tapering; lateral and mesial surfaces smooth; terrupted row of rugae and submarginal ele- dorsal margin proximally rugose, with long vated ridge ventrally, rugae and ridge with plumose and short simple setae; ventral mar- long plumose setae; margins with long plu- gin with short simple setae. Propodal lateral mose setae; mesial surface smooth with scat- surface with numerous short, transverse rows tered patches of long plumose setae in sub- of setose rugae; dorsal margin unarmed; ven- dorsal and submarginal rows. Merus with tral margin distally produced into acute large median decalci®ed window covering spine; cutting edge lacking teeth, lined with nearly all of lateral surface, with few scat- long plumose setae; dorsal margin with long tered setae on surface and margins; mesial plumose setae, ventral margin with short surface nearly smooth, with few setae, and simple setae. Carpus with dorsodistal angle small decalci®ed area near junction with ba- produced into strong corneous-tipped spine; sis-ischium. Basis-ischium incompletely dorsal margin with short transverse grooves fused and unarmed. Coxa unarmed. behind spine; dorsal and distal margins with Pereopod III (®g. 84C) dactylus with base long plumose setae; lateral surface with to heel convex, heel broadly rounded and small distal rugose area, few transverse se- low, heel to tip with broadly concave indent, tose ridges on distal half of surface; mesial tip acute, tip to base smoothly convex dis- surface smooth, with few median rows of se- tally to straight proximally; lateral surface tae, margins with long plumose setae. Merus smooth, with several small tufts of short se- unarmed; lateral surface with scattered trans- tae in generally straight line across medi- verse rows of long plumose setae, margins oproximal surface, dorsodistal margin with with long plumose setae; mesial surface with tufts of short setae; ventral margin with long 258 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Fig. 84. Albunea elioti Benedict, 1904: A±D, (, 12.0 mm cl, WAM 23394; E, (, 10.4 mm cl, MNHN Hi-88; F, (, 16.5 mm cl, USNM 281472; G, &, 16.2 mm cl, USNM 26169, holotype. A. Left pereopod I, lateral view. B. Left pereopod II, lateral view. C. Left pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 3.3 mm (E±G) and 4.4 mm (A±D). plumose setae, dorsal margin with short sim- pus. Carpus produced dorsodistally, exceed- ple and plumose setae; mesial surface ing proximal margin of propodus by approx- smooth, with plumose setae proximally at imately one-third length of propodus; dor- junction with propodus. Propodus not in¯at- solateral margin unarmed; lateral surface ed dorsoventrally; lateral surface smooth, slightly rugose dorsodistally, with mat of with long plumose setae in oblique row, with short setae and row of setae ventrally; mesial simple setae on dorsal margins; dorsolateral surface smooth, with long plumose setae on surface narrow, oblique, ¯attened, with se- margins. Merus smooth, with large decalci- tose mat; mesial surface with scattered long ®ed window covering nearly half of lateral setae on and near distal margin, with small surface medially; dorsal and ventral margins decalci®ed window near junction with car- unarmed, with long plumose setae; latero- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 259 distal margin with long plumose setae; me- short thick setae on posterolateral angle; sial surface smooth, with small decalci®ed pleura thinner and shorter than on somite II, window at junction with basis-ischium. Ba- directed posterolaterally, with setae as in so- sis-ischium incompletely fused and unarmed. mite II; anterolateral angle acute; dorsal sur- Coxa unarmed. Female with large gonopore face obliquely ¯attened anterolaterally. So- on anterior mesial margin of coxa, surround- mite IV similar to somite III, but thinner and ed with short plumose setae; male without shorter; dorsal surface with few thick setae pore. posterolaterally; pleura thinner and shorter Pereopod IV (®g. 84D) dactylus with base than on somite III, directed posterolaterally; to tip convex to concave, tip acute, tip to dorsal surface obliquely ¯attened anterolat- base convex; lateral surface smooth, ventral erally; margins with long plumose setae. So- margin with long plumose setae, dorsal mar- mite V wider than somite IV; lateral margins gin with short simple setae; mesial surface with plumose setae; pleura absent. Somite VI with dorsal decalci®ed region, demarcated subequal to somite V; dorsal surface with ventrally by longitudinal elevated ridge with short transverse rows of setae laterad of mid- row of short setae; with setose punctations line, anteriorly and posteriorly; pleura absent. ventral to decalci®ed window. Propodus ex- Females with uniramous, paired pleopods panded dorsally and ventrally; ventral expan- on somites II±V; males without pleopods. sion almost reaching ventral margin of dac- Telson of male (®g. 84F) elongate trian- tylus, margin with long plumose setae; dorsal gular, two times longer than wide, with expansion with row of long plumose setae pointed rounded tip; median area thickly cal- dorsally, oblique area with mat of short sim- ci®ed, with dorsodistally produced tip; mar- ple setae; lateral and mesial surfaces smooth. ginal area weakly calci®ed, ¯attened and Carpus not produced dorsodistally; ventral thin; median longitudinal groove extending three-fourths of lateral surface and mesial one-half length, ¯anked on distal half by surface smooth, dorsodistal quarter of lateral dense row of short thick simple setae con- surface with mat of short setae; dorsal mar- tinuing to produced tip; proximolateral an- gin with short simple and long plumose se- gles each with patch of short simple setae; tae; ventral margin with short simple setae. margins with long simple setae. Telson of fe- Merus with scattered short transverse rows of male (®g. 84G) similar to male but broader, setae on lateral surface, dorsal and ventro- ¯attened, and evenly calci®ed; median distal margins with long plumose setae; groove similar but ¯anked along distal half proximoventral third of mesial surface with by long thin simple setae; proximolateral an- large decalci®ed window. Basis-ischium in- gle with denser patch of setae than male. completely fused and unarmed. Coxa un- DISTRIBUTION: Known from Madagascar armed. and the Seychelles eastward to Samoa, in up Abdomen (®g. 84E) with somite I wider to 32 m depth. than long, widest posteriorly; dorsal surface MAXIMUM SIZE: Males: 16.65 mm cl; fe- with anterior margin straight; posterior mar- males: 17.0 mm cl. gin curved with elevated submarginal row of TYPE SPECIMEN: USNM 26169 (holotype). short setae; with small transverse decalci®ed TYPE LOCALITY: Samoa. windows laterad of segment median. Somite REMARKS: As previously noted, Thomas- II dorsal surface with submarginal transverse sin's (1969) material of ``Albunea microps'' ridge anteriorly; with small transverse decal- from Madagascar was a mixture of that spe- ci®ed windows laterad of segment median cies and A. elioti. This is clearly shown by just anterior to submarginal ridge; with tuft his illustrations, as ®gures 2 and 3b are A. of setae at posterolateral angle, extending microps, while all of the ®gures on plate 2 onto pleura posteromesially; pleura expanded are A. elioti (see also Boyko and Harvey, and directed slightly anterolaterally; lateral 1999). How many specimens of each taxon margins rounded, anterior and lateral mar- were present in Thomassin's (1969) samples gins with long plumose setae, posterior mar- is unknown, as he did not deposit any spec- gin with short setae. Somite III similar to so- imens in MNHN or any other institution, and mite II, but narrower, shorter; small tuft of they are unavailable for examination. 260 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Calado's (1995) illustration of the cara- with three or four short medial elements be- pace (as A. microps) is too slender and does tween longer supralateral elements of CG4, not accurately depict the subquadrate shape CG5 two short, irregularly convex, trans- typical of this species. verse elements; CG6 and CG7 separate; CG8 This species was considered a synonym of present; CG11 present. Rostrum present, A. microps until Boyko and Harvey (1999) reaching posterior margin of ocular plate. gave the distinguishing characters that sepa- Ocular plate subquadrate. Distal peduncular rate the two species (see under A. microps segments dorsoventrally ¯attened and oblong for details). Albunea microps is the sister in shape, rounded at tip, approximated along species to A. elioti. mesial margins, lateral margins convex, me- sial margins straight. Cornea on lateral mar- Albunea bulla, new species gin at tip. Dactylus of pereopod II with heel Figures 85, 86 produced and rounded. Dactylus of pereopod III with heel produced and rounded. Dactylus Albunea symnista [sic]: Gordon, 1938: 187 (part)* of pereopod IV evenly sinuous from base to (not Albunea symmysta (Linnaeus, 1758)). ?Albuneidae sp. DiSalvo et al., 1988: 458. tip, with shallow indent. Telson of male di- vided into buttonlike proximal two-thirds MATERIAL EXAMINED: Taiwan: Ma-Kung, and mucronate, tapering, distal third; proxi- Peng-Hui Island, Sept. 15, 1996, coll. un- mal two-thirds rounded, heavily calci®ed, known: 1 &, 17.4 mm cl, paratype (NTOU). dorsally in¯ated, row of long setae ¯anking Australia: Queensland: Samovrez Reef, 8 distal half of telson, medial transverse row of m, Sept. 1974, coll. N. Coleman: 1 &, 19.3 long setae present; distal third of telson mm cl, paratype (AM P20456); Cape Bowl- weakly calci®ed, sharply tapering distally ing Green, 16 fms (ϭ 29.3 m), Nov. 23, with produced tip. Telson of female ovate, 1962, coll. W. Goode on R/V ``Dorothea'': 1 tapering slightly distally, tip slightly pro- (, 11.8 mm cl, 1 &, 17.9 mm cl, paratypes duced. (WAM 23387); New South Wales: Off DESCRIPTION: Carapace (®g. 85A) wider Richmond River mouth, off Ballina, 28Њ52ЈS, than long. Anterior margin faintly concave 153Њ34ЈE, 16 m, Oct. 6, 1962, coll. unknown: on either side of ocular sinus, becoming con- 1 (, 15.4 mm cl, holotype (AM P15353); vex laterally with 8±10 large spines (n ϭ 5) Lord Howe Island, 31Њ33ЈS, 159Њ05ЈE, coll. on each side along length. Rostrum a small unknown: 1 &, 18.9 mm cl, allotype (AM acute tooth, reaching proximal margin of oc- P1925). ular plate. Ocular sinus smoothly concave. New Caledonia: ``New Caledonia,'' 1903, Frontal region smooth; setal ®eld broad an- coll. A. Milne Edwards: 1 &, 19.4 mm cl teriorly and narrow posteriorly; posterior lat- (MNHN-Hi 150). eral elements reduced to narrow bands of se- Fiji: Viti Island: 1 (, 7.5 mm cl, paratype tae. CG1 parallel to anterior margin of car- (ZMH K±5137 ex Museum Goddefroy apace, convex, sinuous, strongly crenulate, 2120). divided into medial fragment and curved, Pitcairn Island: Sta. PIT-UI, Haul 13, 1 posteriorly displaced lateral elements. Me- mi northwest of Pitcairn Island, 25Њ02±04ЈS, sogastric region smooth; CG2 present as two 130Њ06±07ЈW, 26±30 fms (ϭ 47.6±54.9 m), or three short elements, anteriorly and pos- Sept. 20, 1967, coll. National Geographic teriorly displaced; CG3 broken into three to Society±Smithsonian-Bishop Museum Mar- six short curved elements between posterior quesas Expedition: 1 unsexable anterior half lateral elements of CG1, medial two ele- with pereopod I, 17.7 mm cl (USNM ments shortest; CG4 with three or four short 304304). medial elements between longer supralateral DIAGNOSIS: Carapace wider than long, cov- elements of CG4, medial one or two ele- ered with strongly setose grooves. Anterior ments anteriorly displaced, lateral short ele- margin with 8±10 spines on either side of ments posteriorly displaced. Hepatic region ocular sinus. Setal ®eld with thin lateral el- smooth, with oblique setose groove at me- ements and concave anterior margin. CG1 dian of lateral margin, nearly reaching pos- with separate posterior lateral elements; CG4 terior element of CG1. Epibranchial region 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 261

Fig. 85. Albunea bulla, n. sp.: A±J, &, 19.3 mm cl, AM P20456, paratype. A. Carapace, branchios- tegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 2.2 mm (B), 3.0 mm (F), 3.3 mm (E, I), 4.4 mm (D, J), 6.0 mm (C), 6.7 mm (G, H), and 7.1 mm (A). 262 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 generally triangular, smooth; posterolateral ¯ated in dorsal view, with plumose setae on margin with one or two oblique rows of short dorsal and ventral margins and scattered on setae. Metagastric region smooth; CG5 pre- mediolateral surface. Segment I wider than sent as two short, irregularly convex, trans- long, unarmed; dorsal quarter of lateral sur- verse elements. CG6 strongly crenulate, face rugose, with long plumose setae; long strongly anteriorly concave medially, and plumose setae on dorsal and ventral margins. sloping out to anteriorly convex lateral Antenna (®g. 85D) with segment V ap- thirds. CG7 oblique, not reaching lateral proximately 2.5 times longer than wide, with margins of median segment of CG6. Cardiac long plumose setae on dorsal and distal mar- region smooth; CG8 present as one long me- gins and scattered on lateral surface; ¯agel- dial element. CG9 present as two short lateral lum with six articles (n ϭ 5), long plumose grooves with broad gap at midline. CG10 setae on dorsal, ventral, and distal margins. present as two short lateral elements. CG11 Segment IV expanded distally, with long plu- present as two or three short elements. Post- mose setae on dorsal, ventral, and distal mar- CG11 element absent. Branchial region with gins, and interrupted row of short setae on numerous short and long transverse rows of dorsolateral surface. Segment III with long setae in anterior half. Posterior margin deep- plumose setae on ventral margin; short sim- ly and evenly convex, with submarginal ple setae on dorsal margin and scattered on groove reaching one-third up margin of pos- surface. Segment II short, widening distally, terior concavity. Branchiostegite with strong dorsally rugose, with plumose setae on mar- anterior submarginal spine; anterior region gins and short simple setae scattered on lat- with scattered short, transverse lines ventral eral surface; antennal acicle long, thin, and to linea anomurica; with many short rows of reaching distal margin of segment IV, with setae and sparsely covered with long plu- long plumose setae on dorsal margin. Seg- mose setae ventrally; posterior region mem- ment I rounded proximally, ¯attened ventro- branous, with numerous irregular fragments laterally, with long plumose setae on dorsal and sparsely covered with long plumose se- and distoventral margins, and short simple tae. setae on surface rugae behind spine; lateral Ocular plate (®g. 85B) triangular with surface with acute spine dorsodistally; low deep median indentation; median peduncular semicircular dorsolateral lobe ventrodistal to segments present as small ovate, calci®ed ar- spine, margin of lobe with long plumose se- eas lateral to ocular plate. Distal peduncular tae; segment with ventromesial antennal segments elongate, subtriangular, with con- gland pore. vex lateral and straight mesial margins, cor- Mandible (®g. 85E) incisor process with nea covering lateral portion of distal tip; lat- two teeth; cutting edge with one tooth. Palp eral margins with faint notch one-fourth dis- three-segmented, with plumose setae on mar- tal from base; mesial margins approximated gins and long, thick, simple setae arising along length; lateral and mesial margins with from bend in second segment and on distal long plumose setae; tuft of plumose setae at margin of terminal segment. proximolateral ventral angles and ventrome- Maxillule (®g. 85F) distal endite proxi- dial longitudinal row of plumose setae ex- mally narrow, widening to in¯ated distal end, tending from tuft to three-fourths length of with thick simple setae on distal margin and peduncle. thin simple setae on dorsal margin. Proximal Antennule (®g. 85C) with segment III nar- endite with thick simple setae on distal mar- row proximally, expanding distally to three gin. Endopodal external lobe truncate distally times proximal width; plumose setae on dor- and curled under; internal lobe reduced with sal and ventral margins and sparsely scat- ®ve thick setae at distolateral margin and one tered on lateral surface; dorsal exopodal ¯a- thick seta in median of lobe. gellum with 67±77 articles (n ϭ 2), long plu- Maxilla (®g. 85G) exopod evenly rounded, mose setae on dorsal and ventral margins; with plumose setae along distal margin. Sca- ventral endopodal ¯agellum with three or phognathite bluntly angled on posterior lobe, four articles (n ϭ 4), plumose setae on dorsal with plumose setae. and ventral margins. Segment II medially in- Maxilliped I (®g. 85H) epipod with plu- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 263 mose setae on margins, distolateral surface, tapering; lateral and mesial surfaces smooth; and mesial surface. Endite tapered distally dorsal margin with long plumose setae; ven- and subequal to ®rst segment of exopod. Ex- tral margin with short simple setae. Propodal opod with two segments: proximal segment lateral surface with numerous short, trans- narrow, margins parallel, with long plumose verse rows of setose rugae; dorsal margin un- setae; distal segment spatulate, longer than armed; ventral margin distally produced into wide, broadest medially, margins and mesio- acute spine; cutting edge lacking teeth, lined dorsal surface with long plumose setae. En- with long plumose setae; dorsal margin with dopod ¯attened and elongate, reaching two- long plumose setae, ventral margin with thirds of distance to distal end of proximal short simple setae. Carpus with dorsodistal exopodal segment; plumose setae on margins angle produced into rounded projection, and median of lateral surface. small dorsally directed acute spine on disto- Maxilliped II (®g. 85I) dactylus evenly dorsal mesial margin; dorsal margin with rounded, length subequal to width, with thick short transverse grooves behind distal projec- simple setae distally and on distolateral sur- tion; dorsal and distal margins with short plu- face. Propodus two times wider than long, mose setae; lateral surface with small distal slightly produced at dorsodistal angle, with rugose area, with few transverse setose ridg- plumose setae on dorsal margin and patch of es on distal half of surface; mesial surface long simple setae on dorsodistal and ven- smooth with medial transverse row of setae, trodistal angles. Carpus not produced dorso- margins with long plumose setae. Merus un- distally, approximately two times longer than armed; lateral surface with scattered trans- wide; long simple setae on dorsal margin and verse rows of long plumose setae, distal and in patches on dorsodistal and ventrodistal an- mediodorsal margins with long plumose se- gles. Merus approximately three times longer tae; mesial side with few scattered setae; ful- than wide, margins parallel; with long simple ly calci®ed. Basis-ischium incompletely and plumose setae on margins and scattered fused, unarmed. Coxa unarmed. on lateral surface. Basis-ischium incomplete- Pereopod II (®g. 86B) dactylus smooth; ly fused, with plumose setae on margins. Ex- base to heel faintly convex, heel low and opod 1.7 times longer than merus, ¯agellum rounded, subquadrate, heel to tip with broad, with one elongate article, approximately as subacute indent, tip subacute, tip to base long as carpus. broadly convex; lateral surface smooth, with Maxilliped III (®g. 85J) dactylus oblong several small tufts of short setae in generally with rounded tip; long plumose setae on mar- straight line across medioproximal surface, gins and in medial row on lateral surface. several widely spaced submarginal tufts of Propodus dorsodistally in¯ated, with longi- short setae dorsodistally; mesial surface tudinal median row of plumose setae on lat- smooth, ventral margin with long plumose eral surface and three or four short patches setae, dorsal margin with short simple setae, of long plumose setae submarginally ven- with patch of long plumose setae at base. trally; dorsal margin with long plumose se- Propodal dorsal surface smooth, with ventral tae. Carpus produced onto propodus approx- margin in¯ated and rounded; oblique row of imately two-thirds length of propodus; lateral long plumose setae on distal margin of lateral surface with two transverse rows of long plu- surface; distal and ventral margins with long mose setae; long plumose setae on dorsal and plumose setae; dorsolateral surface as nar- distal margins. Merus in¯ated, unarmed, with row, oblique, ¯attened shelf, with short setae long plumose setae on margins and few scat- on dorsal margin and long plumose setae on tered small patches on lateral surface. Basis- ventral margin; mesial surface with elevated, ischium incompletely fused, with weak crista curved setose ridge from ventral junction dentata of four teeth. Exopod two-segment- with dactylus almost to ventral proximal ed: proximal segment small; distal segment junction with carpus. Carpus strongly pro- styliform, bluntly rounded distally, approxi- duced, tapered distally, and rounded dorso- mately one-third length of merus; with plu- distally, dorsal margin smooth; lateral sur- mose setae on margins; without ¯agellum. face smooth, produced area smooth, irregu- Pereopod I (®g. 86A) dactylus curved and lar, interrupted row of rugae and submarginal 264 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Fig. 86. Albunea bulla, n. sp.: A±E, G, &, 19.3 mm cl, AM P20456, paratype; F, (, 15.4 mm cl, AM P15353, holotype. A. Left pereopod I, lateral view. B. Left pereopod II, lateral view. C. Left pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 3.0 mm (F), 3.3 mm (G), and 6.0 mm (A±E). elevated ridge ventrally, rugae and ridge with and ventral margins, few scattered long plu- long plumose setae; dorsal margin with short mose setae on surface; mesial surface nearly plumose setae, ventral margin with long plu- smooth with two long rows of setae. Basis- mose setae; mesial surface smooth, with row ischium incompletely fused and unarmed. of long plumose setae distally, ventrally, and Coxa unarmed. subdorsally. Merus with large median decal- Pereopod III (®g. 86C) dactylus with base ci®ed window covering nearly all of lateral to heel straight, heel low and subquadrate, surface, long plumose setae on dorsodistal heel to tip with broadly concave indent, tip 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 265 subacute, tip to base smoothly convex; lateral ventral four-®fths of lateral surface and me- surface smooth, with several small tufts of sial surface smooth, dorsodistal one-®fth of short setae in generally straight line across lateral surface with mat of short simple setae; medioproximal surface, dorsodistal margin dorsal margin with short simple and long with tufts of short setae; ventral margin with plumose setae; ventral margin with short long plumose setae, dorsal margin with short simple setae; mesial surface decalci®ed me- simple and plumose setae; mesial surface dially. Merus lateral surface with scattered smooth, with plumose setae proximally at short transverse rows of setae, dorsal margin junction with propodus. Propodus not in¯at- with long plumose setae; proximoventral ed dorsoventrally; lateral surface smooth, fourth of mesial surface with large decalci- with long plumose setae in oblique row, sim- ®ed window. Basis-ischium incompletely ple setae on dorsal margin, plumose setae on fused and unarmed. Coxa unarmed. ventral margin; dorsolateral surface narrow, Abdomen (®g. 86E) somite I wider than oblique, ¯attened, without setose mat; mesial long, widest posteriorly; dorsal surface with surface smooth. Carpus produced dorsodis- anterior margin straight; posterior margin tally, exceeding proximal margin of propo- curved, with elevated submarginal row of dus by one-half length of propodus; tip sub- short setae; small transverse decalci®ed win- acute, dorsal margin unarmed; lateral surface dows laterad of segment median. Somite II slightly rugose dorsodistally, with mat of dorsal surface with irregular, submarginal, short setae and two interrupted rows of setae transverse ridge anteriorly; with small, trans- ventrally; mesial surface smooth, with long verse, decalci®ed windows laterad of seg- plumose setae on distal margin and in short ment median just anterior to submarginal oblique row on distal surface. Merus smooth, ridge; pleura expanded and directed laterally; narrow, and almost cylindrical, with large de- anterolateral margins rounded, anterior and calci®ed window covering nearly half of lat- lateral margins with long plumose setae, pos- eral surface medially; dorsal and ventral mar- terolateral angle rounded, posterior margin gins unarmed, dorsodistal and ventral mar- with short setae; posteromesial angle with gins with long plumose setae; mesial surface mat of short simple setae. Somite III similar smooth. Basis-ischium incompletely fused to somite II, but narrower, shorter, anterior and unarmed. Coxa unarmed. Female with submarginal windows present; pleura thinner large gonopore on anterior mesial margin of and shorter than on somite II, directed pos- coxa, surrounded with short plumose setae; terolaterally proximally and anterolaterally male with small pore. distally, with setae as in somite II; anterolat- Pereopod IV (®g. 86D) dactylus with base eral angle subacute; dorsal surface obliquely to tip convex proximally, with indistinct heel ¯attened anterolaterally, with submarginal and indent and almost straight from indent to row of short setae. Somite IV similar to so- tip, tip acute, tip to base concave distally to mite III, but thinner and shorter, anterior sub- convex proximally; lateral surface smooth, marginal windows present; pleura thinner ventral margin with long plumose setae, dor- and shorter than on somite III, directed lat- sal margin with short simple setae; mesial erally; dorsal surface obliquely ¯attened an- surface with dorsal decalci®ed region, de- terolaterally; lateral and posterior margins marcated ventrally by longitudinal elevated with long plumose setae, anterior margin ridge with row of short setae; with setose with short simple setae. Somite V wider than punctations ventral to decalci®ed window. somite IV, anterior submarginal windows Propodus expanded dorsally and ventrally; present; lateral margins with plumose setae; ventral expansion exceeding ventral margin pleura absent. Somite VI slightly broader of dactylus, ventral margin with long plu- than somite V, anterior submarginal windows mose setae; dorsal expansion with row of present; dorsal surface with two short trans- long plumose setae dorsally, oblique area verse rows of setae laterad of midline and on with mat of short simple setae; lateral surface posterior margin; pleura absent. smooth, mesial surface smooth, with disto- Females with uniramous, paired pleopods ventral area of few patches of long plumose on somites II±V; males without pleopods. setae. Carpus slightly produced dorsodistally; Telson of male (®g. 86F) divided into a 266 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 subcircular (``buttonlike'') proximal two- This species belongs to the ``carabus- thirds and mucronate tapering distal third; group'' of Albunea and is the sister taxon to proximal two-thirds rounded, heavily calci- the other two species in that group. The dis- ®ed, dorsally in¯ated, with deep median tinctive ``buttonlike'' morphology of the groove and long plumose setae ¯anking dis- male telson separates it from both A. carabus tal third of groove to distal end of calci®ed and A. danai. region, proximolateral angles with few short simple setae, transverse row of long plumose Albunea carabus (Linnaeus, 1758) setae present on either side of median groove Figures 87, 88 approximately one-third proximal from distal Cancer Carabus Linnaeus, 1758: 632. ± Linnae- margin of calci®ed region; distal third of tel- us, 1767: 1052. ± Houttuyn, 1769: 417. ± de son weakly calci®ed, sharply tapering distal- Villers, 1789: 156. ly with produced tip; entire telson lined with Cancer carabus: Statius MuÈller, 1775: 1126± long plumose setae on margins in distal two- 1127. ± Latreille, 1831: 56.ÐH. Milne Ed- thirds. Telson of female (®g. 86G) ovate, ta- wards, 1837a: 112.ÐH. Milne Edwards, 1840: pering slightly distally, tip slightly produced, 112. ± Bolivar, 1875: 20±21. dorsal surface evenly calci®ed; median Cancer (Pagurus) Carabus: Gmelin and LinneÂ, groove present in medial two-thirds of dorsal 1790: 2984. Cancer (Astacus) Carabus: Herbst, 1796: 67±68. surface, lined with short simple setae; prox- Albunea symnista [sic]: Ra®nesque-Schmaltz, imolateral angle with few short simple setae, 1814: 20. ± Lucas, 1849a: 27±28*. ± Lucas, margins with long simple setae. 1849b: pl. 3, ®g. 2*. ± Heller, 1863: 153 (part). DISTRIBUTION: Known from Taiwan; ± Barrois, 1888: 18±19, 75, 82, 89, 93±94. ± Queensland, Australia; New Caledonia; and Bolivar, 1892: 128 (list). ± Ferrer Aledo, 1914: Fiji, in up to 29.3 m depth. Also provision- 68. ± Miranda y Rivera, 1933a: 22. ± Miranda ally from Pitcairn Island, in 47.6±54.9 m y Rivera, 1933b: 1 (list). ± Holthuis, 1954c: 34. depth. ± Holthuis, 1956: 228. ± ICZN, 1958: 222 (not Albunea symmysta (Linnaeus, 1758)). MAXIMUM SIZE: Males: 15.4 mm cl; fe- males, 19.4 mm cl. cancer carabus: Lamarck, 1818: 224. Albunea Guerinii Lucas, 1853: 45±47, pl. 1, ®g. TYPE SPECIMENS: AM P15353 (holotype), 9. ± Bolivar, 1875: 20±21. ± Carus, 1885: 496. AM P1925 (allotype), AM P20456 (para- ± Rodriguez Femenias, 1887: 5. ± de Buen, type), NTOU (paratype), WAM 23387 (2 1887: 425. ± Bolivar, 1916: 251 (list). ± de paratypes), ZMH K±5137 (paratype). Buen, 1916: 359. TYPE LOCALITY: Off Richmond River Albunaea [sic] Guerini: Stimpson, 1858: 230 mouth, off Ballina, New South Wales, Aus- (list). tralia, 28Њ52ЈS, 153Њ34ЈE, 16 m. Albunea sp. LarrinuÂa y Azcona, 1874: 46. ETYMOLOGY: The speci®c name of this tax- ``Albunea?'' Barcelo y Combis, 1875: 63. on is given for the distinctive buttonlike mor- Albunea guerinii: Miers, 1878: 327±328. ± Ahar- phology of the telson of the males. oni, 1937: 1136. ± Aharoni, 1944: 41*. Albunea carabus: Ortmann, 1896: 224 (list). ± REMARKS: The Pitcairn specimen exists Ortmann, 1901: 1276. ± Monod, 1933: 473 only as the carapace and anterior half of the (part). ± Gordon, 1938: 186, ®g. 3b. ± Bouvier, animal with all appendages anterior to and 1940: 179±181, ®gs. 132, 133. ± Holthuis, including pereopod I. Given its poor condi- 1954c: 34. ± Piguet, 1955: 14. ± Holthuis, 1956: tion, it is only provisionally referred to this 228. ± Monod, 1956: 40±42, ®gs 10±14. ± Hol- species, although the characters of the extant thuis and Gottleib, 1958: 77, 116*. ± ICZN, structures are essentially identical. The single 1958: 222. ± Figueira, 1960: 6. ± Gauld, 1960: reference to an Easter Island albuneid (Di- 66 (list). ± Zariquiey Alvarez, 1961: 103±109, Salvo et al., 1988) may also be this taxon, as ®gs. 1±3. ± Crosnier, 1967: 341*. ± PeÂreÂs, 1967: there are several crustacean species which 456. ± Zariquiey Alvarez, 1968: 294±296, ®g. 104. ± Moncharmont, 1969: 434±439, ®gs. 1± are known from both Easter Island and Pit- 3. ± Rubio and Holthuis, 1976: 41±46*. ± Kae- cairn Island (Boyko, in prep.). Despite re- stner, 1980: 336, ®g. 13±26b. ± Riedl, 1983: peated attempts to collect albuneids on Eas- 488, pl. 179. ± Beschin and De Angeli, 1984: ter Island over a 3±week period in August 92, ®g. 2. ± Pretus, 1985±1989a: 258. ± Pretus, 1999, I found no specimens. 1985±1989b: ®g. 92. ± CoeÃlho and Calado, 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 267

1987: 42±43, table 1. ± GarcõÂa Socias and Mas- mm cl (MNHN-Hi 148); Cape Matifou, suti Jaume, 1987: 79, 91 (list). ± GarcõÂa Socias 1840±1842, M. Deshayes: 1 &, 15.5 mm cl, and Gracia, 1988: 55. ± Seridji, 1988: 1293± syntype of A. guerinii (MNHN-Hi 189); 1298, ®gs. 1±3. ± GuilleÂn Nieto, 1990: 165± Cape Matifou, 1840±1842, coll. M. Deshay- 167, ®g. 64. ± Brunet and Vicente, 1992: 168± es: 1 (, 12.4 mm cl, syntype of A. guerinii 169, ®gs. 1, 2. ± GarcõÂa Raso et al., 1992: 131, 133, 258 (list). ± Wirtz and Martins, 1993: 57± (MNHN-Hi 190); Cape Matifou, 1840±1842, 58. ± Calado, 1995: 27±30, pl. 4, ®g. a, pl. 5, coll. M. Deshayes: 2 (, 13.5±16.0 mm cl, ®g. a, pl. 6, ®gs. a±f*. ± GonzaÂlez PeÂrez, 1995: syntypes of A. guerinii (MNHN-Hi 9); Rade 165, ®g. 102. ± Giacobbe and SpanoÁ, 1996: de BoÃne, 15±20 m, 1908, coll. E. Chevraux: 719±725, pl. 1. ± Calado, 1997a: 17, 21±22.Ð 3 &, 13.9±18.1 mm cl (MNHN-Hi 1). SpanoÁ et al., 1999: 617±620. ± d'Udekem Spain: Off San Andres de Llavaneras, d'Acoz, 1999: 171. ± Boyko, 1999: 161. near Arenys de Mar, Barcelona, 4±5 m, Aug. Albunea barbara (Lucas ms) Ortmann, 1896: 224 8, 1973, coll. Tau-Tau: 1 (, 18.0 mm cl (list) (nomen nudum). (RMNH 36273), 1 (, 19.5 mm cl (AMNH Albunea Guerini: A. Milne Edwards and Bouvier, 18077 ex RMNH 36273); between San Pol 1900: 275. ± Ferrer y Galdiano, 1918: 413 (list). Albunea symmista [sic]: Holthuis, 1954c: 34 (not de Mar and San Andres de Llavaneres, near Albunea symmysta (Linnaeus, 1758)). Arenys de Mar, Barcelona, 3±6 m, Nov. 22, ?Albunea sp. B Lebour, 1959: 129, ®g. 15. 1972, March 1973, Aug. 3, 1973, coll. Tau- Albunea Carabus: Seridji, 1988: 1293, 1298. Tau: 1 (, 20.0 mm cl (RMNH 36274), 3 (, Albunea aff. carabus: Fransen, 1991: 48, 77. 19.1±21.0 mm cl (USNM 258373 ex RMNH not Hippa caerulea Risso, 1816: 50±51. ± Risso, 36274); between San Pol de Mar and San 1827: 36±37 (ϭ Gnathia sp. cf. phallonajopsis Andres de Llavaneres, near Arenys de Mar, Monod, 1925). Barcelona, 3±6 m, Aug. 27, 1974, coll. Tau- not Hippa coerulea [sic]: Risso, 1844: 94. ± Tau: 1 (, 20.3 mm cl (USNM 260867); Ar- Hope, 1851: 12 (ϭ Gnathia sp. cf. phallona- enys de Mar, Barcelona, 3±7 m, June 11, jopsis Monod, 1925). not Albunea guerinii: Stebbing, 1914: 281 (ϭ Al- 1971, coll. M. Rubio: 1 (, 18.6 mm cl bunea gibbesii Stimpson, 1859). (RMNH 28713); Canet de Mar, Barcelona, not Albunea carabus: Balss, 1916a: 37*. ± Mo- 5±7 m, June 23, 1972, coll. Tau-Tau: 1 (, nod, 1933: 473 (part) (ϭ Albunea elegans A. 18.4 mm cl, 1 &, 23.5 mm cl (RMNH Milne Edwards and Bouvier, 1898). 28911); Canet de Mar, Barcelona, 5 m, Nov. not Albunea guerinii: Stebbing, 1917: 26 (ϭ Al- 1972, coll. M. Rubio: 2 (, 18.1±19.2 mm cl bunea sp. indet.). (MNHN-Hi 85). not Albunea carabus: Chace, 1966: 635 (ϭ Al- Portugal: Funchal Bay, Madeira, 10±15 bunea gibbesii Stimpson, 1859). m, March 31, 1958, coll. A. Figuera: 1 &, not Albunea carabus: Holthuis, 1977: 61±62 (ϭ 17.7 mm cl (ZMUC 2712); Sta. 25, 36Њ59ЈN, Gnathia sp. cf. phallonajopsis Monod, 1925). 25Њ03ЈW, 10±18 m, May 28, 1981, coll. MATERIAL EXAMINED: Lebanon: Vicinity CANCAP V: 1 oviger, 10.0 mm cl, 1 juve- of Beirut, coll. M. E. Ericksen: 2 &, 16.3± nile, 4.0 mm cl (RMNH 36282). 16.6 mm cl (USNM 260973). Cape Verde: Sta. 7.068, west-southwest Israel: Nahr-Rubin, coll. unknown: 1 &, of IlheÂu Calheta do Visto, west of Boa Vista, 14.7 mm cl (BMNH 1920.12.16.3); Haifa 16Њ11ЈN, 22Њ59ЈW, 40 m, Aug. 27, 1986, Bay, June 18, 1987, coll. unknown: 1 &, 16.5 coll. CANCAP: 1 &, 5.1 mm cl (RMNH mm cl (ZMTAU NS 24665). 38605). Tunisia: Public beach, Raouad, Jan. 30, Ivory Coast: Sta. 17, Gulf of GuineÂe, 27 1973, coll. J. Moncef and R. B. Manning: 1 m, June 21, 1956, coll. ``Calypso'': 8 (, 9.4± (, 14.0 mm cl (USNM 256595). 10.4 mm cl, 6 &, 8.3±10.5 mm cl (MNHN- Algeria: Between Lebon and Hussein, Hi 111). 36Њ44ЈN, 03Њ05ЈE, 2±7 m, June±July 1941, Ghana: Trawl 1, Agassil, coll. R. Bassin- coll. H. Nouvel: 1 (, 4.4 mm cl (RMNH dale: 1 &, 11.4 mm cl (BMNH 1957.12.4.1). 34726); Oran, July 1881, coll. M. Deshayes Togo:CoÃtes du Togo, 06Њ07ЈN, 01Њ34ЈE, on ``Travailleur'': 2 (, 12.1±13.3 mm cl, 1 26 m, Oct. 5, 1963, coll. A. Crosnier: 1 (, &, 18.6 mm cl (MNHN-Hi 10); Gulf of 11.8 mm cl (MNHN-Hi 5). Dahomey: Oran, June 6, 1900, coll. P. Pallary: 1 &, 20.7 06Њ16ЈN, 02Њ26ЈE, 22 M, Oct. 9, 1963, coll. 268 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

A. Crosnier: 1 unsexable, unmeasurable gin of carapace, sinuous, strongly crenulate, specimen (MNHN-Hi 2); 06Њ14ЈN, 02Њ26ЈE, divided into medial fragment and curved, 30 m, Oct. 9, 1963, coll. A. Crosnier: 1 &, posteriorly displaced lateral elements. Me- 4.9 mm cl (MNHN-Hi 3); 06Њ09Ј20ЉN, sogastric region smooth; CG2 absent; CG3 01Њ52ЈE, 35 m, Oct. 4, 1963, coll. A. Cros- broken into ®ve to eight short elements be- nier: 1 (, 11.2 mm cl (MNHN-Hi 4); tween posterior lateral elements of CG1; 06Њ14Ј30ЉN, 01Њ48ЈE, 15 m, Oct. 4, 1963, CG4 with two short and two long medial el- coll. A. Crosnier: 1 juvenile, 3.3 mm cl ements between longer supralateral elements (MNHN-Hi 6). of CG4. Hepatic region smooth, with oblique No Data:4(, 13.0±16.2 mm cl (MNHN- setose groove at median of lateral margin. Hi 90); 3 (, 14.8±15.9 mm cl (MNHN-Hi Epibranchial region generally triangular, 91). smooth; posterolateral margin without rows DIAGNOSIS: Carapace wider than long, cov- of setae. Metagastric region smooth; CG5 ered with lightly setose crenulate grooves. present as two curved, oblique elements. Anterior margin with 9±14 spines on either CG6 strongly crenulate, strongly anteriorly side of ocular sinus. Setal ®eld with narrow concave medially, and sloping out to anteri- lateral elements and straight anterior margin. orly convex lateral thirds. CG7 oblique, not CG1 with separate posterior lateral elements; reaching lateral margins of median segment CG4 with two short and two long medial el- of CG6. Cardiac region smooth; CG8 present ements between longer supralateral elements as two short medial elements. CG9 present of CG4; CG5 present as two curved, oblique as two short elements with wide gap at mid- elements; CG6 and CG7 separate; CG8 bro- line. CG10 present as two long elements. ken; CG11 present. Rostrum present, not CG11 present as two short elements. Post- reaching posterior margin of ocular plate. CG11 element absent. Branchial region with Ocular plate triangular. Distal peduncular numerous short, transverse rows of setae. segments dorsoventrally ¯attened and trian- Posterior margin deeply and evenly convex, gular in shape, tapering at tip, approximated with submarginal groove reaching halfway along mesial margins at base, lateral margins up margin of posterior concavity. Branchios- broadly convex, mesial margins straight. tegite with short anterior submarginal spine; Cornea at tip. Dactylus of pereopod II with anterior region with scattered short, trans- heel produced, subquadrate and rounded. verse lines ventral to linea anomurica; with Dactylus of pereopod III with heel produced, many short rows of setae and sparsely cov- subquadrate and rounded. Dactylus of pereo- ered with long plumose setae ventrally; pos- pod IV sinuous from base to tip, with low terior region membranous, with numerous ir- rounded heel and shallow indent. Telson of regular fragments and sparsely covered with male broadly triangular, as long as wide, with long plumose setae. broadly rounded tip; thickly calci®ed in Ocular plate (®g. 87B) triangular with proximal third and medial strip, in¯ated dor- deep median indentation; median peduncular sally; distolateral two-thirds decalci®ed, me- segments present as small ovate calci®ed ar- dian and transverse rows of thin setae. Telson eas lateral to ocular plate. Distal peduncular of female ¯attened, ovate, and evenly calci- segments elongate, subtriangular, with con- ®ed with slightly truncate tip. vex lateral and straight mesial margins, cor- DESCRIPTION: Carapace (®g. 87A) wider nea covering lateral three-fourths of distal than long. Anterior margin slightly concave tip; lateral margins with notch just proximal on either side of ocular sinus, becoming con- to cornea; mesial margins approximated vex laterally, with 9±14 large spines (n ϭ 6) along length; mesial and lateral margins with along length. Rostrum as small acute tooth, long plumose setae; tuft of plumose setae at not reaching proximal margin of ocular plate. proximolateral ventral angles and ventrome- Ocular sinus smoothly concave, with few dial row of plumose setae extending from small spinules. Frontal region smooth; setal tuft to base of cornea. ®eld narrow anteriorly and posteriorly; pos- Antennule (®g. 87C) with segment III nar- terior lateral elements reduced to narrow row proximally, expanding distally to twice bands of setae. CG1 parallel to anterior mar- proximal width; with plumose setae on dor- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 269

Fig. 87. Albunea carabus (Linnaeus, 1758): A, (, 21.0 mm cl, USNM 258373; B±J, (, 19.1 mm cl, USNM 258373. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular pedun- cles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 2.2 mm (B), 3.3 mm (E, F), 4.4 mm (I), 6.7 mm (C, D, G, H, J), and 8.5 mm (A). sal and ventral margins and sparsely scat- dorsal and ventral margins and scattered on tered on lateral surface; dorsal exopodal ¯a- ventrolateral third of surface. Segment I wid- gellum with 86±108 articles (n ϭ 6), long er than long, unarmed; dorsal third of lateral plumose setae on dorsal and ventral margins; surface rugose with long plumose setae; long ventral endopodal ¯agellum with three to plumose setae on dorsal and ventral margins. ®ve articles (n ϭ 6), plumose setae on dorsal Antenna (®g. 87D) with segment V ap- and ventral margins. Segment II medially in- proximately two times longer than wide, with ¯ated in dorsal view, with plumose setae on long plumose setae on dorsal and ventral 270 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 margins and scattered on lateral surface; ¯a- rounded, length equal to width, with thick gellum with seven articles (n ϭ 6), long plu- simple setae distally and on distolateral sur- mose setae on dorsal, ventral, and distal mar- face. Propodus two times wider than long, gins. Segment IV expanded distally, with slightly produced at dorsodistal angle, with long plumose setae on dorsal, ventral, and plumose setae on dorsal margin and patch of distal margins, and row of setae on dorsolat- long simple setae on lateral surface and ven- eral surface. Segment III with long plumose trolateral angle. Carpus not produced dorso- setae on ventral margin and in short row on distally, approximately two times longer than surface. Segment II short, widening distally, wide; long simple setae on dorsal margin and rugose, with plumose setae on margins and in short row on lateral surface. Merus ap- scattered on lateral surface; antennal acicle proximately three times longer than wide, long, thin, and reaching to distal margin of margins parallel; with simple and plumose segment IV, with long plumose setae on dor- setae on margins and scattered on surface. sal margin. Segment I rounded proximally, Basis-ischium incompletely fused with plu- ¯attened ventrolaterally, with long plumose mose setae on margins. Exopod one-third setae on margins and scattered on surface ru- longer than merus, ¯agellum with one elon- gae behind spine; lateral surface with acute gate article, longer than carpus. spine dorsodistally (occasionally two spines), Maxilliped III (®g. 87J) dactylus with with low semicircular dorsolateral lobe ven- rounded tip; long plumose setae on margins trodistal to spine; segment with ventromesial and lateral surface. Propodus dorsodistally antennal gland pore. in¯ated, with longitudinal median row of Mandible (®g. 87E) incisor process with plumose setae on lateral surface; margins two teeth; cutting edge with one tooth. Palp with plumose setae. Carpus produced onto three-segmented, with plumose setae on mar- propodus almost one-half length of propo- gins and long, thick, simple setae arising dus; lateral surface with two rows of plumose from bend in second segment and on distal setae; plumose setae on margins. Merus in- margin of terminal segment. ¯ated, unarmed, with plumose setae on mar- Maxillule (®g. 87F) distal endite proxi- gins and scattered on lateral surface. Basis- mally narrow, widening to in¯ated distal end, ischium incompletely fused, without crista with thick simple setae on distal margin and dentata. Exopod two-segmented: proximal thin simple setae on dorsal margin. Proximal segment small; distal segment styliform, ta- endite with thick simple setae on distal mar- pering, approximately one-third length of gin. Endopodal external lobe truncate distally merus; with plumose setae on margins; with- and curled under; internal lobe reduced with out ¯agellum. three thick setae at distolateral margin. Pereopod I (®g. 88A) dactylus curved and Maxilla (®g. 87G) exopod evenly rounded, tapering; lateral and mesial surfaces smooth; with plumose setae along distal margin. Sca- dorsal margin with few rugae proximally and phognathite bluntly angled on posterior lobe, long plumose and short simple setae; ventral with plumose setae. margin with short simple setae. Propodal lat- Maxilliped I (®g. 87H) epipod with plu- eral surface with numerous short, transverse mose setae on margins, distolateral surface, rows of setose rugae; dorsal margin un- and mesial surface. Endite tapered distally armed; ventral margin distally produced into and subequal to ®rst segment of exopod. Ex- acute spine; cutting edge lacking teeth, lined opod with two segments: proximal segment with long plumose setae; dorsal margin with narrow, margins parallel with plumose setae; long plumose setae, ventral margin with distal segment spatulate, longer than wide, short simple setae. Carpus with dorsodistal broadest medially, margins and mesioventral angle produced into strong corneous-tipped surface with long plumose setae. Endopod spine; dorsal margin with short transverse ¯attened and elongate, reaching two-thirds of grooves behind spine; dorsal and distal mar- distance to distal end of proximal exopodal gins with short plumose setae; lateral surface segment; plumose setae on margins and me- with small distal rugose area, with few trans- dian of lateral surface. verse setose ridges on distal half of surface; Maxilliped II (®g. 87I) dactylus evenly mesial surface smooth, with medial trans- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 271

Fig. 88. Albunea carabus (Linnaeus, 1758): A, B, F, (, 19.1 mm cl, USNM 258373; C, D, (, 21.0 mm cl, USNM 258373; E, (, 19.1 mm cl, USNM 258373; G, &, 23.5 mm cl, RMNH 28911. A. Left pereopod I, lateral view. B. Right pereopod II, lateral view. C. Left pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 3.3 mm (F, G), 6.7 mm (A, B), and 8.4 mm (C±E). verse and subdorsal row of setae, margins and not produced, heel to tip with narrow, with long plumose setae. Merus unarmed; acute indent, tip acute, tip to base broadly lateral surface with scattered transverse rows convex; lateral surface smooth, with several of long plumose setae, margins with long small tufts of short setae in roughly straight plumose setae; mesial side with few scattered line across medioproximal surface, several setae; fully calci®ed. Basis-ischium incom- widely spaced submarginal tufts of short se- pletely fused, unarmed. Coxa unarmed. tae dorsodistally; mesial surface smooth, Pereopod II (®g. 88B) dactylus smooth; ventral margin with long plumose setae, dor- base to heel slightly concave, heel rounded sal margin with short simple setae, with 272 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 patch of long plumose setae at base. Propo- large decalci®ed window covering nearly dal dorsal surface smooth, ventral margin in- half of lateral surface medially; dorsal and ¯ated and rounded; oblique row of long plu- ventral margins unarmed, with long plumose mose setae on distal margin of lateral sur- setae; distolateral margin with long plumose face; distal and ventral margins with long setae; mesial surface smooth. Basis-ischium plumose setae; dorsolateral surface as nar- incompletely fused and unarmed. Coxa un- row, oblique, ¯attened shelf, with short setae armed. Female with large gonopore on an- on dorsal margin and long plumose setae on terior mesial margin of coxa, surrounded ventral margin; mesial surface with elevated, with short plumose setae; male without pore. curved setose ridge from ventral junction Pereopod IV (®g. 88D) dactylus with base with dactylus almost to ventral proximal to tip convex proximally, concave distally, junction with carpus. Carpus strongly pro- with shallow indent, tip acute, tip to base duced and rounded dorsodistally, dorsal mar- convex; lateral surface smooth, ventral mar- gin smooth proximally, serrate distally; lat- gin with long plumose setae, dorsal margin eral surface smooth, with small setose mat at with short simple setae; mesial surface with tip of produced area and irregular, broken dorsal decalci®ed region, demarcated ven- row of rugae and submarginal elevated ridge trally by longitudinal elevated ridge with row ventrally, rugae and ridge with long plumose of short setae; with setose punctations ventral setae; margins with long plumose setae; me- to decalci®ed window. Propodus expanded sial surface smooth with row of long plu- dorsally and ventrally; ventral expansion mose setae distally, ventrally, and subdorsal- reaching ventral margin of dactylus, margin ly. Merus with large median decalci®ed win- with long plumose setae; dorsal expansion dow covering nearly all of lateral surface, with row of long plumose setae dorsally, with few scattered long plumose setae on oblique area smooth; lateral and mesial sur- surface and margins; mesial surface nearly faces smooth. Carpus slightly produced dor- smooth with two long rows of setae. Basis- sodistally; ventral three-fourths of lateral sur- ischium incompletely fused and unarmed. face and mesial surface smooth, with few Coxa unarmed. scattered setae, dorsodistal quarter of lateral Pereopod III (®g. 88C) dactylus with base surface with mat of short setae; dorsal mar- to heel slightly concave, heel rounded and gin with short simple and long plumose se- not produced, heel to tip with broadly con- tae; ventral margin with short simple setae; cave indent, tip acute, tip to base smoothly mesial surface decalci®ed medially. Merus convex; lateral surface smooth, with several lateral surface with scattered short transverse small tufts of short setae in roughly straight rows of setae, dorsal and ventrodistal mar- line across medioproximal surface, dorsodis- gins with long plumose setae; proximoven- tal margin with tufts of short setae; ventral tral half of mesial surface with large decal- margin with long plumose setae, dorsal mar- ci®ed window. Basis-ischium incompletely gin with short simple and plumose setae; me- fused and unarmed. Coxa unarmed. sial surface smooth with plumose setae prox- Abdomen (®g. 88E) with somite I length imally at junction with propodus. Propodus and width subequal, widest posteriorly; dor- not in¯ated dorsoventrally; lateral surface sal surface with anterior margin straight; pos- smooth, with long plumose setae in oblique terior margin curved, with elevated submar- row, simple setae on dorsal margin; dorso- ginal row of short setae; small transverse de- lateral surface narrow, oblique, ¯attened; me- calci®ed windows laterad of segment medi- sial surface smooth. Carpus produced dor- an. Somite II dorsal surface with submarginal sodistally, exceeding proximal margin of transverse ridge anteriorly; with small trans- propodus by one-half length of propodus; verse decalci®ed windows laterad of segment dorsolateral margin unarmed; lateral surface median just anterior to submarginal ridge; slightly rugose dorsodistally, with mat of pleura expanded and directed anterolaterally; short setae and two interrupted rows of setae lateral margins angled, anterior and lateral ventrally; mesial surface smooth, with long margins with long plumose setae, posterior plumose setae on distal margin and in row margin with short setae; posteromesial angle on distal half of surface. Merus smooth, with with mat of short simple setae. Somite III 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 273 similar to somite II, but narrower, shorter; syntypes and possess the correct data. It may pleura thinner and shorter than on somite II, be that MNHN-Hi 90 and MNHN-Hi 91 are directed posterolaterally but with distal tips also syntypes of A. guerinii, but they have swept forward, with setae as in somite II; no data associated with them that would con- anterolateral angle subacute; dorsal surface ®rm this. obliquely ¯attened anterolaterally. Somite IV TYPE LOCALITIES: Cancer carabus: ``M. similar to somite III, but thinner and shorter; Mediterraneo'' (Linnaeus, 1758). As pointed dorsal surface with few short setae antero- out by Rubio and Holthuis (1976), the col- laterally; pleura thinner and shorter than on lector of the type(s), Erik Brander, was sta- somite III, directed posterolaterally with dis- tioned in Algeria, and this is probably where tal tips slightly swept forward; dorsal surface the type(s) came from, although it cannot be obliquely ¯attened anterolaterally; margins considered the type locality in the absence of with long plumose setae. Somite V wider a primary type specimen. Albunea guerinii: than somite IV; lateral margins with plumose Algeria. setae; pleura absent. Somite VI slightly REMARKS: At least part of the dif®culty broader than somite V; dorsal surface and early authors had in identifying this taxon as distal margin with short transverse rows of an Albunea was caused by Linnaeus' (1758) setae laterad of midline; pleura absent. placement of the species within Cancer. Females with uniramous, paired pleopods Cancer Carabus (Linnaean no. 48) was sep- on somites II±V; males without pleopods. arated by eight intervening taxa from Cancer Telson of male (®g. 88F) broadly triangular, Symmysta (Linnaean no. 39), the only other as long as wide, with broadly rounded tip; albuneid described by Linnaeus (1758). thickly calci®ed in proximal third and medial These eight intermediate taxa were an as- strip, in¯ated dorsally; distolateral two-thirds sortment of anomurans, cray®sh, and shrimp decalci®ed; median longitudinal groove ex- which, by themselves, offered no clue as to tending one-half length, rows of long simple the relationships of carabus and symmysta. setae demarcating juncture of calci®ed and By 1767, Linnaeus had moved the two taxa decalci®ed regions and in tuft at distal end; closer together, to no. 68 and no. 70 on his proximolateral angles without setae; margins list, and separated them by a galatheid. The with long simple setae. Telson of female (®g. ``rostrum dentibus 2 parallelis mobilibus de- 88G) ¯attened, ovate, and evenly calci®ed with slightly truncate tip; median groove pressis'' described by Linnaeus (1758) for similar to male, row of short simple setae on this species is actually the ocular peduncles either side of median groove from midpoint rather than the rostrum. This was probably of telson continuing distally to three-fourths the key morphological feature that caused length of telson and in short transverse sub- confusion as to what sort of crustacean Lin- marginal lateral rows approximately one- naeus was referring to (Miers, 1878). Lin- fourth distal from proximal margin; proxi- naeus (1758) cited this species as ``Testa molateral angle without setae, margins with magnitudine extimi pollicis'' (as long as long simple setae. the last joint of the thumb), and this has DISTRIBUTION: From Lebanon westward been extrapolated to a carapace length of ap- through the Mediterranean and from the proximately 17 mm (Rubio and Holthuis, Azores and Madeira southward to Dahomey, 1976), although larger specimens are known. in up to 40 m depth. Calado (1995) erroneously cited BMNH MAXIMUM SIZE: Males: 21.0 mm cl; fe- 1920.12.16.3 as the ``holoÂtipo.'' No Linnaean males: 23.5 mm cl. material of this species exists, no neotype has TYPE SPECIMENS: The type(s) of Cancer ever been designated, and no neotype is carabus are lost. MNHN-Hi 189 (syntype of needed, as this species is the only albuneid A. guerinii), MNHN-Hi 190 (syntype of A. in the Mediterranean and is only distantly re- guerinii), MNHN-Hi 9 (2 syntypes of A. lated to the other albuneid species in the east guerinii). Lucas (1849a, 1853) cited 16 total Atlantic. The species has been called specimens (ϭ syntypes) of A. guerinii. Only ``Leeuw-Krab'' (Houttuyn, 1769), ``Kahn four specimens in the MNHN are labeled as Krabbe'' (Statius MuÈller, 1775), and ``LoÈw- 274 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 enkrebs'' (Herbst, 1796), but no general poorly known, this seems a premature con- common name has been adopted. clusion. The west African larvae reported by Lucas (1853), in describing A. guerinii, Lebour (1959) as ``Albunea sp. B'' resemble was the ®rst to recognize and discuss the sex- the early (stage II or III) zoeae of A. carabus ual dimorphism in telson shape within the in the shape of the posterior telson margin. genus. Zariquiey Alvarez's (1961: ®g. 1) il- The coloration of adults is: carapace dark lustration of this species is one of the most purple with brownish areas, dorsal regions of accurate drawings published for any albuneid pereopods and abdomen purple, antennular before 1999 (®g. 1C, herein), and his illus- ¯agellae ringed with purple and yellowish trations of the male and female telsons are brown (Rubio and Holthuis, 1976). another rare example of recognition of the This species typi®es the ``carabus-group'' sexual dimorphism present in this genus. of Albunea, which also includes A. danai and Brunet and Vicente (1992) incorrectly cit- A. bulla. It is the sister species to A. danai. ed Monod as the ®rst author to unequivocally assign this taxon to Albunea; that assignation Albunea asymmetrica (P. MuÈller, 1979) was ®rst indicated by Ortmann (1896). Ort- Figure 89 mann (1896) also introduced the manuscript name ``Albunea barbara Lucas'' in synony- Mioranina asymmetrica P. M uÈller, 1979: 278±279, my with A. carabus. That name is both a pls. 9, 10*. Albunea asymmetrica:P.MuÈller, 1984: 62, pl. 28, nomen nudum and an unavailable name. The ®gs. 4±6*. specimen cited from ANSP by Ortmann (1896) and identi®ed as A. barbara Lucas is MATERIAL EXAMINED: Hungary: MGy±2± not extant (Boyko, personal obs.). 1, Upper Badenian, Budapest, GyakorloÂuÂt, Holthuis's (1977) placement of Hippa ca- coll. P. MuÈller: 1 carapace fragment, 10.5 ϫ erulea Risso in synonymy with A. carabus is 13.6 mm (l ϫ w), holotype (plaster cast of incorrect, as Hippa caerulea is actually a HNHM); MGy±2±2, Upper Badenian, Bu- gnathiid isopod (see appendix 2). dapest, GyakorloÂuÂt, coll. P. MuÈller: 1 cara- Although there are repeated statements in pace fragment, 13.8 ϫ 12.2 mm (l ϫ w), the literature that this species is ``muy raro'' paratype (plaster cast of HNHM; pl. 9, ®g. 3 (Zariquiey Alvarez, 1968; Brunet and Vi- in P. MuÈller, 1979); MGy±2±2, Upper Bad- cente, 1992), it is likely more common in its enian, Budapest, GyakorloÂuÂt, coll. P. MuÈller: preferred sand bottom habitats than current 1? merus, 13.0 ϫ 8.0 mm (l ϫ w), paratype data suggest. The apparent rarity of this spe- (plaster cast of HNHM; pl. 9, ®g. 2 in P. cies in the eastern basin of the Mediterranean MuÈller, 1979); MGy±2±2, Upper Badenian, is likely due to limited collecting efforts rath- Budapest, GyakorloÂuÂt, coll. P. MuÈller: 1 car- er than to unspeci®ed ``ecological factors'' apace fragment, 12.5 ϫ 9.5 mm (l ϫ w), (see SpanoÁ et al., 1999). The species is also paratype (plaster cast of HNHM; pl. 10, ®g. probably more widely distributed in the east- 2inP.MuÈller, 1979); MGy±2, Upper Bad- ern basin than current data indicate. In fact, enian, Budapest, GyakorloÂuÂt, coll. P. MuÈller: A. carabus is occasionally washed ashore in 1 carapace fragment, 20.2 ϫ 10.1 mm (l ϫ large numbers, possibly due to mortality w), paratype (latex cast of HNHM; pl. 10, from freshwater runoff (Piguet, 1955). This ®g.1inP.MuÈller, 1979); MGy±2, Upper is apparently the only species of albuneid in Badenian, Budapest, GyakorloÂuÂt, coll. P. the Azores, but is only known from a few MuÈller: 6 fragments (HNHM); MR8±16, Up- specimens from that location (Figueira, per Badenian, Budapest-RaÂkos, Layer 8, coll. 1960). P. M uÈller: 2 fragments, 9.5 ϫ 9.8 mm, 7.5 ϫ Seridji (1988) described zoeal stages I, II, 8.0 mm (l ϫ w) (HNHM). and III from plankton collected off Algeria. DIAGNOSIS: Carapace as long as wide, cov- Seridji (1988) speculated that A. carabus ered with crenulate grooves. Anterior margin might have the same number of zoeal stages with 10 spines on either side of ocular sinus. (®ve) as A. symmysta, but as these two spe- Setal ®eld with thin lateral elements and cies are not particularly closely related and straight anterior margin. CG1 with separate larval development in the genus is overall posterior lateral elements; CG4 with two 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 275

Fig. 89. Albunea asymmetrica (MuÈller, 1979): A. Anterolateral left quarter of carapace, 10.5 ϫ 13.6 mm (l ϫ w), HNHM MGy-2±1, holotype. B. Posterolateral right quarter of carapace, 13.8 ϫ 12.2 mm (l ϫ w), HNHM, paratype. C. Right half of carapace, 20.2 ϫ 10.1 mm (l ϫ w), HNHM MGy-2, paratype. D. Anterior section of right branchiostegite, 12.5 ϫ 9.5 mm (l ϫ w), HNHM. Scale ϭ 6.7 mm (B±D) and 7.2 mm (A). long medial elements between longer supra- triangular, smooth. Metagastric region lateral elements of CG4; CG5 unknown; smooth; CG5 presence unknown. CG6 CG6 and CG7 fused; CG8 unknown; CG11 strongly crenulate, strongly anteriorly con- unknown. Rostrum present. cave medially and sloping out to anteriorly DESCRIPTION: Carapace (®g. 89A±C) ap- convex lateral thirds. CG7 oblique, not proximately as wide as long. Anterior margin reaching lateral margins of median segment slightly concave on either side of ocular si- of CG6. Cardiac region unknown; CG8 un- nus, becoming convex laterally with 10 large known. CG9 unknown. CG10 unknown. spines (n ϭ 1) along length. Rostrum as CG11 unknown. Branchial region with nu- small acute tooth. Ocular sinus smoothly merous short, transverse rows of setae. Bran- concave and unarmed. Frontal region chiostegite (®g. 89D) with short anterior sub- smooth; setal ®eld narrow anteriorly and marginal spine; anterior region with scattered widening posteriorly; posterior lateral ele- rugae. ments reduced to narrow bands of setae. CG1 Pereopod I subchelate. Dactylus unknown. parallel to anterior margin of carapace, sin- Propodal lateral surface with numerous short, uous, strongly crenulate, divided into medial transverse rows of setose rugae; dorsal mar- fragment and curved, posteriorly displaced gin unarmed. lateral elements. Mesogastric region smooth; DISTRIBUTION: Known only from the Up- CG2 presence unknown; CG3 present but per Badenian formation, Budapest, Hungary. composition unknown; CG4 with two long TYPE SPECIMENS: HNHM MGy±2±1 (ho- medial elements between longer supralateral lotype), HNHM MGy±2±2 (8 paratypes). elements of CG4. Hepatic region smooth, TYPE LOCALITY: MGy±2±1, Upper Bad- with oblique setose groove at median of lat- enian, Budapest, GyakorloÂuÂt, Hungary. eral margin. Epibranchial region generally REMARKS: The species was ®rst described 276 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 as the type of the ``raninid'' genus Mioranina m), Sept. 9, 1959, coll. Pele Expedition: 1 &, P. M uÈller, 1979, illustrating once again the 13.0 mm cl, paratype (BPBM S6777); Kan- convergent morphology between raninds and oehe Bay, Oahu, 1924, coll. unknown: 1 (, albuneids. None of the material of this taxon 4.0 mm cl (BPBM S7806). is well preserved, especially when compared DIAGNOSIS: Carapace slightly longer than to other known albuneid fossils (e.g., Bes- wide, covered with lightly setose grooves. chin and De Angeli, 1984). This material is Anterior margin with eight or nine spines. quite similar to A. carabus in its carapace Setal ®eld with narrow lateral elements and grooves, shortness of branchiostegite spine, slightly concave anterior margin; posterior and proportions of length and width. The lateral elements not extending to posterior number and size of the anterior carapace lateral elements of CG1. CG1 with separate margin spines of A. asymmetrica are not ap- posterior lateral elements; CG4 with two to preciably different from A. carabus, contrary four short medial elements; CG5 divided into to the statements of P. MuÈller (1984). The two lateral elements, not nearly reaching relatively recent age (15 mybp, Tertiary: margins of CG6; CG6 and CG7 separate, but Miocene) of this fossil material also suggests almost approximate; CG8 with one or two that it may be conspeci®c with an extant tax- posteriorly displaced median elements sepa- on. Additional better preserved material is rated from lateral elements; CG11 absent. needed to determine if A. asymmetrica and Rostrum present, not reaching proximal mar- A. carabus are synonymous or closely related gin of ocular plate. Ocular plate subquadrate. species. Distal peduncular segments dorsoventrally ¯attened and elongate, pointed at tip, ap- Albunea danai Boyko, 1999 proximate along mesial margins; lateral mar- Figures 90, 91 gins convex; mesial margins straight proxi- mally, convex distally. Cornea at lateral mar- Albunea danai Boyko, 1999: 145 (list), 155±161, ®gs. 5, 6*. gin of tip. Antennule with 87±92 ¯agellar ex- opodal and three or four endopodal articles. MATERIAL EXAMINED: USA: Hawaii: Hal- Antenna with seven ¯agellar articles; acute onu Blow Hole dive site, south shore, Oahu, spine on dorsolateral surface of peduncle 12.2±13.7 m, April 4, 1997, coll. R. Holcom: segment I. Dactyli of pereopods II and III 1 (, 16.7 mm cl, holotype (QM W23105); with heels low and smoothly rounded. Coxa Kailua, Oahu, March 1938, coll. unknown: 1 of pereopod III of males with small pore. &, 16.8 mm cl, allotype (BPBM S11782); off Telson of male triangular, dorsoventrally ¯at- Eaa Beach, near Pearl Harbour, 15 fms (ϭ tened laterally and distally, in¯ated medially. 27.4 m), July 5, 1964, coll. B. R. Wilson on Telson of female ¯attened, rounded at tip. ``Pele'': 1 &, 10.8 mm cl, paratype (WAM DESCRIPTION: Carapace (®g. 90A) slightly 10422); off Waikiki, Oahu, 20 ft (ϭ 6.1 m), wider than long. Anterior margin slightly May 30, 1948, coll. Allen and V. Smith: 1 concave on either side of ocular sinus, be- &, 10.8 mm cl, paratype (AMNH 17716 ex coming convex laterally with eight or nine BPBM S5348); off Sand Island, Oahu, 16± large spines along length. Rostrum as small 25 fms (ϭ 29.3±45.7 m), July 17, 1959, coll. acute tooth, extending only one-half of the Pele Expedition: 1 (, 10.4 mm cl, paratype distance between distal margin of the ocular (AMNH 17717 ex BPBM S6776); off Hon- sinus and ocular plate. Ocular sinus smoothly olulu, 15±22 fms (ϭ 27.4±40.2 m), Feb.± concave and unarmed. Frontal region March 1962, coll. T. Richert: 1 (, 16.4 mm smooth; setal ®eld broad posteriorly, narrow- cl, 1 &, 15.5 mm cl, paratypes (WAM ing anteriorly, with narrow anterior lateral el- 23390); Waikiki, Oahu, 75 ft (ϭ 22.7 m), ements and slightly concave anterior margin; May 23, 1948, coll. Smith and Allen: 1 (, posterior lateral elements thin and not reach- 11.6 mm cl, paratype (BPBM S5343); Ka- ing to posterior lateral elements of CG1. hana Bay, Oahu, 25±30 fms (ϭ 45.7±54.9 CG1 parallel to anterior margin of carapace, m), July 25, 1959, coll. Pele Expedition: 1 faintly sinuous, strongly crenulate, divided (, 8.6 mm cl, paratype (BPBM S6775); Di- into medial fragment and curved, posteriorly amond Head, Oahu, 25±45 fms (ϭ 45.7±82.3 displaced lateral elements. Mesogastric re- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 277

Fig. 90. Albunea danai Boyko, 1999: A, (, 16.7 mm cl, QM W23105, holotype; B±F, H±J, oviger, 10.8 mm cl, WAM 10422, paratype; G, ( 11.6 mm cl, BPBM S5343, paratype. A. Carapace, bran- chiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Right antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 1.2 mm (C, D, G, H, J), 1.6 mm (B, E, F), 2.2 mm (I), and 6.7 mm (A). gion smooth; CG2 absent; CG3 broken into ements spaced approximately equally be- six short elements approximately equally tween longer lateral elements of CG4. He- spaced between posterior lateral elements of patic region smooth, with oblique setose CG1; CG4 with two to four short medial el- groove at median of lateral margin. Epibran- 278 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 chial region generally triangular, smooth; articles and plumose setae on dorsal and ven- posterolateral margin with two short rows of tral margins. Segment II medially in¯ated in setae. Metagastric region smooth; CG5 di- dorsal view, with plumose setae on dorsal vided into two short lateral elements. CG6 and ventral margins and scattered on ventro- strongly crenulate, strongly anteriorly con- lateral third of surface. Segment I wider than cave medially and sloping out to anteriorly long, unarmed; dorsal third of lateral surface convex lateral thirds. Cervical groove strong- rugose with long plumose setae; long plu- ly concave at median and sloping out to ir- mose setae on dorsal and ventral margins. regularly convex lateral thirds; median and Antenna (®g. 90D) with segment V ap- lateral thirds separated by short setae-free proximately two times longer than wide, with gap lateral to small depressions. CG7 long plumose setae on dorsal margin and oblique, almost reaching lateral margins of scattered on distal half of lateral surface; ¯a- median segment of CG6. Cardiac region gellum with seven articles, long plumose se- smooth; CG8 present as two very short la- tae on dorsal, ventral, and distal margins. teromedial elements displaced posteriorly Segment IV expanded distally, with long plu- from longer lateral elements. CG9 present as mose setae on dorsal, ventral, and distal mar- two short lateral grooves with gap at midline. gins and two rows of setae on dorsolateral CG10 present as two curved lateral frag- surface. Segment III with long plumose setae ments with gap between fragments approxi- on dorsal and ventral margin. Segment II mately equal to length of single fragment. short, widening distally, with plumose setae CG11 absent. Branchial region with numer- on margins and scattered on lateral surface; ous short, transverse rows of setae. Posterior antennal acicle long, thin and exceeding dis- margin deeply and evenly convex, with sub- tal margin of segment IV by one-fourth of marginal groove reaching approximately length of segment IV, with long plumose se- halfway up either side of posterior concavity. tae on dorsal margin. Segment I rounded Branchiostegite with short anterior submar- proximally, ¯attened ventrolaterally with ginal; anterior region with scattered, short, long plumose setae on margins; lateral sur- transverse lines ventral to linea anomurica; face with acute spine dorsally, with low with many short rows of setae and sparsely semicircular dorsolateral lobe ventrodistal to covered with long plumose setae ventrally; spine; segment with ventromesial antennal posterior region membranous, with numer- gland pore. ous irregular fragments and sparsely covered Mandible (®g. 90E) incisor process with with long plumose setae. one tooth; cutting edge with one tooth. Palp Ocular plate (®g. 90B) subquadrate with three-segmented, with plumose setae on mar- median indentation; median peduncular seg- gins and long, thick, simple setae arising ments reduced to small rounded calci®ed from bend in second segment. area on either side of ocular plate. Distal pe- Maxillule (®g. 90F) distal endite proxi- duncular segments elongate, with distally mally narrow, widening to in¯ated distal end, convex lateral margins, tapering to rounded with thick simple setae on distal margin. distal cornea located in lateral notch; mesial Proximal endite with thick simple setae on margins approximated almost all of length; distal margin. Endopodal external lobe trun- mesial and proximolateral margins of seg- cate distally and curled under; internal lobe ment with sparse row of long plumose setae; reduced, with three thick setae at distolateral tuft of plumose setae at proximolateral ven- margin. tral angle. Maxilla (®g. 90G) exopod evenly rounded, Antennule (®g. 90C) with segment III nar- with plumose setae along distal margin. Sca- row proximally, expanding distally to two phognathite bluntly angled on posterior lobe, times proximal width; with plumose setae on with plumose setae. dorsal and ventral margins and sparsely scat- Maxilliped I (®g. 90H) epipod with plu- tered on lateral surface; dorsal exopodal ¯a- mose setae on margins, distolateral surface, gellum with 87±92 articles, long plumose se- and mesial surface (epipod shown curled). tae on dorsal and ventral margins; ventral en- Endite tapered distally and subequal to ®rst dopodal ¯agellum short with three or four segment of exopod. Exopod with two seg- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 279 ments; proximal segment narrow, margins otherwise unarmed; dorsal and distal margins parallel, with plumose setae; distal segment with long plumose setae; lateral surface with spatulate, approximately as long as wide, small distal rugose area, with few transverse broadest medially, margins and mesioventral setose ridges on distal half of surface; mesial surface with long plumose setae. Endopod surface smooth, with few median rows of se- ¯attened and elongate, reaching to distal end tae, margins with long plumose setae. Merus of proximal exopodal segment; plumose se- unarmed; lateral surface with scattered trans- tae on margins and median of lateral surface. verse rows of long plumose setae, margins Maxilliped II (®g. 90I) dactylus evenly with long plumose setae; mesial side with rounded, length equal to width, with thick few short rows of setae; proximal quarter of simple setae distally and on distolateral sur- mesial surface with decalci®ed window. Ba- face. Propodus two times wider than long, sis-ischium incompletely fused, unarmed. slightly produced at dorsodistal angle, with Coxa unarmed. plumose setae on dorsal margin and long Pereopod II (®g. 91B) dactylus smooth; simple setae on dorsodistal margin. Carpus base to heel straight, heel smoothly rounded, not produced dorsodistally, approximately heel to tip with wide, acute indent, tip acute, two times longer than wide; long simple se- tip to base broadly convex distally and con- tae on dorsal and distal margins. Merus ap- cave proximally; lateral surface smooth, with proximately three times longer than wide, several small tufts of short setae in generally margins parallel; with simple setae on ven- straight line across medioproximal surface, trolateral margin and plumose setae on dor- several widely spaced submarginal tufts of solateral margin. Basis-ischium incompletely short setae dorsodistally; mesial surface fused with plumose setae on margins. Exo- smooth, ventral margin with long plumose pod one-third longer than merus, with ¯a- setae, dorsal margin with short simple setae, gellum one-segmented. Maxilliped III (®g. patch of long plumose setae at base. Propo- 90J) dactylus with rounded tip; long plumose dal dorsal surface smooth, with ventral mar- setae on margins and lateral surface. Propo- gin in¯ated and rounded; oblique row of long dus with longitudinal median row of plumose plumose setae on distal margin of lateral sur- setae on lateral surface; margins with plu- face; distal and ventral margin with long plu- mose setae. Carpus slightly produced onto mose setae; dorsolateral surface as narrow, propodus; lateral surface with row of plu- mose setae ventromedially; plumose setae on oblique, ¯attened shelf, with short setae on margins. Merus unarmed, with plumose setae dorsal margin and long plumose setae on on margins and scattered on lateral surface. ventral margin; mesial surface with elevated, Basis-ischium incompletely fused, with weak curved, setose ridge from ventral junction crista dentata of approximately two teeth. with dactylus almost to ventral proximal Exopod two-segmented: proximal segment junction with carpus; decalci®ed region just small; distal segment styliform, tapering, ap- distal to junction with carpus. Carpus slightly proximately one-third length of merus; with produced and gently rounded dorsodistally, plumose setae on margins; without ¯agellum. dorsal margin unarmed; lateral surface Pereopod I (®g. 91A) dactylus curved and smooth, with irregular, interrupted row of ru- tapering; lateral and mesial surfaces smooth; gae and submarginal elevated ridge ventrally, dorsal margin with long plumose and short rugae and ridge with long plumose setae; simple setae; ventral margin with short sim- margins with long plumose setae; mesial sur- ple setae. Propodal lateral surface with nu- face smooth, with long plumose setae in scat- merous short, transverse rows of setose ru- tered patches on dorsal half of surface and gae; dorsal margin unarmed; ventral margin on margins. Merus with large median decal- distally produced into acute spine; cutting ci®ed window covering nearly all of lateral edge lacking teeth, lined with long plumose surface, with few scattered setae on surface setae; dorsal margin with long plumose setae, and margins; mesial surface nearly smooth, ventral margin with short simple setae. Car- with few setae, decalci®ed area on proximal pus with dorsodistal angle produced into quarter near junction with basis-ischium. Ba- strong corneous-tipped spine; dorsal margin sis-ischium incompletely fused and unarmed. 280 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Fig. 91. Albunea danai Boyko, 1999: A, F, (, 16.7 mm cl, QM W23105, holotype; B±E, (, 11.6 mm cl, BPBM S5343, paratype; G, &, 16.8 mm cl, BPBM S11782, allotype. A. Left pereopod I, lateral view. B. Right pereopod II, lateral view. C. Right pereopod III, lateral view. D. Right pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 3.0 mm (F, G) and 4.4 mm (A±E).

Coxa with one small spine on anterior mar- smooth, with several small tufts of short se- gin. tae in generally straight line across medi- Pereopod III (®g. 91C) dactylus with base oproximal surface, dorsodistal margin with to heel straight, heel broadly rounded and tufts of short setae; ventral margin with long low, heel to tip with broadly concave indent, plumose setae, dorsal margin with short sim- tip acute, tip to base smoothly convex dis- ple and plumose setae; mesial surface tally to straight proximally; lateral surface smooth, with plumose setae proximally at 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 281 junction with propodus. Propodus not in¯at- ischium incompletely fused and unarmed. ed dorsoventrally; lateral surface smooth, Coxa unarmed. with long plumose setae distally, simple se- Abdomen (®g. 91E) with somite I approx- tae on dorsal margins; dorsolateral surface imately as long as wide, widest posteriorly; narrow, oblique, ¯attened; mesial surface dorsal surface with anterior margin straight; with scattered long setae on and near distal posterior margin straight with elevated sub- margin, with decalci®ed window near junc- marginal row of short setae; with small trans- tion with carpus. Carpus produced dorsodis- verse, decalci®ed windows laterad of seg- tally, exceeding proximal margin of propo- ment median. Somite II dorsal surface with dus by approximately one-third length of submarginal transverse ridge anteriorly; with propodus, rounded; dorsolateral margin un- small transverse decalci®ed windows laterad armed; lateral surface slightly rugose dorso- of segment median just anterior to submar- distally, with mat of short setae and row of ginal ridge; with tuft of setae at posterolateral setae ventrally; mesial surface smooth, with angle, extending onto pleura posteromesially; long plumose setae on margins and scattered pleura expanded and directed slightly ante- on surface. Merus smooth, with large decal- riorly; lateral margins rounded, anterior and ci®ed window covering nearly half of lateral lateral margins with long plumose setae, pos- surface medially; dorsal and ventral margins terior margin with short setae. Somite III unarmed, with long plumose setae; latero- similar to somite II, but narrower, shorter, distal margin with long plumose setae; me- and lacking anterior submarginal ridge; small sial surface smooth, with decalci®ed window tuft of short thick setae on posterolateral an- at junction with basis-ischium. Basis-ischium gle; pleura thinner and shorter than on somite incompletely fused and unarmed. Coxa with II, directed posterolaterally, with setae as in tubercle on anterior margin. Female with somite II; anterolateral angle acute; dorsal large gonopore on anterior mesial margin of surface obliquely ¯attened anterolaterally. coxa, surrounded with short plumose setae; Somite IV similar to somite III, but thinner and shorter; dorsal surface with few thick se- male with small pore. tae posterolaterally; pleura thinner and short- Pereopod IV (®g. 91D) dactylus with base er than on somite III, directed posterolater- to tip convex to concave, tip acute, tip to ally; dorsal surface obliquely ¯attened an- base straight distally, becoming convex prox- terolaterally; margins with long plumose se- imally; lateral surface smooth, ventral mar- tae. Somite V wider than somite IV; lateral gin with long plumose setae, dorsal margin margins with plumose setae; pleura absent. with short simple setae; mesial surface with Somite VI subequal to somite V in length but dorsal decalci®ed region, demarcated ven- wider; dorsal surface with short transverse trally by longitudinal elevated ridge with row rows of setae laterad of midline anteriorly of short setae; with setose punctations ventral and posteriorly; lateral margins with long to decalci®ed window. Propodus expanded plumose setae; pleura absent. dorsally and ventrally; ventral expansion ex- Females with uniramous, paired pleopods ceeds ventral margin of dactylus, margin on somites II±V; males without pleopods. with long plumose setae; dorsal expansion Telson of male (®g. 91F) triangular, slight- with row of long plumose setae medially; lat- ly longer than wide, with smoothly rounded eral and mesial surfaces smooth. Carpus not tip; proximal half heavily calci®ed, distal produced dorsodistally; ventral three-fourths half weakly calci®ed except for large median of lateral surface and mesial surface smooth, region; median longitudinal groove extend- dorsal fourth of lateral surface with mat of ing to distal end of calci®ed area, line with short setae; dorsal margin with short simple long, thin, simple setae; junction of proximal and long plumose setae; ventral margin with and distal regions demarcated by strong line short simple setae. Merus with scattered, of long setae laterally; calci®ed plate slightly short, transverse rows of setae on lateral sur- elevated medially but without ridge. Telson face, dorsal and ventrodistal margins with of female (®g. 91G) ovate, longer than wide, long plumose setae; mesial surface with large rounded distally; dorsal surface smooth, with decalci®ed window proximoventrally. Basis- median longitudinal groove anteriorly; with 282 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 row of setose punctae lateral to midline from Albunea marquisiana Boyko, 2000 posterior end of longitudinal groove to three- Figures 92, 93 fourths of length of telson; margins with long plumose setae. Albunea sp. cf. symnysta [sic] Poupin, 1996b: 26, pl. 12, ®g. c*. ± Poupin, 1998: 39*. DISTRIBUTION: Known only from Oahu, Albunea sp. Tudge et al., 1999: 2±5, ®gs. 1±3*. Hawaii, USA, 4.8±40.2 m depth. Albunea marquisiana Boyko, 2000a: 109±115, MAXIMUM SIZE: Males: 16.7 mm cl; fe- ®gs. 2, 3*. males: 16.8 mm cl. TYPE SPECIMENS: QM W23105 (holotype), MATERIAL EXAMINED: Marquises Islands: BPBM S11782 (allotype), WAM 10422 Sta. TH X, Haul 1, Haava Straits betweenà Ile (paratype), AMNH 17716 (paratype), Tahuata and IÃle Hiva Oa, 09Њ52ЈS, 139Њ04ЈW, AMNH 17717 (paratype), WAM 23390 (2 40 fms (ϭ 73 m), Oct. 1, 1967, coll. D. M. paratypes), BPBM S5343 (paratype), BPBM Devaney for National Geographic Society± S6775 (paratype), BPBM S6777 (paratype). Smithsonian-Bishop Museum Marquesas Ex- TYPE LOCALITY: Halonu Blow Hole dive pedition: 1 (, 10.2 mm cl, holotype (USNM site, south shore, Oahu, Hawaii, USA, 12.2± 268577); Sta. DW 1279, IÃle Eiao, 07Њ59.4ЈS, 13.7 m. 140Њ42.2ЈW, 23±70 m, Sept. 6, 1997, coll. REMARKS: The coloration of this species is MUSORSTOM 9 N/O ``Alis'' Campagne (P. brownish with reddish-brown setae in life Bouchet, B. Dayrat, B. Richer De Forges): 1 (from color transparencies), and a uniform &, 9.3 mm cl, allotype (MNHN-Hi 220); Sta. off-white to tan in preservative. CP 1304, IÃle Nuku Hiva, 08Њ54.4ЈS, Based on all available data, this species 140Њ13.9ЈW, 50±58 m, Sept. 10, 1997, coll. appears to be a true Hawaiian endemic. MUSORSTOM 9 N/O ``Alis'' Campagne (P. However, this conclusion should be accepted Bouchet, B. Dayrat, B. Richer De Forges): 1 cautiously, given the ``endemic'' label ap- (, 7.5 mm cl, paratype (AMNH 17819); Sta. plied to A. speciosa prior to the ®ndings of D 85, IÃle Fatu Hiva, 10Њ29.4ЈS, 138Њ46.5ЈW, Boyko (1999). The holotype of A. danai was 100 m, Jan. 29, 1991, coll. J. Poupin: 8 (, collected with specimens of A. speciosa (QM 5.2±10.0 mm cl, 2 &, 5.6±6.9 mm cl, 5 an- W22285) and the two species are sympatric terior half carapaces, 4.2±7.0 mm cl in Hawaii in at least part of their bathymetric (MNHN-Hi 243), 1 &, 6.4 mm cl, paratype ranges. (AMNH 17820); Sta. NH-VIII, Haul 5, Baie This species belongs to the ``carabus- Hatuatua, IÃle Nuku Hiva, 08Њ51ЈS, 140Њ00ЈW, group'' and is the sister species to A. cara- 26 fms (ϭ 48 m), Sept. 18, 1967, coll. Na- bus, from the Mediterranean and western Af- tional Geographic Society±Smithsonian- rica, and shares with it a similar shape of the Bishop Museum Marquesas Expedition: 1 &, dactyli of pereopods II±IV and telson mor- 5.2 mm cl, paratype (USNM 260948); Sta. phology. Albunea carabus can be easily sep- UP II, Haul 4, off west coast of IÃle Ua Pou, arated from A. danai by its CG8 of four me- 40±45 fms (ϭ 73±82 m), Sept. 23, 1967, dial elements, more strongly crenulated CGs, coll. National Geographic Society±Smithson- more pronounced heel on the dactyli of pe- ian-Bishop Museum Marquesas Expedition: reopods II and III, and a less in¯ated merus 1 (, 8.5 mm cl, paratype (USNM 260949); of maxilliped III. Albunea danai can be dis- Sta. FH I, Haul 1, west coast ofà Ile Fatu tinguished from other Indo-West Paci®c spe- Hiva, 10Њ27±30ЈS, 138Њ40ЈW, 41±43 fms (ϭ cies, except A. bulla, by the triangular shape 75±79 m), Sept. 25, 1967, coll. National of the telson of the male, the rounded dac- Geographic Society±Smithsonian-Bishop tylus of pereopod III, and setal patterns on Museum Marquesas Expedition: 1 anterior the carapace of both sexes. Albunea danai third (USNM 260950); Sta. EO I, Haul 1, off can be separated from A. bulla, which is the northern coast of IÃle Eiao, 08Њ00ЈS, sister taxon to the clade containing A. danai 140Њ50ЈW, 28±29 fms (ϭ 51±53 m), Sept. 21, and A. carabus, by the different pattern of 1967, coll. National Geographic Society± carapace grooves and the distinctly ``button- Smithsonian-Bishop Museum Marquesas Ex- like'' morphology of the telson of the males pedition: 1 (, 7.9 mm cl, paratype (USNM of A. bulla. 260952); Sta. TH IX, Haul 1, off Hana Moe 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 283

Noe, northwest coast of IÃle Tahuata, 09Њ54ЈS, STOM 9 N/O ``Alis'' Campagne (P. Bouchet, 139Њ07ЈW, 37 fms (ϭ 68 m), Oct. 1, 1967, B. Dayrat, B. Richer De Forges): 1 (, 7.2 coll. National Geographic Society±Smithson- mm cl, paratype (MNHN-Hi 229); Sta. DR ian-Bishop Museum Marquesas Expedition: 1245, IÃle Fatu Hiva, 10Њ29.2ЈS, 138Њ36.2ЈW, 1 (, 7.9 mm cl, 1 &, 8.4 mm cl, paratypes 85±130 m, Sept. 1, 1997, coll. MUSOR- (USNM 268578); Sta. DW 1143,Ã Ile Ua Pou, STOM 9 N/O ``Alis'' Campagne (P. Bouchet, 09Њ20.9ЈS, 140Њ02.7ЈW, 18±55 m, Aug. 22, B. Dayrat, B. Richer De Forges): 1 (, 10.1 1997, coll. MUSORSTOM 9 N/O ``Alis'' mm cl, 1 &, 9.4 mm cl, paratypes (MNHN- Campagne (P. Bouchet, B. Dayrat, B. Richer Hi 230); Sta. DW 1256, IÃle Ua Pou, De Forges): 1 (, 4.5 mm cl, paratype 09Њ25.4ЈS, 140Њ07.9ЈW, 70±72 m, Sept. 3, (MNHN-Hi 221); Sta. DW 1162,Ã Ile Nuku 1997, MUSORSTOM 9 N/O ``Alis'' Cam- Hiva, 08Њ56.2ЈS, 140Њ06.1ЈW, 45±64 m, Aug. pagne (P. Bouchet, B. Dayrat, B. Richer De 24, 1997, coll. MUSORSTOM 9 N/O ``Alis'' Forges): 6 (, 5.2±8.7 mm cl, 2 &, 6.0±7.4 Campagne (P. Bouchet, B. Dayrat, B. Richer mm cl, 1 posterior half of oviger (MNHN- De Forges): 1 anterior half, 4.7 mm cl Hi 231); Sta. DW 1260, IÃle Ua Pou, (MNHN-Hi 222); Sta. DW 1185,Ã Ile Nuku 09Њ25.4ЈS, 140Њ07.3ЈW, 49±100 m, Sept. 3, Hiva, 08Њ48.9ЈS, 140Њ03.4ЈW, 31±33 m, Aug. 1997, coll. MUSORSTOM 9 N/O ``Alis'' 26, 1997, coll. MUSORSTOM 9 N/O ``Alis'' Campagne (P. Bouchet, B. Dayrat, B. Richer Campagne (P. Bouchet, B. Dayrat, B. Richer De Forges): 2 (, 8.4±8.9 mm cl, 1 &, 9.7 De Forges): 1 (, 7.6 mm cl, 1 anterior half, mm cl, paratypes (MNHN-Hi 232); Sta. DW 9.5 mm cl (MNHN-Hi 222 bis); Sta. DW 1266, IÃle Eiao, 07Њ57.3ЈS, 140Њ42.6ЈW, 84 m, 1180, IÃle Nuku Hiva, 08Њ46.2ЈS, 140Њ04.6ЈW, Sept. 4, 1997, coll. MUSORSTOM 9 N/O 80±82 m, Aug. 26, 1997, coll. MUSOR- ``Alis'' Campagne (P. Bouchet, B. Dayrat, B. STOM 9 N/O ``Alis'' Campagne (P. Bouchet, Richer De Forges): 1 (, 5.6 mm cl, paratype B. Dayrat, B. Richer De Forges): 1 (, 10.2 (MNHN-Hi 233); Sta. DW 1279,Ã Ile Eiao, mm cl, paratype (MNHN-Hi 223); Sta. CP 07Њ59.4ЈS, 140Њ42.2ЈW, 23±70 m, Sept. 6, 1187, IÃle Nuku Hiva, 08Њ49.2ЈS, 140Њ03.5ЈW, 1997, coll. MUSORSTOM 9 N/O ``Alis'' 25±30 m, Aug. 26, 1997, coll. MUSOR- Campagne (P. Bouchet, B. Dayrat, B. Richer STOM 9 N/O ``Alis'' Campagne (P. Bouchet, De Forges): 4 &, 3.5±8.1 mm cl, paratypes B. Dayrat, B. Richer De Forges): 3 (, 8.9± (MNHN-Hi 234); Sta. DW 1283,Ã Ile Mutu 10.8 mm cl, paratypes (MNHN-Hi 224); Sta. One, Hatutaa, 07Њ53.8ЈS, 140Њ34.5ЈW, 55±56 DW 1213, IÃle Hiva Oa, 09Њ50.3ЈS, m, Sept. 7, 1997, coll. MUSORSTOM 9 N/ 139Њ03.2ЈW, 18±20 m, Aug. 29, 1997, coll. O ``Alis'' Campagne (P. Bouchet, B. Dayrat, MUSORSTOM 9 N/O ``Alis'' Campagne (P. B. Richer De Forges): 1 (, 5.4 mm cl, para- Bouchet, B. Dayrat, B. Richer De Forges): 1 type (MNHN-Hi 235); Sta. DW 1292, &, 5.3 mm cl (MNHN-Hi 225); Sta. DW 08Њ54.1ЈS, 139Њ37.8ЈW, Ile Ua Huka, 95±100 1214, IÃle Hiva Oa, 09Њ49.8ЈS, 139Њ03.1ЈW, m, Sept. 8, 1997, coll. MUSORSTOM 9 N/ 25±40 m, Aug. 29, 1997, coll. MUSOR- O ``Alis'' Campagne (P. Bouchet, B. Dayrat, STOM 9 N/O ``Alis'' Campagne (P. Bouchet, B. Richer De Forges): 1 (, 6.1 mm cl B. Dayrat, B. Richer De Forges): 1 anterior (MNHN-Hi 257); Sta. DW 1297,Ã Ile Ua half, 7.1 mm cl (MNHN-Hi 226); Sta. DW Huka, 08Њ54.2ЈS, 139Њ37.4ЈW, 90±150 m, 1217, IÃle Hiva Oa, 09Њ44.5ЈS, 138Њ49.9ЈW, Sept. 8, 1997, coll. MUSORSTOM 9 N/O 85±87 m, Aug. 30, 1997, coll. MUSOR- ``Alis'' Campagne (P. Bouchet, B. Dayrat, B. STOM 9 N/O ``Alis'' Campagne (P. Bouchet, Richer De Forges): 1 (, 7.9 mm cl, paratype B. Dayrat, B. Richer De Forges): 1 (, 5.1 (MNHN-Hi 236), 1 &, 4.8 mm cl, paratype mm cl, paratype (MNHN-Hi 227); Sta. DW (MNHN-Hi 237); Sta. CP 1304, IÃle Nuku 1241, IÃle Fatu Hiva, 10Њ27.8ЈS, 138Њ40.6ЈW, Hiva, 08Њ54.4ЈS, 140Њ13.9ЈW, 50±58 m, Sept. 85±130 m, Sept. 1, 1997, coll. MUSOR- 10, 1997, coll. MUSORSTOM 9 N/O ``Alis'' STOM 9 N/O ``Alis'' Campagne (P. Bouchet, Campagne (P. Bouchet, B. Dayrat, B. Richer B. Dayrat, B. Richer De Forges): 1 (, 6.2 De Forges): 3 (, 6.3±8.0 mm cl, paratypes mm cl, paratype (MNHN-Hi 228); Sta. DW (MNHN-Hi 238); Sta. 24, CoÃte NW Baie 1242, IÃle Fatu Hiva, 10Њ28.1ЈS, 138Њ41.1ЈW, Haahue,Ã Ile Ua Huka, 08Њ53.6ЈS, 139Њ37ЈW, 119±122 m, Sept. 1, 1997, coll. MUSOR- 9±25 m, Oct. 1997, coll. R. Von Cosel, J. 284 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

TroÈndleÂ, and J. Tardy: 1 (, 4.3 mm cl, 4 &, posterior lateral elements of CG1. CG1 par- 5.8±10.9 mm cl, 6 juveniles, 3.1±3.8 mm cl, allel to anterior margin of carapace, sinuous, 2 unsexable, unmeasurable specimens slightly crenulate, divided into medial sinu- (MNHN-Hi 239); Sta. 29, Baie de Hane, IÃle ous fragment and curved, posteriorly dis- Ua Huka, 08Њ55.7ЈS, 139Њ32.0ЈW, 7±11 m, placed lateral elements. Mesogastric region Oct. 1997, coll. R. Von Cosel, J. TroÈndleÂ, and smooth; CG2 absent; CG3 broken into four J. Tardy: 2 &, 5.0±5.1 mm cl (MNHN-Hi to six short elements; CG4 with two long lat- 240); Sta. 34, Baie Haavei, Pointe Tenoni, IÃle eral elements terminating above apex of Teuaua, IÃle Ua Huka, 08Њ56.8ЈS, 139Њ35.7ЈW, CG6, occasionally with two short medial el- 10±15 m, Oct. 1997, coll. R. Von Cosel, J. ements. Hepatic region smooth, with long se- TroÈndleÂ, and J. Tardy: 1 (, 6.3 mm cl, para- tose groove at median of lateral margin. Epi- type (MNHN-Hi 241); Sta. D 77, IÃle Eiao, branchial region generally triangular, smooth. 07Њ56.7ЈS, 139Њ30.8ЈW, 54 m, Jan. 21, 1991, Metagastric region smooth; CG5 broken into coll. J. Poupin: 1 (, 10.0 mm cl, paratype two short oblique elements. CG6 strongly (MNHN-Hi 242). crenulate, strongly anteriorly concave medi- DIAGNOSIS: Carapace wider than long, cov- ally and sloping out to anteriorly convex lat- ered with lightly setose grooves. Anterior eral thirds. CG7 present as two long oblique margin with 8±10 spines on either side of elements and separate from CG6. Cardiac re- ocular sinus. Setal ®eld with narrow lateral gion smooth; CG8 with two to four median elements and sinuous anterior margin. CG1 elements and two slightly longer lateral ele- with separate posterior lateral elements; CG4 ments. CG9 present as two short, widely sep- with two long lateral elements terminating arated elements. CG10 present as two above apex of CG6, occasionally with two straight lateral fragments, with gap between short medial elements; CG5 broken into two fragments greater than length of single frag- short oblique elements; CG6 and CG7 sep- ment. CG11 present as two or three irregu- arate; CG8 broken; CG11 present. Rostrum larly spaced short elements. Branchial region present, not reaching posterior margin of oc- with numerous short, transverse rows of se- ular plate. Ocular plate triangular. Distal pe- tae. Posterior margin deeply and evenly con- duncular segments dorsoventrally ¯attened vex, with submarginal groove reaching ap- and subtriangular in shape, tapering at tip, proximately three-fourths up either side of approximated along mesial margins, lateral posterior concavity. Branchiostegite with margins convex except slightly concave at short anterior submarginal spine; anterior re- tip, mesial margins straight. Cornea at tip. gion with scattered short, transverse lines Dactylus of pereopod II with heel produced ventral to linea anomurica; with many short and subquadrate. Dactylus of pereopod III rows of setae and sparsely covered with long with heel slightly produced and acute. Dac- plumose setae ventrally; posterior region tylus of pereopod IV evenly sinuous from membranous, with numerous irregular frag- base to tip. Telson of male spatulate, tip ments and sparsely covered with long plu- rounded, dorsal surface in¯ated medially mose setae. with strong medial ridge lined with short Ocular plate (®g. 92B) subquadrate, with thick setae; lateral margins decalci®ed. Tel- shallow median indentation; median pedun- son of female ovate. cular segments reduced to small rounded cal- DESCRIPTION: Carapace (®g. 92A) slightly ci®ed area on either side of ocular plate. Dis- wider than long. Anterior margin concave on tal peduncular segments elongate, with prox- either side of ocular sinus, becoming convex imally convex and distally concave lateral laterally, with 8±10 large spines along length. margins, tapering to produced distal cornea; Rostrum as small acute tooth, not reaching mesial margins approximated along entire proximal margin of ocular plate. Ocular sinus length; mesial and ventral margins of seg- smoothly concave and unarmed. Frontal re- ment with sparse row of long plumose setae; gion smooth; setal ®eld broad posteriorly, few small tufts of plumose setae on proximal narrowing anteriorly, with narrow anterior quarter of segment. lateral elements and sinuous anterior margin; Antennule (®g. 92C) with segment III sub- posterior lateral elements not reaching to cylindrical; with plumose setae on dorsal and 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 285

Fig. 92. Albunea marquisiana Boyko, 2000: A, (, 10.2 mm cl, USNM 268577, holotype; B±J, (, 7.9 mm cl, USNM 268578, paratype. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 1.6 mm (B, E, F, I), 2.2 mm (D, G, H, J), and 3.3 mm (A, C). 286 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 ventral margins; dorsal exopodal ¯agellum with two segments; proximal segment nar- with 110±118 articles, long plumose setae on row, margins parallel, with plumose setae; dorsal and ventral margins; ventral endopo- distal segment spatulate, approximately as dal ¯agellum short with two articles and plu- long as wide, broadest medially, margins mose setae on dorsal and ventral margins. with long plumose setae. Endopod ¯attened Segment II medially in¯ated in dorsal view, and elongate, reaching to distal end of prox- with plumose setae on dorsal and ventral imal exopodal segment, with plumose setae margins and scattered on ventrolateral third on margins. of surface. Segment I wider than long, un- Maxilliped II (®g. 92I) dactylus evenly armed; dorsal third of lateral surface rugose rounded, length slightly greater than width, with long plumose setae; long plumose setae with thick simple setae distally. Propodus on dorsal and ventral margins. two times wider than long, with plumose se- Antenna (®g. 92D) with segment V ap- tae on dorsal margin and long simple setae proximately two times longer than wide, with on distal margin. Carpus not strongly pro- long plumose setae on dorsal margins; ¯a- duced dorsodistally, approximately three gellum with seven articles, long plumose se- times longer than wide, with long simple se- tae on dorsal, ventral, and distal margins. tae on dorsal margin. Merus approximately Segment IV expanded distally, with long plu- three times longer than wide, margins par- mose setae on dorsal, ventral, and distal mar- allel, with simple setae on ventrolateral mar- gins and row of setae on dorsolateral margin. gin and scattered on surface, plumose setae Segment III with long plumose setae on dor- on dorsolateral margin. Basis-ischium in- sal and ventral margins. Segment II short, completely fused, with plumose setae on widening distally, with plumose setae on margins. Exopod one-fourth longer than mer- margins; antennal acicle long, thin, truncate us, ¯agellum with one elongate article. distally, slightly exceeding distal margin of Maxilliped III (®g. 92J) dactylus evenly segment IV, with long plumose setae on dor- rounded; with long plumose setae on margins sal margin. Segment I rounded proximally, and lateral surface. Propodus with longitu- ¯attened ventrolaterally, with long plumose dinal median row of plumose setae on lateral setae on margins; lateral surface with acute surface; margins with plumose setae. Carpus spine subdorsally, with low semicircular, dor- slightly produced onto propodus; lateral sur- solateral lobe ventrodistal to spine; segment face with row of plumose setae ventromedi- with ventromesial antennal gland pore. ally; plumose setae on margins. Merus un- Mandible (®g. 92E) incisor process with- armed, with plumose setae on margins and out teeth; cutting edge with one tooth. Palp scattered on surface. Basis-ischium incom- three-segmented, with plumose setae on mar- pletely fused, with faint crista dentata of few gins and long, thick, simple setae arising small low teeth. Exopod two-segmented: from bend in second segment. proximal segment small; distal segment sty- Maxillule (®g. 92F) distal endite proxi- liform, tapering, approximately one-third mally narrow, widening to in¯ated distal end, length of merus, with plumose setae on mar- with thick simple setae on distal margin; plu- gins; without ¯agellum. mose setae on dorsal margin. Proximal endite Pereopod I (®g. 93A) dactylus curved and with thick simple setae on distal margin. En- tapering; lateral and mesial surfaces smooth; dopodal external lobe truncate distally and dorsal margin with long plumose and short curled under; internal lobe reduced with simple setae; ventral margin with short sim- three thick setae at distolateral margin. ple setae. Propodal lateral surface with nu- Maxilla (®g. 92G) exopod evenly rounded, merous short, transverse rows of setose ru- with plumose setae along distal margin. Sca- gae; dorsal margin unarmed; ventral margin phognathite bluntly angled on posterior lobe, distally produced into acute spine; cutting with plumose setae. edge lacking teeth, lined with long plumose Maxilliped I (®g. 92H) epipod with plu- setae; dorsal margin with long plumose setae, mose setae on distal margin and on distola- ventral margin with short simple setae. Car- teral surface. Endite tapered distally and sub- pus with dorsodistal angle produced into equal to ®rst segment of exopod. Exopod small spine, dorsal margin smooth; dorsal 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 287

Fig. 93. Albunea marquisiana Boyko, 2000: A±F, (, 7.9 mm cl, USNM 268578, paratype; G, &, 5.2 mm cl, USNM 260948, paratype. A. Left pereopod I, lateral view. B. Left pereopod II, lateral view. C. Right pereopod III, lateral view. D. Right pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 1.1 mm (G), 1.6 mm (F), 3.3 mm (A±D), and 6.7 mm (E). and distal margins with long plumose setae; Pereopod II (®g. 93B) with dactylus lateral surface with small distal rugose area, smooth; with base to heel slightly concave, few transverse setose ridges on distal half of heel smoothly rounded and slightly pro- surface; mesial surface smooth, with median duced, heel to tip with acute, narrow indent, row of long plumose setae, margins with tip acute, tip to base broadly convex; lateral long plumose setae. Merus unarmed; lateral surface smooth, with several small tufts of surface with scattered transverse rows of short setae in generally straight line across long plumose setae, margins with long plu- medioproximal surface, several widely mose setae; mesial surface with few short spaced submarginal tufts of short setae dor- rows of setae; proximal fourth of mesial sur- sodistally; mesial surface smooth, ventral face with decalci®ed window. Basis-ischium margin with long plumose setae, dorsal mar- incompletely fused, unarmed. Coxa un- gin with short simple setae, with patch of armed. long plumose setae at base. Propodus with 288 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 dorsal surface smooth, ventral margin in¯at- margin unarmed; lateral surface slightly ru- ed and rounded; oblique row of long plu- gose dorsodistally, with mat of short setae mose setae on distal margin of lateral sur- and two longer rows of setae ventrally; me- face; distal and ventral margin with long plu- sial surface smooth, with long plumose setae mose setae; dorsolateral surface as narrow, on margins and in transverse row on surface. oblique, ¯attened shelf, with short setae on Merus smooth, with large decalci®ed area dorsal margin and long plumose setae on near median of lateral surface; dorsal and ventral margin; mesial surface with elevated, ventral margins unarmed, with long plumose curved, setose ridge from ventral junction setae; distolateral margin with long plumose with dactylus almost to ventral proximal setae; mesial surface smooth, with decalci- junction with carpus; decalci®ed region just ®ed window at junction with basis-ischium. distal to junction with carpus. Carpus slightly Basis-ischium incompletely fused and un- produced dorsodistally; lateral surface nearly armed. Coxa unarmed. Female with large smooth, with irregular, interrupted row of ru- gonopore on anterior mesial surface of coxa, gae and submarginal elevated ridge ventrally, surrounded with short plumose setae; male rugae and ridge with long plumose setae; without pore. margins with long plumose setae; mesial sur- Pereopod IV (®g. 93D) dactylus with base face smooth with long plumose setae in scat- to tip convex to concave, tip acute, tip to tered patches on surface and on margins. base straight distally, becoming convex prox- Merus lateral surface with large decalci®ed imally; lateral surface smooth, ventral mar- area in median, few scattered setae on sur- gin with long plumose setae, dorsal margin face and margins and large patch of long with short simple setae; mesial surface with simple setae at distolateral margin; mesial dorsal decalci®ed region, demarcated ven- surface nearly smooth, with many median se- trally by longitudinal elevated ridge with row tae, with decalci®ed area on proximal fourth of short setae; with setose punctae ventral to near junction with basis-ischium. Basis-is- decalci®ed window. Propodus expanded dor- chium incompletely fused and unarmed. sally and ventrally; ventral expansion not ex- Coxa with one small tubercle on anterior ceeding ventral margin of dactylus, margin margin. with long plumose setae; dorsal expansion Pereopod III (®g. 93C) dactylus with base with row of long plumose setae medially and to heel concave, heel broadly rounded and mat of short simple setae; lateral and mesial slightly produced, heel to tip with broad, surfaces smooth, mesial surface with large evenly rounded indent, tip acute, tip to base decalci®ed area. Carpus not produced dor- smoothly convex to straight; lateral surface sodistally; lateral and mesial surfaces smooth, with several small tufts of short se- smooth; dorsal margin with short simple and tae in generally straight line across medi- long plumose setae; ventral margin with oproximal surface, dorsodistal margin with short simple setae and small mat of short tufts of short setae; ventromesial margin with simple setae dorsally. Merus with scattered, long plumose setae, dorsal margin with short short, transverse rows of setae on lateral sur- simple and plumose setae; mesial surface face, dorsal and ventrodistal margins with smooth, with plumose setae proximally at long plumose setae; mesial surface with large junction with propodus. Propodus not in¯at- decalci®ed window proximoventrally. Basis- ed dorsoventrally; lateral surface smooth, ischium incompletely fused and unarmed. with long plumose setae distally, with simple Coxa unarmed. setae on dorsal margins, and long plumose Abdomen (®g. 93E) with somite I approx- setae on ventral margin; dorsolateral surface imately as long as wide, widest posteriorly; narrow, oblique, ¯attened; mesial surface dorsal surface with anterior margin straight; with scattered long setae on and near distal posterior margin concave, with elevated sub- margin, with decalci®ed window near junc- marginal row of short setae; with small trans- tion with carpus. Carpus produced dorsodis- verse, decalci®ed windows laterad of seg- tally, exceeding proximal margin of propo- ment median. Somite II dorsal surface with dus by approximately one-fourth length of submarginal transverse ridge anteriorly; with propodus, pointed but not acute; dorsolateral small transverse, decalci®ed windows laterad 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 289 of segment median just anterior to submar- USNM 268578 (2 paratypes), MNHN-Hi ginal ridge; with tuft of setae at posterolateral 221 (paratype), MNHN-Hi 223 (paratype), angle, extending onto pleura posteromesially; MNHN-Hi 224 (3 paratypes), MNHN-Hi posterior margin with indistinct punctate sub- 227 (paratype), MNHN-Hi 228 (paratype), marginal groove laterally; pleura expanded MNHN-Hi 229 (paratype), MNHN-Hi 230 (2 and directed slightly anteriorly; lateral mar- paratypes), MNHN-Hi 232 (3 paratypes), gins rounded, anterior and lateral margins MNHN-Hi 233 (paratype), MNHN-Hi 234 (4 with long plumose setae, posterior margin paratypes), MNHN-Hi 235 (paratype), with short setae. Somite III similar to somite MNHN-Hi 236 (paratype), MNHN-Hi 237 II, but narrower, shorter, and lacking anterior (paratype), MNHN-Hi 238 (3 paratypes), submarginal ridge; small tuft of short thick MNHN-Hi 241 (paratype), MNHN-Hi 242 setae on posterolateral angle; pleura thinner (paratype). and shorter than on somite II, directed an- TYPE LOCALITY: Haava Straits between IÃle terolaterally, with setae as in somite II; an- Tahuata and IÃle Hiva Oa, Marquises Islands, terolateral angle acute; dorsal surface 09Њ52ЈS, 139Њ04ЈW, 73 m. obliquely ¯attened anterolaterally. Somite IV REMARKS: Juvenile specimens are virtually similar to somite III, but thinner and shorter; lacking in pigment and appear almost white. dorsal surface with thick setae posterolater- Adults are a uniform tan, with reddish-brown ally; pleura thinner and shorter than on so- setae. Larger specimens show a markedly in- mite III, directed posterolaterally; dorsal sur- creased reddish tone on the carapace (espe- face obliquely ¯attened anterolaterally; mar- cially the anterior region), ocular peduncles, gin with long plumose setae. Somite V sub- antennae, and antennules (from preserved equal to somite IV; lateral margins with material and a color transparency made of a plumose setae; pleura absent. Somite VI sub- live specimen [MNHN-Hi 223]; see also equal to somite V in width but longer; dorsal Poupin, 1996b). surface with short transverse rows of setae Little is known about the biology of this laterad of midline anteriorly; pleura absent. species. Ovigerous females, unfortunately, Females with uniramous, paired pleopods are only known from the posterior portion of on somites II±V; males lacking pleopods. a single specimen. Additionally, the mor- Telson of male (®g. 93F) spatulate, with phology of the spermatozoa of this species length greater than width, rounded distally; weakly calci®ed at margins of large calci®ed has been studied by Tudge et al. (1999), median plate; median longitudinal groove based on one of the paratypes (MNHN-Hi short, extending one-fourth length of telson; 236). thick elevated ridge continuing from end of Albunea marquisiana is most similar to A. groove to distal end of telson, lined with holthuisi and A. groeningi. All three species dense row of thick simple setae. Telson of share the distinctive thick median row of se- female (®g. 93G) ovate, longer than wide, tae on the telson of the male, as well as a dorsal surface smooth, with median longitu- general similarity in the shape of the pereo- dinal groove reaching almost to distal mar- podal dactyli and can be collectively consid- gin; with row of setose punctae lateral to ered to form the ``holthuisi-group'' of spe- midline along whole length of median cies. They can easily be separated by the groove except proximal fourth; margins with number of elements of CG8 (one long me- long plumose setae. dian element in A. holthuisi, three or four DISTRIBUTION: Known only from the Mar- short elements in A. marquisiana and A. quises Islands, in 7±130 m depth. groeningi), and CG11 (absent in A. groen- MAXIMUM SIZE: Males: 10.8 mm cl; fe- ingi, one in A. holthuisi, two or three in A. males: 10.9 mm cl. marquisiana), the relative thickness of the TYPE SPECIMENS: USNM 268577 (holo- proximal blade of pereopod III (thicker in A. type), MNHN-Hi 220 (allotype), AMNH marquisiana and A. groeningi), and the distal 17819 (paratype), AMNH 17820 (paratype), margins of both the male and female telsons USNM 260948 (paratype), USNM 260949 (truncate in A. holthuisi, smoothly rounded (paratype), USNM 260952 (paratype), in A. marquisiana, indented in A. groeningi). 290 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Albunea holthuisi Boyko and Harvey, 1999 DIAGNOSIS: Carapace slightly longer than Figures 94, 95 wide, covered with lightly setose grooves. Anterior margin with 8±11 spines on either ?Albunea symnista [sic]: Ward, 1942: 52 (list), 63 side of ocular sinus. Setal ®eld with narrow (not Albunea symmysta (Linnaeus, 1758)). ?Albunea steinitzi: Thomassin, 1969: 143±146, pl. lateral elements and concave anterior margin. 3, ®gs. 1±8, text-®gs. 3c, 4 (not Albunea stein- CG1 with separate posterior lateral elements; itzi Holthuis, 1958). CG4 with short element on either side of me- Albunea holthuisi Boyko and Harvey, 1999: 386± dian with missing elements at midline and 391, 400 (list), 401 (key), ®gs. 6, 7*. between median and laterals; CG5 present only as short lateral elements; CG6 and CG7 MATERIAL EXAMINED: Zanzibar: Sta. 651, separate; CG8 complete; CG11 present. Ros- dredged grass and shell, 1.5 mi west-south- trum present, not reaching posterior margin west of Ras Nungwa, 8 fms (ϭ 14.6 m), Feb. of ocular plate. Ocular plate triangular. Distal 20, 1957, coll. A. J. Ostheimer III: 1 (, 7.7 peduncular segments dorsoventrally ¯attened mm cl, paratype (ANSP CA4644); Sta. 651, and oblong in shape, tapering at tip, approx- ®ne grass and shell, 1.5 mi west-southwest imated along distal two-thirds of mesial mar- of Ras Nungwa, 8 fms (ϭ 14.6 m), Feb. 20, gins, lateral margins convex except slightly 1957, coll. A. J. Ostheimer III: 1 &, 6.7 mm concave at tip, mesial margins convex. Cor- cl, paratype (ANSP CA4645). nea at tip. Dactylus of pereopod II with heel Madagascar: Environs de Nosy Be, CoÃte slightly produced, low and rounded. Dacty- southwest, 13Њ37.7ЈS, 47Њ49.6ЈE, 25 m, coll. lus of pereopod III with heel slightly pro- unknown: 1 (, 8.1 mm cl, holotype jecting, acute. Dactylus of pereopod IV sin- (MNHN-Hi 202); Andilana, Nosy Be, CoÃte uous from base to tip, with slight indent. Tel- southwest, Sept. 1959, coll. A. Crosnier: 1 son of male spatulate, tip broadly truncate, &, 8.1 mm, allotype (MNHN-Hi 203); vois- dorsal surface with elevated, median, longi- inage de Nosy Be, CoÃte northwest, tudinal ridge bearing short thick setae prox- 13Њ38.3ЈS, 42Њ49.6ЈE, 34 m, coll. A. Cros- imally and long thick setae distally. Telson nier: 1 (, 8.0 mm cl, paratype (MNHN-Hi of female ¯attened and spatulate, longitudi- 204); Sakatia, Envoi II, 1921, coll. G. Petit: nal row of short, thin setae medially. 1 (, 6.7 mm cl (MNHN-Hi 19). DESCRIPTION: Carapace (®g. 94A) slightly Seychelles: Sta. 5, 33 m, Sept. 4, 1980, wider than long. Anterior margin concave on coll. ORSTOM-Reves 2: 1 (, 9.2 mm cl either side of ocular sinus, becoming convex (MNHN-Hi 249); MaheÂ, July±Aug. 1972, laterally, 8±11 large spines on concave re- coll. Mission Zoologique MRAC-ULB: 1 &, gion, ventral row of long plumose setae sub- 5.7 mm cl (MRAC 57.459). marginally. Rostrum as small acute tooth, not Indonesia: Sta. 522, west side of Samber- reaching to proximal margin of ocular plate. baban Bay, Japen Island, Irian Jaya, Feb. 14, Ocular sinus smoothly concave and unarmed. 1956, coll. C. T. Abbot on ``Gloria Maris'': Frontal region smooth; setal ®eld broad pos- 1 (, 11.3 mm cl, paratype (RMNH 23703); teriorly, narrowing anteriorly, with narrow Corindon II, Sta. B255, Makassar, 01Њ56.5ЈS, lateral elements and concave anterior margin. 119Њ17.3ЈE, 13 m, Nov. 6, 1980, coll. R/V CG1 parallel to anterior margin of carapace, ``Coriolis'': 1 &, 7.1 mm cl, paratype sinuous, slightly crenulate, divided into me- (MNHN-Hi 205); Corindon II, Sta. B256, dial fragment and curved lateral elements Makassar, 01Њ56.5ЈS, 119Њ17.2ЈE, 24 m, Nov. that are displaced posteriorly. Mesogastric 6, 1980, coll. R/V ``Coriolis'': 1 (, 4.9 mm region smooth, CG2 short; CG3 broken into cl, 1 &, 5.4 mm cl (MNHN-Hi 206). six short elements approximately equally Malaysia: Juara Bay, Pulau Tioman, Pa- spaced between posterior elements of CG1; hang, May 26, 1985, coll. P.K.L. Ng: 1 (, CG4 fragmented into four elements with gap 7.5 mm cl (ZRC 1989.3827). at midline and between median and lateral Australia: Queensland: Little Trunk elements. Hepatic region smooth, with short Reef, 18Њ20ЈS, 146Њ46ЈE, 9.1±12.2 m, Nov. setose groove at median of lateral margin. 5, 1990, coll. K. Lamprell: 1 (, 9.6 mm cl Epibranchial region generally triangular, (QM W24961). smooth, posterolateral margin with short row 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 291

Fig. 94. Albunea holthuisi Boyko and Harvey, 1999: A, B, (, 8.1 mm cl, MNHN Hi-202, holotype; C, (, 7.7 mm cl, ANSP CA4644, paratype; D±J, (, 8.0 mm cl, MNHN Hi-204, paratype. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 1.1 mm (B), 1.6 mm (E, F, I), and 3.3 mm (A, C, D, G, H, J). of setae. Metagastric region smooth; CG5 gle fragment; CG11 present. Branchial re- present only as short lateral elements directly gion with numerous short, transverse rows of posterior to median elements of CG4; CG6 setae. Posterior margin deeply and evenly slightly crenulate, strongly concave medially convex, with submarginal groove interrupted and sloping out to convex lateral thirds; CG7 medially. Branchiostegite with short anterior transverse and separate from CG6. Cardiac submarginal spine, anterior region with scat- region smooth; CG8 uninterrupted; CG9 pre- tered short, transverse lines ventral to linea sent only as lateral short lines; CG10 present anomurica, with many short rows of setae in two fragments, separated by length of sin- and covered with long plumose setae ven- 292 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 trally, posterior region membranous, with nu- gins and long, thick, simple setae arising merous irregular fragments, and covered from bend in second segment. with long plumose setae. Maxillule (®g. 94F) distal endite proxi- Ocular plate (®g. 94B) triangular, with mally narrow, widening to in¯ated distal end, shallow median indentation; median pedun- with thick simple setae on distal margin. cular segments present as small, ovate cal- Proximal endite with thick simple setae on ci®ed areas lateral to ocular plate. Distal pe- distal margin. Endopodal external lobe trun- duncular segments elongate, with distally cate distally and curled under, notched prox- convex lateral margins, tapering to rounded imally; internal lobe reduced with two thick distolateral cornea, mesial margins approxi- setae at distolateral margin. mated along entire length, mesial and ventral Maxilla (®g. 94G) exopod evenly rounded, margins of segment with sparse row of long with plumose setae along distal margin. Sca- plumose setae, tuft of plumose setae at prox- phognathite bluntly angled on posterior lobe, imal lateral ventral angle, ventral surface with plumose setae. with oblique row of long plumose setae from Maxilliped I (®g. 94H) with epipod with proximal lateral angle almost to distal mesial plumose setae on distal margin and on dis- margin. tolateral surface. Endite tapered distally and Antennule (®g. 94C) with segment III nar- subequal to ®rst segment of exopod. Exopod row proximally, expanding distally to twice with two segments; proximal segment nar- proximal width; plumose setae on dorsal and row, margins parallel, with plumose setae; ventral margins, dorsal exopodal ¯agellum distal segment spatulate, slightly longer than with 76±104 articles, long plumose setae on wide, broadest medially, margins with long dorsal and ventral margins, ventral endopo- plumose setae. Endopod ¯attened and elon- dal ¯agellum short, with two or three articles, gate, reaching to distal end of proximal ex- plumose setae on dorsal and ventral margins. opodal segment, with plumose setae on mar- Segment II medially in¯ated in dorsal view, gins. plumose setae on dorsal and ventral margins Maxilliped II (®g. 94I) dactylus evenly and scattered on lateral surface. Segment I rounded, length equal to width, with thick wider than long, unarmed, lateral surface simple setae distally. Propodus 1.5 times with long plumose setae dorsally and on dor- wider than long, with plumose setae on dor- sal and ventral margins. sal margin and long simple setae on distal Antenna (®g. 94D) with segment V ap- margin. Carpus not strongly produced dor- proximately three times longer than wide, sodistally, approximately two times longer long plumose setae on dorsal and ventral than wide, with long simple setae on dorsal margins, ¯agellum with seven articles, long margin. Merus approximately three times plumose setae on dorsal, ventral, and distal longer than wide, margins parallel, with sim- margins. Segment IV expanded distally, long ple setae on ventrolateral margin and plu- plumose setae on dorsal, ventral, and distal mose setae on dorsolateral margin. Basis-is- margins and simple setae on dorsolateral chium incompletely fused, with plumose se- margin. Segment III with long plumose setae tae on margins. Exopod one-fourth longer on ventral margin. Segment II short, widen- than merus, ¯agellum with one article. ing distally, plumose setae on margins, an- Maxilliped III (®g. 94J) dactylus rounded tennal acicle long, thin, exceeding base of at tip, long plumose setae on margins and segment V by approximately one-half length lateral surface. Propodus with longitudinal of segment V, long plumose setae on dorsal median row of plumose setae on lateral sur- margin. Segment I rounded proximally, ¯at- face, margins with plumose setae. Carpus tened ventromesially, long plumose setae on slightly produced onto propodus, lateral sur- margins; lateral surface with acute spine dor- face with row of plumose setae ventromedi- sally, with low, semicircular dorsolateral lobe ally; plumose setae on margins. Merus un- ventrodistal to spine. armed, plumose setae on margins. Basis in- Mandible (®g. 94E) incisor process with completely fused with ischium; weak crista one tooth; cutting edge with one tooth. Palp dentata of two or three teeth. Exopod two- three-segmented, with plumose setae on mar- segmented, proximal segment small, distal 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 293

Fig. 95. Albunea holthuisi Boyko and Harvey, 1999: A, B, (, 8.0 mm cl, MNHN Hi-204, paratype; C, D, H, &, 8.1 mm cl, MNHN Hi-203, allotype; E±G, (, 8.1 mm cl, MNHN Hi-202, holotype. A. Left pereopod I, lateral view. B. Left pereopod II, lateral view. C. Left pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of (, lateral view. H. Telson of &, dorsal view. Scale ϭ 1.7 mm (F, H) and 3.3 mm (A±E, G). segment styliform, tapering, approximately dorsal margin with long plumose and short one-third length of merus, plumose setae on simple setae, short simple setae on ventral margins; ¯agellum absent. margin. Propodal lateral surface with numer- Pereopod I (®g. 95A) dactylus curved and ous short, transverse rows of setose rugae; tapering; lateral and mesial surfaces smooth; dorsal margin unarmed; ventral margin dis- 294 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 tally produced into acute spine; cutting edge to heel concave, heel produced in short, acute lacking teeth, lined with long plumose setae; spur, heel to indent nearly straight, indent lateral, mesial, and ventral margins with long broadly concave, tip acute, tip to base setae. Carpus with dorsodistal angle pro- smoothly convex to straight; lateral surface duced into small corneous spine; dorsal and smooth, dorsodistal margin with tufts of distal margins with long plumose setae; lat- short setae, ventromesial margin with long eral surface with distal rugose area, few plumose setae, dorsal margin with short sim- transverse setose ridges on distal two-thirds ple and plumose setae; mesial surface of surface; mesial surface smooth, with scat- smooth, plumose setae proximally at junction tered rows of long plumose setae, margins with propodus. Propodus weakly in¯ated; with long plumose setae. Merus unarmed; lateral surface smooth, long plumose setae lateral surface with scattered transverse rows distally, simple setae on margins, long plu- of long plumose setae, margins with long mose setae on ventral margin, dorsolateral plumose setae; mesial surface with few short surface narrow, oblique, ¯attened; mesial rows of setae. Basis-ischium incompletely surface with scattered long setae on and near fused, unarmed. Coxa unarmed. distal margin. Carpus produced dorsodistally, Pereopod II (®g. 95B) dactylus smooth; exceeding proximal margin of propodus by base to heel concave, heel with smoothly approximately one-third length of propodus, rounded low spur, heel to tip acutely indented broadly rounded, dorsolateral margin un- and narrow, tip acute, tip to base broadly armed; lateral surface slightly rugose dorso- convex; lateral surface smooth, several small distally, many short and two longer rows of tufts of short setae in generally straight line setae ventrally; mesial surface smooth, long across medioproximal surface, several wide- plumose setae on margins and scattered on ly spaced submarginal tufts of short setae surface. Merus smooth, dorsal and ventral dorsodistally; mesial surface smooth, ventral margins unarmed, long plumose setae, dis- margin with long plumose setae, dorsal mar- tolateral margin with long plumose setae; lat- gin with short plumose setae, patch of long eral surface with decalci®ed area anteriorly; plumose setae at base (not illustrated). Pro- mesial surface smooth. Basis-ischium incom- podal dorsal surface smooth, ventral margin pletely fused and unarmed. Coxa with one in¯ated and rounded; oblique row of long small spine on anterior margin. Female with plumose setae on distal margin of lateral sur- large gonopore on median mesial surface of face; distal and ventral margins with long coxa, surrounded with short plumose setae; plumose setae; dorsolateral surface as nar- male without pore. row, oblique, ¯attened shelf, short setae on Pereopod IV (®g. 95D) dactylus with base dorsal margin and long plumose setae on to tip convex to straight, tip acute, tip to base ventral margin; mesial surface with elevated, convex distally, becoming broadly concave curved, setose ridge from ventral junction proximally; lateral surface smooth, ventral with dactylus almost to ventral proximal margin with long plumose setae, dorsal mar- junction with carpus. Carpus slightly pro- gin with short simple setae; mesial surface duced, gently rounded; lateral surface nearly with median decalci®ed area, demarcated smooth, with irregular, broken row of rugae ventrally by longitudinal elevated ridge with and submarginal elevated ridge ventrally, ru- row of long plumose setae, setose punctae gae and ridge with long plumose setae, mar- ventral to decalci®ed window. Propodus ex- gins with long plumose setae; mesial surface panded dorsally and ventrally, ventral expan- smooth, long plumose setae on margins and sion exceeds ventral margin of dactylus, mar- in scattered patches on surface. Merus with gin with long plumose setae, dorsal expan- medial decalci®ed area on lateral surface, sion with row of long plumose setae medi- long plumose setae on lateral margins; me- ally; lateral and mesial surfaces smooth. sial surface nearly smooth, with few setae. Carpus not produced dorsodistally; lateral Basis-ischium incompletely fused and un- and mesial surfaces smooth, dorsal margin armed. Coxa with one small spine on anterior with short simple and long plumose setae, margin. ventral margin with short plumose setae. Pereopod III (®g. 95C) dactylus with base Merus with scattered short, transverse rows 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 295 of setae on lateral surface, dorsal and ven- cate posteriorly, dorsal surface smooth, with trodistal margins with long plumose setae, median longitudinal groove anteriorly, row slightly rugose ventrodistally, with short se- of setose punctae lateral to midline from me- tae; mesial surface with large decalci®ed dian of longitudinal groove almost to poste- window proximoventrally. Basis-ischium in- rior margin; margins with long plumose se- completely fused and unarmed. Coxa un- tae. armed. DISTRIBUTION: Madagascar, Zanzibar, and Abdomen (®g. 95E) with somite I approx- the Seychelles; also from Indonesia and imately as wide as long, widest posteriorly; Queensland, Australia, in 9.1±34.0 m depth. dorsal surface with anterior margin concave, MAXIMUM SIZE: Males: 11.3 mm cl; fe- posterior margin concave, with submarginal males 8.1 mm cl. row of short setae, small transverse decalci- TYPE SPECIMENS: MNHN-Hi 202 (holo- ®ed submedial windows. Somite II dorsal type), MNHN-Hi 203 (allotype), MNHN-Hi surface with submarginal transverse ridge an- 204 (paratype), MNHN-Hi 205 (paratype), teriorly, tuft of setae at posterolateral angle, ANSP CA4644 (paratype), ANSP CA4645 extending onto pleura posteromesially, pos- (paratype), RMNH 23703 (paratype). terior margin with indistinct punctate sub- TYPE LOCALITY: Environs de Nosy Be, marginal groove laterally; pleura expanded CoÃte southwest, Madagascar, 13Њ37.7ЈS, and directed slightly anteriorly, margins ®ne- 47Њ49.6ЈE, 25 m. ly toothed, lateral margins rounded, anterior REMARKS: As with all older records of Al- and lateral margins with long plumose setae, bunea species lacking descriptive informa- posterior margin with short setae. Somite III tion or illustrations, it is dif®cult to be certain similar to somite II, but narrower, shorter, what species Ward (1942) was dealing with and lacking anterior submarginal ridge, small from Mauritius. I have examined no speci- tuft of short thick setae on posterolateral an- mens from that locality, and the only species gle; pleura thinner and shorter than on somite known from nearby ReÂunion is A. speciosa, II, directed anterolaterally, with setae as in which no doubt Ward (1942) would have rec- somite II, anterolateral angle acute, dorsal ognized as quite different from A. symmysta. surface obliquely ¯attened anterolaterally. Albunea holthuisi is the best candidate for Somite IV similar to somite III, but thinner Ward's (1942) record, as it super®cially re- and shorter; dorsal surface with thick setae sembles A. symmysta and is known from posterolaterally; pleura thinner and shorter nearby Madagascar and the Seychelles. The than on somite III, directed laterally, dorsal other taxa known from the vicinity of Maur- surface obliquely ¯attened anterolaterally, itius (A. elioti and A. microps) are unlikely margins with long plumose setae. Somite V to have been confused with A. symmysta by wider than somite IV, lateral margins with an experienced carcinologist such as Ward. plumose setae; pleura absent. Somite VI sub- As noted by Boyko and Harvey (1999), equal to somite V in width but longer, dorsal the record of ``Albunea steinitzi'' from Mad- surface with short oblique rows of setae lat- agascar given by Thomassin (1969) is not erad of midline anteriorly, lateral margins that species. This record is tentatively placed with long plumose setae; pleura absent. in synonymy with A. holthuisi, which is also Females with uniramous, paired pleopods known from Madagascar, even though cer- on somites II±V, males lacking pleopods. tain morphological details shown in Tho- Telson of male (®g. 95F, G) spatulate, massin's (1969) drawings do not ®t perfectly truncate posteriorly, weakly calci®ed, except with this species. Thomassin's (1969) draw- for large triangular anterior plate, margins ing of the habitus has a produced distal mar- with long plumose setae; median longitudinal gin of the (presumably male) telson, while groove very short, restricted to anterior of the indent of the dactylus of pereopod II is calci®ed plate, calci®ed plate with thick el- too broad. These may, however, be due to the evated medial ridge (®g. 95G) covered with relatively poor quality of Thomassin's (1969) short thick simple setae, small tuft of setae drawings (see under A. speciosa). Thomas- at anterolateral margin. Telson of female (®g. sin's (1969) record may represent an unde- 95H) ovate, longer than wide, slightly trun- scribed species, but as his material is un- 296 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 available for examination (see under A. spe- Philippines: Busuanga, Palawan, May ciosa), no de®nite conclusions can be made 10±30, 1963, coll. J. E. Norton and F. E. at this time. Dayrit: 1 (, 6.4 mm cl, paratype (NMCR Although Boyko and Harvey (1999) indi- 1151b); Calapan, Mindoro, Dec. 1932±Jan. cated that this species is most similar to A. 1933, coll. P. de Mesa: 1 &, 7.9 mm cl, para- steinitzi, that relationship was based on the type (MCZ 9625). species described at that time. In fact, A. hol- Singapore: Singapore, coll. unknown: 1 thuisi typi®es its own group of Albunea spe- (, 9.6 mm cl, paratype (BMNH 1937.6.1.8); cies, and it is actually the sister taxon to A. Morib Beach, Selengar, March 8, 1993, coll. groeningi. It is also closely related to A. mar- A. Sasekumar: 1 oviger, 10.7 mm cl, para- quisiana. type (ZRC 1995.571). Malaysia: Bedok, Feb. 7, 1959, coll. P. Albunea groeningi, new species Yeoh: 1 (, 10.8 mm cl (ZRC 2000.1799). Figures 96, 97 Vietnam: Sta. 51, off Nhatrang, 15 m, Jan. 14, 1960, coll. R/V ``Gallardo'': 1 (, 3.8 mm Albunea symnista [sic]: Gordon, 1938: 187 (part)*. ± Miyake, 1978: 152±154, ®gs. 59, cl (ZMUC 2717); Sta. 215, off Nhatrang, 16 60a* (not Albunea symmysta (Linnaeus, 1758)). m, March 7, 1960, coll. R/V ``Gallardo'': 1 Albunea symmysta: Miyake et al., 1962: 125 (part; (, 3.0 mm cl (ZMUC 2718); Sta. 54, off Toyama Bay material) (not Albunea symmysta Nhatrang, 14 m, Jan. 14, 1960, coll. R/V (Linnaeus, 1758)). ``Gallardo'': 1 &, 3.7 mm cl (ZMUC 2719); Albunea steinitzi: SereÁne and Umali, 1965: 97± Sta. 220, off Nhatrang, 16 m, March 7, 1960, 102, pl. 1, ®g. 2, pl. 2, ®g. 2, pl. 3, ®gs. 3±4b, coll. R/V ``Gallardo'': 1 unsexable, unmea- pl. 4, ®g. 2, text-®gs. 1b, 2b, 4b, 5c, 6c, cЈ, 7a, surable specimen (ZMUC 2721). 9b, c*. ± Haig, 1974: 447 (list) (not Albunea Australia: Western Australia: ``North- steinitzi Holthuis, 1958). west Australia,'' coll. unknown: 1 (, 6.0 mm ?Albunea symnista [sic]: Kikuchi, 1932: 10 (? not Albunea symmysta (Linnaeus, 1758)). cl, paratype (BMNH 1932.11.30.65); Rose- mary Island, Dampier Archipelago, 7 fms (ϭ MATERIAL EXAMINED: Japan: Mikawa-Is- 12.8 m), Aug. 22, 1963, coll. F.R.V. Lance- shiki, Aichi Prefecture, Honshu, Sept. 1, lin: 3 (, 9.2±11.4 mm cl, 2 &, 10.6±11.3 mm 1941, coll. T. Sakai: 1 (, 9.4 mm cl, holo- cl, paratypes (WAM 23386); Bernier Island, type (RMNH ex 32052), 1 &, 12.5 mm cl, Shark Bay, May 16, 1960, coll. R. W. allotype (RMNH 32052); Sakai, Tottori Pre- George: 1 &, 13.4 mm cl, paratype (WAM fecture, March 1, 1964, coll. T. Senda: 1 &, 23391); Shark Bay, Feb. 1963, coll. L. R. 16.4 mm cl (ZLKU 12836); Fushiki, Toyama Thomas: 1 (, 12.9 mm cl, paratype (WAM Bay, Aug. 6±26, 1950, coll. Hori and Mori: 23395); North Island, 17±25 fms (ϭ 31.1± 1 (, 8.7 mm cl, 1 &, 7.0 mm cl, paratypes 45.7 m), May 1965, coll. R. Seabrook: 1 (, (ZLKU 5125±5126); Mimase, Kochi City, 12.9 mm cl, paratype (WAM 23396); Mis- Tosa Bay, Shikoku Island, Dec. 24, 1959, sion Bay, Napier Broome Bay, 1±3 m, Aug. coll. K. Sakai: 1 &, 17.0 mm cl, paratype 14, 1991, coll. L. Wharsh: 2 (, 6.4±8.3 mm (ZLKU 7438); Tosa Bay, Shikoku Island, cl, 1 &, 8.1 mm cl, paratype (WAM 20667); Feb. 19, 1960, coll. S. Nakayama: 1 (, 11.6 Queensland: Ball Bay, May 24, 1964, coll. mm cl, paratype (ZLKU 7511). Zoology Department of the University of Taiwan: ``Formosa,'' coll. M. Maki: 1 &, Queensland: 2 (, 10.5±12.4 mm cl, 4 &, 13.8 mm cl, paratype (USNM 59074); Kao- 7.8±15.4 mm cl, paratypes (WAM 23389); hsiung, 25 m, June 7, 1992, coll. unknown: Rockhampton, Keppel Bay, 23Њ10ЈS, 2 (, 9.2±9.8 mm cl, 1 &, 11.2 mm cl, para- 150Њ55ЈE, 9 m, Sept. 6, 1967, coll. B. J. types (NTOU); ``Southern Taiwan,'' 30 m, Smith: 1 unsexable, unmeasurable specimen Jan. 1, 1992, coll. unknown: 1 (, 8.8 mm cl, (MOV J47044); Lucinda, Hinchinbrook 1 &, 9.2 mm cl, paratypes (NTOU); Tung- Channel, 18Њ29ЈS, 146Њ16ЈE, Oct. 1969, coll. Shiao, Miao±Li County, Aug. 30, 1999, coll. E. Gossner: 1 (, 11.0 mm cl, paratype unknown: 1 &, 8.4 mm cl (NTOU); Tan- (MOV J14552); Rudder Reef, 30 mi north- Shui, Taipei County, June 20, 1999, coll. un- east of Mossman, 16Њ11ЈS, 145Њ40ЈE, Oct. known: 1 (, 4.1 mm cl (NTOU). 1973, coll. R. J. Plant: 1 (, 8.5 mm cl, para- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 297 type (MOV J44728); Rockhampton, Keppel erally with 8±11 large spines (n ϭ 6) on each Bay, 23Њ10ЈS, 150Њ55ЈE, 9 m, Sept. 6, 1967, side along length. Rostrum as small acute coll. B. J. Smith: 1 (, 14.1 mm cl, paratype tooth, reaching proximal margin of ocular (MOV J44732); Dunk Island, 17Њ57ЈS, plate. Ocular sinus smoothly concave. Fron- 146Њ09ЈE, coll. E. J. Ban®eld: 2 (, 8.9±9.2 tal region smooth; setal ®eld narrow anteri- mm cl, paratypes (AM P5340); off North orly and posteriorly; posterior lateral ele- Head, Port Denison, 20Њ01ЈS, 148Њ15ЈE, coll. ments reduced to narrow bands of setae. CG1 E. H. Rainford: 1 (, 14.4 mm cl, paratype parallel to anterior margin of carapace, con- (AM P7029); Queen's Beach, Bowen, coll. vex, sinuous, strongly crenulate, divided into E. N. Rainworth: 1 (, 9.1 mm cl, paratype medial fragment and curved, posteriorly dis- (QM W143); Kinkuna National Park, 20 km placed lateral elements. Mesogastric region south of Bundaberg, 25Њ00ЈS, 152Њ30ЈE, smooth; CG2 present as none to two short April 1992, coll. M. Hines: 1 unsexable spec- elements; CG3 broken into four widely sep- imen, 12.5 mm cl (QM W18006); Weipa, arated short elements between posterior lat- July 1961, coll. G. Webster: 1 (, 9.1 mm cl, eral elements of CG1; CG4 with two to six 1 &, 9.6 mm cl, paratypes (QM W2221); short, oblique medial elements (rarely ab- Bowen, Feb. 27, 1934, coll. unknown: 1 (, sent) between longer supralateral elements of 11.6 mm cl, paratype (QM W489); Towns- CG4. Hepatic region smooth, with oblique ville, Oct. 7, 1938, coll. G. Coates: 1 &, 10.5 setose groove at median of lateral margin. mm cl, paratype (QM W873); Victoria: Mud Epibranchial region generally triangular, Island, Port Phillip Bay, April 1977, coll. R. smooth; posterolateral margin with single Willington: 1 &, 13.9 mm cl, paratype (MOV oblique row of short setae. Metagastric re- J40194). gion smooth; CG5 present as two short trans- DIAGNOSIS: Carapace wider than long, cov- verse elements. CG6 strongly crenulate, ered with lightly setose grooves. Anterior strongly anteriorly concave medially and margin with 8±11 spines on either side of sloping out to anteriorly convex lateral ocular sinus. Setal ®eld with narrow lateral thirds. CG7 almost transverse, not reaching elements and concave anterior margin. CG1 lateral margins of median segment of CG6. with separate posterior lateral elements; CG4 Cardiac region smooth; CG8 present as two with two to six short, oblique medial ele- to four short medial elements and two longer ments (rarely absent) between longer supra- lateral elements. CG9 present as two to four lateral elements of CG4; CG5 present as two short lateral grooves with broad gap at mid- short transverse elements; CG6 and CG7 line. CG10 present as two long lateral ele- separate; CG8 broken; CG11 absent. Ros- ments. CG11 absent. Post-CG11 element ab- trum present, reaching posterior margin of sent. Branchial region with three to six short, ocular plate. Ocular plate triangular. Distal oblique rows of setae in anterior half. Pos- peduncular segments dorsoventrally ¯attened terior margin deeply and evenly convex, with and subtriangular in shape, tapering at tip, submarginal groove reaching two-thirds up approximated at base of mesial margins, lat- margin of posterior concavity. Branchioste- eral margins convex except slightly concave gite with short anterior submarginal spine; at tip, mesial margins straight. Cornea at tip. anterior region with scattered, short, trans- Dactylus of pereopod II with heel produced verse lines ventral to linea anomurica; with and acute. Dactylus of pereopod III with heel many short rows of setae and sparsely cov- thin, produced and acute. Dactylus of pereo- ered with long plumose setae ventrally; pos- pod IV with produced, subquadrate heel. Tel- terior region membranous, with numerous ir- son of male elongate, ovate, tip rounded with regular fragments and sparsely covered with medial indentation, dorsal surface in¯ated long plumose setae. medially, with medial row of long thickened Ocular plate (®g. 96B) triangular, with setae; lateral margins decalci®ed. Telson of shallow median indentation; median pedun- female similar to male, evenly calci®ed. cular segments present as small ovate calci- DESCRIPTION: Carapace (®g. 96A) wider ®ed areas lateral to ocular plate. Distal pe- than long. Anterior margin concave on either duncular segments elongate, subtriangular, side of ocular sinus, becoming convex lat- with convex lateral and straight mesial mar- 298 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Fig. 96. Albunea groeningi, n. sp.: A±J, (, 9.2 mm cl, AM P5340, paratype. A. Carapace, bran- chiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Right maxilliped III, lateral view. Scale ϭ 1.1 mm (B), 1.6 mm (E, F, I), 2.2 mm (C, D, H), and 3.3 mm (A, G, J). 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 299 gins, cornea covering lateral portion of distal three-segmented, with plumose setae on mar- tip; lateral margin with faint notch one-fourth gins and long, thick, simple setae arising distal from base; mesial margins approxi- from bend in second segment and on distal mated at base; mesial margins with long plu- margin of terminal segment. mose setae; tuft of plumose setae at proxi- Maxillule (®g. 96F) distal endite proxi- molateral ventral angles and ventromedial mally narrow, widening to in¯ated distal end, oblique row of plumose setae extending from with thick simple setae on distal margin and tuft to three-fourths length of segment. thin simple setae on dorsal margin. Proximal Antennule (®g. 96C) with segment III nar- endite with thick simple setae on distal mar- row proximally, expanding distally to three gin. Endopodal external lobe truncate distally times proximal width; plumose setae on dor- and curled under; internal lobe reduced with sal and ventral margins and sparsely scat- two thick setae at distolateral margin. tered on lateral surface; dorsal exopodal ¯a- Maxilla (®g. 96G) exopod evenly rounded, gellum with 104±127 articles (n ϭ 6), long with plumose setae along distal margin. Sca- plumose setae on dorsal and ventral margins; phognathite bluntly angled on posterior lobe, ventral endopodal ¯agellum with two articles with plumose setae. (n ϭ 6), plumose setae on dorsal and ventral Maxilliped I (®g. 96H) epipod with plu- margins. Segment II medially in¯ated in dor- mose setae on margins, distolateral surface, sal view, with plumose setae on dorsal and and mesial surface. Endite tapered distally ventral margins. Segment I wider than long, and subequal to ®rst segment of exopod. Ex- unarmed; dorsal third of lateral surface ru- opod with two segments: proximal segment gose with long plumose setae; long plumose narrow, margins parallel, with plumose setae; setae on dorsal and ventral margins. distal segment spatulate, longer than wide, Antenna (®g. 96D) with segment V ap- broadest medially, margins and mesioventral proximately two times longer than wide, with surface with long plumose setae. Endopod long plumose setae on dorsal and ventral ¯attened and elongate, reaching two-thirds to margins and scattered on lateral surface; ¯a- distal end of proximal exopodal segment; gellum with seven articles (n ϭ 6), long plu- plumose setae on margins and median of lat- mose setae on dorsal, ventral, and distal mar- eral surface. gins. Segment IV expanded distally, with Maxilliped II (®g. 96I) dactylus evenly long plumose setae on dorsal, ventral, and rounded, length subequal to width, with thick distal margins, and row of short setae on dor- simple setae distally and on distolateral sur- solateral surface. Segment III with long plu- face. Propodus two times wider than long, mose setae on ventral margin; short simple slightly produced at dorsodistal angle, with setae on dorsal margin and in short row on plumose setae on dorsal margin and patch of surface. Segment II short, widening distally, long simple setae on dorsodistal and ven- rugose, with plumose setae on margins and trodistal angles. Carpus not produced dorso- short simple setae scattered on lateral sur- distally, approximately two times longer than face; antennal acicle long, thin, and exceed- wide; long simple setae on dorsal margin. ing distal margin of segment IV by one-third Merus approximately three times longer than length of segment IV, with long plumose se- wide, margins parallel; with simple and plu- tae on dorsal margin. Segment I rounded mose setae on margins. Basis-ischium in- proximally, ¯attened ventrolaterally, with completely fused with plumose setae on mar- long plumose setae on dorsal and distoven- gins. Exopod one-half longer than merus, ¯a- tral margins, and short simple setae in short gellum with one elongate article, approxi- row on surface rugae behind spine; lateral mately as long as carpus. surface with acute spine dorsodistally; low Maxilliped III (®g. 96J) dactylus oblong semicircular dorsolateral lobe ventrodistal to with rounded tip; long plumose setae on mar- spine, margin of lobe with long plumose se- gins and lateral surface. Propodus dorsodis- tae; segment with ventromesial antennal tally in¯ated, with longitudinal median row gland pore. of plumose setae on lateral surface; margins Mandible (®g. 96E) incisor process with with plumose setae. Carpus produced onto three teeth; cutting edge with one tooth. Palp propodus approximately one-third length of 300 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Fig. 97. Albunea groeningi, n. sp.: A±F, (, 9.2 mm cl, AM P5340, paratype; G, &, 13.9 mm cl, MOV J40194, paratype. A. Left pereopod I, lateral view. B. Left pereopod II, lateral view. C. Left pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 1.7 mm (F), 2.2 mm (G), and 3.3 mm (A±E). propodus; lateral surface with two rows of ing, approximately one-third length of mer- plumose setae; plumose setae on margins. us; with plumose setae on margins; without Merus in¯ated, unarmed, with plumose setae ¯agellum. on margins and few scattered small areas on Pereopod I (®g. 97A) dactylus curved and lateral surface. Basis-ischium incompletely tapering; lateral and mesial surfaces smooth; fused, with weak crista dentata of one or two dorsal margin with long plumose setae; ven- teeth. Exopod two-segmented: proximal seg- tral margin with short simple setae. Propodal ment small; distal segment styliform, taper- lateral surface with numerous short, trans- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 301 verse rows of setose rugae; dorsal margin un- large median decalci®ed window covering armed; ventral margin distally produced into nearly all of lateral surface, long plumose se- acute spine; cutting edge lacking teeth, lined tae on dorsodistal and ventral margins, few with long plumose setae; dorsal margin with scattered long plumose setae on surface; me- long plumose setae, ventral margin with sial surface nearly smooth, with two long short simple setae. Carpus with dorsodistal rows of setae. Basis-ischium incompletely angle produced into strong corneous-tipped fused and unarmed. Coxa unarmed. spine; dorsal margin with short transverse Pereopod III (®g. 97C) dactylus with base grooves behind spine; dorsal and distal mar- to heel deeply concave, heel narrow and gins with long plumose setae; lateral surface acutely produced, heel to tip with broadly with small distal rugose area, few transverse concave indent and small concave region at setose ridges on distal half of surface; mesial midpoint of proximal margin, tip acute, tip surface smooth, with medial transverse row to base smoothly convex; lateral surface of setae, margins with long plumose setae. smooth, with several small tufts of short se- Merus unarmed; lateral surface with scat- tae in generally straight line across medi- tered transverse rows of long plumose setae, oproximal surface, dorsodistal margin with distal margin with long plumose setae; me- tufts of short setae; ventral margin with long sial surface with few scattered setae; fully plumose setae, dorsal margin with short sim- calci®ed. Basis-ischium incompletely fused, ple and plumose setae; mesial surface unarmed. Coxa unarmed. smooth, with plumose setae proximally at Pereopod II (®g. 97B) dactylus smooth; junction with propodus. Propodus not in¯at- base to heel strongly and smoothly concave, ed dorsoventrally; lateral surface smooth, heel produced, broad and subacute, heel to with long plumose setae in oblique row, sim- tip with narrow, acute indent, tip acute, tip ple setae on dorsal margin, plumose setae on to base broadly convex; lateral surface ventral margin; dorsolateral surface narrow, smooth, with several small tufts of short se- oblique, ¯attened, with sparse setose mat; tae in generally straight line across medi- mesial surface smooth. Carpus produced dor- oproximal surface, several widely spaced sodistally, exceeding proximal margin of submarginal tufts of short setae dorsodistally; propodus by one-third length of propodus; mesial surface smooth, ventral margin with tip subacute, dorsolateral margin unarmed; long plumose setae, dorsal margin with short lateral surface slightly rugose dorsodistally, simple setae, with patch of long plumose se- with mat of short setae and two interrupted tae at base. Propodal dorsal surface smooth, rows of setae ventrally; mesial surface with ventral margin in¯ated and rounded; smooth, with long plumose setae on distal oblique row of long plumose setae on distal margin and in oblique row on surface. Merus margin of lateral surface; distal and ventral smooth, with large decalci®ed window cov- margins with long plumose setae; dorsolat- ering nearly half of lateral surface medially; eral surface as narrow, oblique, ¯attened dorsal and ventral margins unarmed, dorso- shelf, with short setae on dorsal margin and distal and ventrodistal margins with long plu- long plumose setae on ventral margin; mesial mose setae; mesial surface smooth. Basis-is- surface with elevated, curved, setose ridge chium incompletely fused and unarmed. from ventral junction with dactylus almost to Coxa unarmed. Female with large gonopore ventral proximal junction with carpus. Car- on anterior mesial margin of coxa, surround- pus strongly produced and rounded dorsodis- ed with short plumose setae; male without tally, dorsal margin smooth; lateral surface pore. smooth, produced area smooth, irregular, in- Pereopod IV (®g. 97D) dactylus with base terrupted row of rugae and submarginal ele- to tip straight proximally, with subquadrate vated ridge ventrally, rugae and ridge with heel and broadly rounded, strongly concave long plumose setae; dorsodistal margin with indent and almost straight from indent to tip, long plumose setae, proximodorsal and ven- tip acute, tip to base concave distally to con- tral margins with short plumose setae; mesial vex proximally; lateral surface smooth, ven- surface smooth, with row of long plumose tral margin with long plumose setae, dorsal setae distally and subdorsally. Merus with margin with short simple setae; mesial sur- 302 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 face with dorsal decalci®ed region, demar- obliquely ¯attened anterolaterally; lateral and cated ventrally by longitudinal elevated ridge posterior margins with long plumose setae, with row of short setae; with setose puncta- anterior margin with short simple setae. So- tions ventral to decalci®ed window. Propo- mite V wider than somite IV, anterior sub- dus expanded dorsally and ventrally; ventral marginal windows present; lateral margins expansion reaching ventral margin of dacty- with plumose setae; pleura absent. Somite VI lus, ventral margin with long plumose setae; slightly broader than somite V, anterior sub- dorsal expansion with row of long plumose marginal windows present; dorsal surface setae dorsally, oblique area with mat of short with short transverse rows of setae laterad of simple setae; lateral surface smooth, mesial midline and on posterior margin; pleura ab- surface smooth, with distoventral area of few sent. patches of long plumose setae. Carpus slight- Females with uniramous, paired pleopods ly produced dorsodistally; ventral four-®fths on somites II±V; males without pleopods. of lateral surface and mesial surface smooth, Telson of male (®g. 97F) ovate and elon- dorsodistal one-®fth of lateral surface with gated, length greater than width, distal tip mat of short setae; dorsal margin with short rounded, with median indentation; thickly simple and long plumose setae; ventral mar- calci®ed medially, in¯ated dorsally; distal gin with short simple setae; mesial surface two-thirds with lateral decalci®ed region; decalci®ed medially. Merus with scattered median longitudinal groove extending one- short transverse rows of setae on lateral sur- half length, row of long, thick, simple setae face, dorsal margin with long plumose setae; of either side of median groove beginning at proximoventral half of mesial surface with median and continuing to distal margin of large decalci®ed window. Basis-ischium in- calci®ed area; proximolateral angles with completely fused and unarmed. Coxa un- long simple setae; margins with long simple armed. setae. Telson of female (®g. 97G) similar to Abdomen (®g. 97E) somite I wider than male, but more truncate distally and with long, widest posteriorly; dorsal surface with smaller median indentation, dorsal surface anterior margin straight; posterior margin evenly calci®ed; median groove similar to curved with elevated submarginal row of male, but with shorter setae; proximolateral short setae; small transverse decalci®ed win- angle with few short simple setae, margins dows laterad of segment median. Somite II with long simple setae. dorsal surface with irregular, submarginal, DISTRIBUTION: From southern Japan south- transverse ridge anteriorly; with small trans- ward in a narrow band to Western Australia verse, decalci®ed windows laterad of seg- and Victoria, Australia, in up to 45.7 m ment median just anterior to submarginal depth. ridge; pleura expanded and directed antero- MAXIMUM SIZE: Males: 14.4 mm cl; fe- laterally; anterolateral margins angled, ante- males: 17.0 mm cl. rior and lateral margins with long plumose TYPE SPECIMENS: RMNH ex 32052 (holo- setae, posterolateral angle rounded, posterior type), RMNH 32052 (allotype); AM P5340 margin with short setae; posteromesial angle (2 paratypes), AM P7029 (paratype), with mat of short simple setae. Somite III BMNH,1932.11.30.65 (paratype), BMNH similar to somite II, but narrower, shorter, an- 1937.6.1.8 (paratype), MCZ 9625 (paratype), terior submarginal windows present; pleura MOV J14552 (paratype), MOV J40194 thinner and shorter than on somite II, direct- (paratype), MOV J44728 (paratype), MOV ed posterolaterally proximally and anterolat- J44732 (paratype), NMCR 1151b (paratype), erally distally, with setae as in somite II; an- NTOU (3 paratypes), NTOU (2 paratypes), terolateral angle subacute; dorsal surface QM W143 (paratype), QM W489 (paratype), obliquely ¯attened anterolaterally, with sub- QM W873 (paratype), QM W2221 (2 para- marginal row of short setae. Somite IV sim- types), USNM 59074 (paratype), WAM ilar to somite III, but thinner and shorter, an- 20667 (3 paratypes), WAM 23386 (5 para- terior submarginal windows present; pleura types), WAM 23389 (6 paratypes), WAM thinner and shorter than on somite III, di- 23391 (paratype), WAM 23395 (paratype), rected posterolaterally; dorsal surface WAM 23396 (paratype), ZLKU 5125±5126 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 303

(2 paratypes), ZLKU 7438 (paratype), ZLKU (part). ± Gmelin and LinneÂ, 1790: 2994. ± Grif- 7511 (paratype), ZRC 1995.571 (paratype). ®th and Pidgeon, 1833: 178. TYPE LOCALITY: Mikawa-Isshiki, Aichi Cancer dorsipes: Herbst, 1791: 5±8, pl. 22, ®g. 2. Prefecture, Honshu, Japan. ± Herbst, 1796: 197±198, pl. 45, ®gs. 1±7 (not Notopus dorsipes (Fabricius, 1793)). ETYMOLOGY: This species is named after Albunea symnista [sic]: Weber, 1795: 94. ± Fa- Matt Groening, cartoonist and creator of the bricius, 1798: 397. ± Herbst, 1804: 31 (list). ± television program ``The Simpsons,'' to hon- Latreille, 1806: 44. ± Lamarck, 1818: 224. ± or his extensive promotion of crustacean is- Desmarest, 1823: 283, unnumbered pl., ®g. 3. sues in the popular media. ``The Simpsons'' ± Desmarest, 1825: 173, pl. 29, ®g. 3. ± GueÂrin has exposed people to a diversity of crusta- MeÂneville, 1829: 12 (list), pl. 15, ®g. 1. ± La- cean species, including Lisa's hermit crab, treille, 1831: 56. ± Brewster, 1832: 234. ± H. Patty and Selma's hermit crab cleaning tech- Milne Edwards, 1837a: 111±112, explanation inques, and, of course, ``Pinchy'' the lobster. for pl. 42, pl. 42, ®g. 3. ± H. Milne Edwards, 1837b: 203±204, pl. 42, ®g. 3. ± H. Milne Ed- The speci®c name is pronounced ``gray-ning- wards, 1840: 111±112. ± White, 1847: 57. ± de i.'' Haan, 1849: viii, pl. Q. ± Lucas, 1853: 45±47, REMARKS: This species is more common pl. 1, ®g. 8. ± Heller, 1863: 153 (part). ± Heller, in the Philippines than indicated by the above 1865: 72. ± Chenu and Desmarest, 1877: 32. ± material examined. SereÁne and Umali (1965) Miers, 1878: 326±327. ± Lucas, 1881: 54±55*. cited 54 specimens from various Philippine ± Albert, 1883: 523±524, pl. 31, ®gs. 1, 6, 17. locations (as A. steinitzi). It is possible that ± de Man, 1887: 425. ± F. MuÈller, 1890: 472. ± some of SereÁne and Umali's (1965) material Ortmann, 1892: 536. ± Henderson, 1893: 338, is not this species, but the identi®cation of 409*. ± Stebbing, 1893: 152 (list). ± Southwell, 1910: 183. ± Boschma, 1931: 351, 354*. ± their material is based on their descriptions Boschma, 1937: 204*. ± Menon, 1937: 10±15, and illustrations and a single specimen cited ®gs. 46±82. ± Gordon, 1938: 187 (part), ®gs. by them that was available for examination 1e, 3f, 4c*. ± Bouvier, 1940: 181, pl. 6, ®g. 1. (NMCR 1151b). Curiously, SereÁne and Uma- ± Gravely, 1941: 75, 104, ®g. 27±1. ± Gurney, li (1965) stated that all of their specimens 1942: 263, ®g. 110k. ± Boschma, 1955: 14, 57, were males. However, their ``female'' of A. 65, 67*. ± Sarojini, 1962: 191, ®g. 1j. ± SereÁne thurstoni (pl. 5, ®g. 1a) is clearly a male, so and Umali, 1965: 90±95 (part), pl. 1, ®g. 1, pl. they may have been in error regarding other 2, ®g. 1, pl. 3, ®g. 1, 1a, pl. 4, ®g. 1, text-®gs. specimens as well. The record of Kikuchi 1a, 2a, 3, 4a, 5a, b, 6a, b, bЈ*. ± Thomassin, 1969: 138±140, pl. 1, text-®g. 3a. ± Boonruang (1932) from Toyama Bay, Japan, is probably and Phasuk, 1975: 1±2, 8, 15, 17, ®g. 4. ± An- this species, the only Albunea known with antaraman and Subramoniam, 1976: 192±199. certainty from that location. ± Naiyanetr, 1978: 333, ®g. 1. ± Naiyanetr, One & specimen from Kaoasiung, Taiwan 1980: 22. ± Panneerselvam and Subramoniam, (NTOU) is host to an undescribed species of 1983: 1±8, ®gs. 1±8. ± Subramoniam, 1984: Albunione Markham and Boyko (Isopoda: 78±94, ®gs. 1a, 2±14. ± CoeÃlho and Calado, Bopyridae). 1987: 43, table 1. ± Seridji, 1988: 1298. ± Sub- This species belongs to the ``holthuisi- ramoniam, 1993: 133, 152±155, 158, 197±198, group'' of species and is the sister-taxon to ®gs. 14, 15. ± Calado, 1997a: 17. ± Naiyanetr, 1998: 51. ± K. Sakai, 1999: 9, pl. 1, ®g. d. A. holthuisi. albunea symnista [sic]: Latreille, 1803: 172, pl. 51, ®g. 4. Albunea symmysta (Linnaeus, 1758) Albunea dorsipes: Herbst, 1804: 31 (list) (not No- Figures 98, 99 topus dorsipes (Fabricius, 1793)). Cancer lymnista [sic]: Froriep, 1806: 183 (list). Cancer Symmysta Linnaeus, 1758: 630. ± Linnae- Albunea Symniste [sic]: DumeÂril, 1816: 431. us, 1764: 453. Albunaea [sic] symnista [sic]: Stimpson, 1858: Cancer symnista [sic]: Linnaeus, 1767: 1053. 230 (list). Cancer Gymnista [sic]: Houttuyn, 1769: 422. Albunea (symnista): Claus, 1885: 69, 108, pl. 6, Cancer gymnista [sic]: Statius MuÈller, 1775: 1128. ®g. 52. ± Claus, 1886: 69, 108, pl. 6, ®g. 52. Hippa symnista [sic]: Fabricius, 1787: 329 (part). Albunea symmysta: Ortmann, 1896: 224. ± Ort- ± Fabricius, 1793: 474. mann, 1901: 1276, pl. 72, ®g. 3, pl. 93, ®g. 4, Cancer Symnista [sic]: de Villers, 1789: 157±158 pl. 104, ®g. 2. ± Holthuis, 1956: 238. ± ICZN, 304 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

1958: 234. ± Subramoniam and Panneerselvam, not Albunea symnista [sic]: Rathbun, 1924: 29 (ϭ 1985: 226±227. ± Boyko and Harvey, 1999: Albunea sp. indet.). 391, 396, 400 (list), 401 (key)*. ± Boyko, 1999: not Albunea symnista [sic]: Nakazawa, 1927: 145 (list). 1051, ®g. 2025. ± Nakazawa et al., 1949: 741, Cancer symmysta: Holthuis, 1956: 237±238. ± ®g. 2144. ± Nakazawa et al., 1951: 741, ®g. ICZN, 1958: 213, 215, 233±234. ± Melville and 2144 (ϭ? Paralbunea dayriti (SereÁne and Uma- Smith, 1987: 298. Albunea Symnista [sic]: Ser- li, 1965)). idji, 1988: 1298. not Albunea symnista [sic]: Kikuchi, 1932: 10 (ϭ Albunea symnysta [sic]: Chace and Kensley, Albunea? groeningi, n. sp.). 1992: 446, ®g. 2n. not Albunea symnista [sic]: T. Sakai, 1935: 61 (ϭ Cancer symmista [sic]: Calado, 1995: 71. Albunea sp. indet.). ``Albunea sp. n.`` Calado, 1995: 76±78, pl. 4, ®g. not Albunea symnista [sic]: Gordon, 1938: 187 l, pl. 5, ®g. l, pl. 23, ®g. a, pl. 24, ®gs. a±e*. (part)*. ± SereÁne and Umali, 1965: 90±95 Albunea symnestra [sic]: Dexter, 1996: 12. (part)*. ± Miyake, 1991: 157, pl. 53, ®g. 2. ± Albunea symmista [sic]: Fransen et al., 1997: Asakura, 1995: 376, ®g. 21±285 (ϭ Albunea 161*. ± Richmond, 1997: 214, unnumberef ®g. occultus, n. sp.). on p. 215. not Albunea symnista [sic]: Gordon, 1938: 187 Albunea edsoni Calado, 1997a: 18±21, ®gs. 1, 2* (part)*. ± Miyake, 1978: 152±154, ®gs. 59, (NEW SYNONYMY). 60a* (ϭ Albunea groeningi, n. sp.). ?Albunea oxyophthalma: Southwell, 1910: 184 not Albunea symnista [sic]: Gordon, 1938: 187 (not Albunea oxyophthalma Miers, 1878 ϭ A. (part)* (ϭ Albunea bulla, n. sp.). paretii GueÂrin MeÂneville, 1853). not A. symnista [sic]: Ward, 1942: 52 (list), 63 ?AIbunea [sic] symnista [sic]: Menon, 1937: 10. (ϭ? Albunea holthuisi Boyko and Harvey, ?Albunea [sp.] Menon, 1943: 331±332. 1999). ?Albunea symnista [sic]: Wang, 1989: 39. ± Sun not Albunea symnista [sic]: Barnard, 1950: 405± and Wang, 1996: 31 (list) (? ϭ Albunea sp. in- 406. ± Kensley, 1981: 35 (list) (ϭ Albunea sp. det.). indet.). not Hippa symnista [sic]: Fabricius, 1787: 329 not Albunea symnista [sic]: Kikuchi, 1959: 49 (part) (ϭ Corystes cassivelaunus (Pennant, (list). ± Kikuchi, 1961: 5 (list) (ϭ Albunea sp. 1777)). indet.). not Cancer Symnista [sic]: de Villers, 1789: 157± not Albunca [sic] symnista [sic]: Kikuchi, 1961: 5 158 (part) (ϭ Corystes cassivelaunus (Pennant, (list) (ϭ Albunea sp. indet.). 1777)). not Albunea symmysta: Miyake, 1961: 12.ÐMi- not Albunea symnista [sic]: Ra®nesque-Schmaltz, yake et al., 1962: 125 (part) (ϭ? Paralbunea 1814: 20. ± Lucas, 1849a: 27±28*. ± Lucas, dayriti (SereÁne and Umali, 1965)). 1849b: pl. 3, ®g. 2*. ± Heller, 1863: 153 (part). not Albunea symmysta: Miyake et al., 1962: 125 ± Barrois, 1888: 18±19, 75, 82, 89, 93±94. ± (part, Toyama Bay material) (ϭ Albunea groen- Bolivar, 1892: 128 (list). ± Ferrer Aledo, 1914: ingi, n. sp.). 68. ± Miranda y Rivera, 1933a: 22. ± Miranda not Albunea symmista [sic]: Miyake, 1965: 651* y Rivera, 1933b: 1 (list) (ϭ Albunea carabus (ϭ Albunea occultus, n. sp.). (Linnaeus, 1758)). not Albunea symmista [sic]: Miyake, 1965: ®g. not Albunea symnista [sic]: BrulleÂ, 1836±1844: 1111 (ϭ? Paralbunea dayriti (SereÁne and Uma- 17. ± Ozorio, 1888: 186 (ϭ Albunea elegans A. li, 1965)). Milne Edwards and Bouvier, 1898). not Albunea symnista [sic]: SereÁne, 1977: 47. ± not Albunea symnista [sic]: Gibbes, 1850a: 24* Calado, 1995: 71±73, pl. 4, ®g. i, pl. 5, ®g. h, (ϭ Albunea paretii GueÂrin MeÂneville, 1853). pl. 21, ®gs. a, b, pl. 22, ®gs. a±f* (ϭ Albunea not Albunea symnista [sic]: Gibbes, 1850b: 187 microps Miers, 1878). (ϭ Albunea catherinae, n. sp.). not Albunea symmysta: Kikuchi and Miyake, not Albunea symnista [sic]: A. Milne Edwards, 1978: 31 (list) (ϭ Albunea sp. indet.) 1862: F±12* (ϭ Albunea speciosa Dana, 1852). not Albunea symmysta: Nurul Huda et al., 1989: not Albanea [sic] symnista [sic]: Hoffman, 1874: 88±89 (ϭ Albunea? thurstoni Henderson, 42 (list) (ϭ Albunea speciosa Dana, 1852). 1893). not Albunea symnista [sic]: Cano, 1889a: 95, 104. ± Cano, 1889b: 263. (ϭ Albunea steinitzi Hol- MATERIAL EXAMINED: India: ``Indian thuis, 1958). Seas,'' coll. unknown: 1 (, 13.6 mm cl, 2 not Albunea symmysta: Nobili, 1906: 142±143*. &, 19.0±19.8 mm cl (BMNH 1883.25); ± Ramadan, 1936: 3 (list) (ϭ Albunea steinitzi ``Ganjam,'' coll. unknown: 1 &, 15.0 mm cl Holthuis, 1958). (BMNH 1889.6.17.151); Colachel, coll. un- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 305 known: 1 (, 14.1 mm cl (BMNH Thailand: Phuket Island, July 1970, coll. 1903.4.6.210); Madras, coll. J. R. Hender- unknown: 1 &, 19.4 mm cl (CASIZ 109242). son: 1 (, 16.2 mm cl, 7 &, 14.2±21.9 mm Philippines: Padada Beach, Gulf of Da- cl (BMNH 1892.7.15.125±132); Madras, vao, Sept. 1939, coll. G. R. Oesch: 1 oviger, coll. unknown: 3 &, 17.2±19.6 mm cl 12.9 mm cl (AMNH 10393); Morong, Ba- (BMNH 1903.4.6.211±212); [Madras], coll. taan, March 1, 1999, coll. J. D. Williams: 2 unknown, 1 larva, 2.8 mm cl (BMNH (, 8.9±12.6 mm cl, 1 juvenile, 5.2 mm cl 1949.12.2.544 ex Menon via Gurney); Ma- (AMNH 18092); Morong, Bataan, Feb. 20, dras, coll. Museum Wien: 1 (, 14.8 mm cl, 1999, coll. J. D. Williams: 1 molt, 7.1 mm neotype (ZMO F17538); Pondicherry, 1881, cl (AMNH 18093); Levgayen Gulf, 500 ft (ϭ coll. M. Maindron: 1 &, 18.0 mm cl 151.5 m), May 1909, coll. ``Albatross'': 1 &, (MNHN-Hi 115); Pondicherry, 1881, coll. 14.8 mm cl (USNM 68613); Calapan, Min- M. Maindron: 3 (, 12.5±13.4 mm cl doro, Jan. 6, 1933, coll. P. de Mesa: 1 (, 12.1 (MNHN-Hi 116); Pondicherry, Jan. 1881, mm cl (MCZ 19654); Calapan, Mindoro, coll. M. Maindron: 2 &, 15.3±18.0 mm cl coll. P. de Mesa: 1 (, 8.2 mm cl (MCZ (MNHN-Hi 16); Pondicherry, Nov. 28, 1942, 9615). coll. unknown: 1 &, 19.5 mm cl (MNHN-Hi Australia: Queensland:15Њ28ЈS, 15); Pondicherry, coll. M. Leschenault: 1 (, 145Њ15ЈE, Cooktown, June 1, 1985, coll. S. 17.4 mm cl, 1 &, 17.8 mm cl (MNHN-Hi W. Gunn: 1 (, 9.7 mm cl (MOV J44730); 182); Pondicherry, coll. M. Leschenault: 1 Cardwell, Port Douglas, Jan. 20, 1972, coll. &, 14.7 mm cl (MNHN-Hi 183); Pondicher- E. M. Grant: 1 &, 15.1 mm cl (QM W3848); ry, coll. M. Leschenault: 1 &, 16.4 mm cl New South Wales: Lord Howe Island, coll. (MNHN-Hi 184); Pondicherry, coll. M. Les- unknown: 1 &, 17.8 mm cl, holotype of A. chenault: 1 (, 13.0 mm cl (MNHN-Hi 185); edsoni (BMNH 1912.11.22.106). [Pondicherry], coll. M. Leschenault: 1 (, Limited Data: Sta. 1679, Mission de , 14.1 mm cl (MNHN-Hi 186); Pondicherry, l'Indochine, 1912, coll. A. Kreuff: 4 ( 12.2±13.2 mm cl, 2 &, 13.6±15.5 mm cl Aug. 1901, coll. M. Maindron: 25 (, 10.6± (MNHN-Hi 86); ``Asia,'' pre±1866, coll. L. 15.5 mm cl, 16 &, 11.0±20.8 mm cl de Jeude: 1 &, 18.7 mm cl (MOV J14551); (MNHN-Hi 196); Pondicherry, Bay of Ben- ``Asia,'' coll. L. de Jeude: 1 (, 11.9 mm cl gal, 11 59 N, 79 50 E, 1901, coll. [M. Maid- Њ Ј Њ Ј (MOV J44726); ``Yanaon, Indes Orientales,'' ron]: 1 , 14.6 mm cl (AM P5404); Pondi- ( coll. unknown: 1 &, 18.6 mm cl (MNHN Hi cherry, 1901, coll. [M. Maidron]: 1 (, 15.5 187). mm cl (AM P5405 ex MNHN); Pondicherry, No Data:1(, 12.4 mm cl (MNHN-Hi 1901, coll. [M. Maidron]: 1 (, 13.0 mm cl 17); 1 &, 17.0 mm cl (MOV J14554). (AM P5406 ex MNHN). DIAGNOSIS: Carapace wider than long, cov- Sri Lanka (Ceylon): ``Ceylon,'' coll. E. ered with lightly setose grooves. Anterior E. Green: 2 (, 15.3±15.9 mm cl, 1 &, 16.2 margin with 11±17 spines on either side of mm cl (BMNH 1904.11.28.4±6); ``Beutkofe, ocular sinus. Setal ®eld with narrow lateral SW Ceylon,'' Aug. 2, 1909, coll. G. DuÈnker: elements and concave anterior margin. CG1 1 &, 17.7 mm cl (ZMH K±32584). with separate posterior lateral elements; CG4 Indonesia: Padang, Sumatra, Oct. 14, with two short, anteriorly displaced medial 1828±Nov. 10, 1829, coll. S. MuÈller: 2 &, elements between longer supralateral ele- 15.0±16.0 mm cl (RMNH 23029); Benkoe- ments of CG4; CG5 of two triangular ele- len, Sumatra, Nov. 1925, coll. H. C. Kellers: ments; CG6 and CG7 separate; CG8 broken; 1 &, 19.1 mm cl (USNM 68615); ``Buiten- CG11 absent. Rostrum present, not reaching zorg,'' Java, June 1906, coll. H. Jensen: 16 posterior margin of ocular plate. Ocular plate (, 10.2±14.1 mm cl, 24 &, 11.2±19.2 mm cl triangular. Distal peduncular segments dor- (ZMUC 2713); ``Moluccas,'' 1841±1843, soventrally ¯attened and triangular, tapering coll. E. A. Forsten: 1 (, 15.5 mm cl, 1 &, at tip, approximated along mesial margins at 13.5 mm cl (RMNH 23030); ``Moluccas,'' base, lateral margins broadly convex, mesial coll. unknown: 8ϩ unsexable specimens, margins straight. Cornea at tip. Dactylus of 12.5±15.3 mm cl (RMNH 853). pereopod II with heel produced, tapered, and 306 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 subacute. Dactylus of pereopod III with heel short, transverse lines ventral to linea ano- tapering, projecting, acute. Dactylus of pe- murica; with many short rows of setae and reopod IV sinuous from base to tip, with pro- sparsely covered with long plumose setae duced subacute heel and deep indent. Telson ventrally; posterior region membranous, with of male broadly triangular, laterally pro- numerous irregular fragments and sparsely duced, length subequal to width, with broad- covered with long plumose setae. ly rounded tip; thickly calci®ed medially, in- Ocular plate (®g. 98B) triangular, with ¯ated dorsally; distal two-thirds with lateral deep median indentation; median peduncular decalci®ed region, median row of thin setae. segments present as small ovate calci®ed ar- Telson of female ¯attened, ovate, and evenly eas lateral to ocular plate. Distal peduncular calci®ed with slightly produced tip. segments elongate, subtriangular, with DESCRIPTION: Carapace (®g. 98A) slightly strongly convex lateral and straight mesial wider than long. Anterior margin slightly margins, cornea covering lateral portion of concave on either side of ocular sinus, be- distal tip; lateral margins with notch one- coming convex laterally, with 11±17 large fourth distal from base; mesial margins ap- spines (n ϭ 9) along length. Rostrum as proximated at base, with long plumose setae; small acute tooth, not reaching proximal tuft of plumose setae at proximolateral ven- margin of ocular plate. Ocular sinus smooth- tral angles and ventromedial oblique row of ly concave, with three or four small spinules. plumose setae extending from tuft to three- Frontal region smooth; setal ®eld narrow an- fourths length of segment. teriorly and posteriorly; posterior lateral el- Antennule (®g. 98C) with segment III nar- ements reduced to narrow bands of setae. row proximally, expanding distally to two CG1 parallel to anterior margin of carapace, times proximal width; with plumose setae on sinuous, strongly crenulate, divided into me- dorsal and ventral margins and sparsely scat- dial fragment and curved, posteriorly dis- tered on lateral surface; dorsal exopodal ¯a- placed lateral elements. Mesogastric region gellum with 121±131 articles (n ϭ 6), long smooth; CG2 absent; CG3 broken into ®ve plumose setae on dorsal and ventral margins; or six short elements between posterior lat- ventral endopodal ¯agellum with three to eral elements of CG1, medial elements an- ®ve articles (n ϭ 6), plumose setae on dorsal teriorly displaced; CG4 with two short, an- and ventral margins. Segment II medially in- teriorly displaced medial elements between ¯ated in dorsal view, with plumose setae on longer supralateral elements of CG4. Hepatic dorsal and ventral margins and scattered on region smooth, with oblique setose groove at ventrolateral third of surface. Segment I wid- median of lateral margin. Epibranchial re- er than long, unarmed; dorsal third of lateral gion generally triangular, smooth; posterolat- surface rugose with long plumose setae; long eral margin without rows of setae. Metagas- plumose setae on dorsal and ventral margins. tric region smooth; CG5 present as two tri- Antenna (®g. 98D) with segment V ap- angular elements. CG6 strongly crenulate, proximately two times longer than wide, with strongly anteriorly concave medially and long plumose setae on dorsal and ventral sloping out to anteriorly convex lateral margins and scattered on lateral surface; ¯a- thirds. CG7 oblique, not reaching lateral gellum with seven articles (n ϭ 6), long plu- margins of median segment of CG6. Cardiac mose setae on dorsal, ventral, and distal mar- region smooth; CG8 present as two to four gins. Segment IV expanded distally, with short medial elements. CG9 present as two long plumose setae on dorsal, ventral, and short lateral grooves with gap at midline. distal margins and row of setae on dorsolat- CG10 absent. CG11 absent. Post-CG11 ele- eral surface. Segment III with long plumose ment absent. Branchial region with numerous setae on dorsal and ventral margin and in short, transverse rows of setae in anterior short row on surface. Segment II short, wid- half. Posterior margin deeply and evenly ening distally, rugose, with plumose setae on convex, with submarginal groove reaching margins and scattered on lateral surface; an- two-thirds up margin of posterior concavity. tennal acicle long, thin, and exceeding distal Branchiostegite with short anterior submar- margin of segment IV by one-sixth length of ginal spine; anterior region with scattered, segment IV, with long plumose setae on dor- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 307

Fig. 98. Albunea symmysta (Linnaeus, 1758): A, &, 14.8 mm cl, USNM 68613; B±J, &, 16.9 mm cl, ZMUC 2713. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 2.2 mm (F), 3.0 mm (B, E), 3.3 mm (I), 4.4 mm (D, H), 5.0 mm (A), 5.9 mm (C), and 6.7 mm (G, J). 308 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 sal margin. Segment I rounded proximally, pod one-third longer than merus, ¯agellum ¯attened ventrolaterally, with long plumose with one elongate article, approximately as setae on margins and in short row on surface long as carpus. rugae behind spine; lateral surface with acute Maxilliped III (®g. 98J) dactylus oblong spine dorsodistally, with low semicircular with rounded tip; long plumose setae on mar- dorsolateral lobe ventrodistal to spine; seg- gins and lateral surface. Propodus dorsodis- ment with ventromesial antennal gland pore. tally in¯ated, with longitudinal median row Mandible (®g. 98E) incisor process with of plumose setae on lateral surface; margins three teeth; cutting edge with one tooth. Palp with plumose setae. Carpus produced onto three-segmented, with plumose setae on mar- propodus almost one-third length of propo- gins and long, thick, simple setae arising dus; lateral surface with two rows of plumose from bend in second segment and on distal setae on surface; plumose setae on margins. margin of terminal segment. Merus in¯ated, unarmed, with plumose setae Maxillule (®g. 98F) distal endite proxi- on margins and scattered on lateral surface. mally narrow, widening to in¯ated distal end, Basis-ischium incompletely fused, with weak with thick simple setae on distal margin and crista dentata of three or four teeth. Exopod thin simple setae on dorsal margin. Proximal two-segmented: proximal segment small; endite with thick simple setae on distal mar- distal segment styliform, tapering, approxi- gin. Endopodal external lobe truncate distally mately two-®fths length of merus; with plu- and curled under; internal lobe reduced with mose setae on margins; without ¯agellum. four thick setae at distolateral margin. Pereopod I (®g. 99A) dactylus curved and Maxilla (®g. 98G) exopod evenly rounded, tapering; lateral and mesial surfaces smooth; with plumose setae along distal margin. Sca- dorsal margin with long plumose setae; ven- phognathite bluntly angled on posterior lobe, tral margin with short simple setae. Propodal with plumose setae. lateral surface with numerous short, trans- Maxilliped I (®g. 98H) epipod with plu- verse rows of setose rugae; dorsal margin un- mose setae on margins, distolateral surface, armed; ventral margin distally produced into and mesial surface. Endite tapered distally acute spine; cutting edge lacking teeth, lined and subequal to ®rst segment of exopod. Ex- with long plumose setae; dorsal margin with opod with two segments; proximal segment long plumose setae, ventral margin with narrow, margins parallel with plumose setae; short simple setae. Carpus with dorsodistal distal segment spatulate, longer than wide, angle produced into strong corneous-tipped broadest medially, margins and mesioventral spine; dorsal margin with short transverse surface with long plumose setae. Endopod grooves behind spine; dorsal and distal mar- ¯attened and elongate, reaching two-thirds to gins with long plumose setae; lateral surface distal end of proximal exopodal segment; with small distal rugose area, few transverse plumose setae on margins and median of lat- setose ridges on distal half of surface; mesial eral surface. surface smooth, with medial transverse row Maxilliped II (®g. 98I) dactylus evenly of setae, margins with long plumose setae. rounded, length slightly greater than width, Merus unarmed; lateral surface with scat- with thick simple setae distally and on dis- tered transverse rows of long plumose setae, tolateral surface. Propodus 1.5 times wider margins with long plumose setae; mesial side than long, slightly produced at dorsodistal with few scattered setae; fully calci®ed. Ba- angle, with plumose setae on dorsal margin sis-ischium incompletely fused, unarmed. and patch of long simple setae on dorsodistal Coxa unarmed. and ventrodistal angles. Carpus not produced Pereopod II (®g. 99B) dactylus smooth; dorsodistally, approximately two times lon- base to heel slightly concave, heel produced, ger than wide; long simple setae on dorsal broad and subacute, heel to tip with narrow, margin. Merus approximately three times acute indent, tip acute, tip to base broadly longer than wide, margins parallel; with sim- convex; lateral surface smooth, with several ple and plumose setae on margins and scat- small tufts of short setae in generally straight tered on surface. Basis-ischium incompletely line across medioproximal surface, several fused with plumose setae on margins. Exo- widely spaced submarginal tufts of short se- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 309

Fig. 99. Albunea symmysta (Linnaeus, 1758): A±E, G, &, 16.9 mm cl, ZMUC 2713; F, (, 14.1 mm cl, ZMUC 2713. A. Right pereopod I, lateral view. B. Left pereopod II, lateral view. C. Left pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 3.3 mm (F, G) and 5.9 mm (A±E). tae dorsodistally; mesial surface smooth, dorsal margin and long plumose setae on ventral margin with long plumose setae, dor- ventral margin; mesial surface with elevated, sal margin with short simple setae, patch of curved setose ridge from ventral junction long plumose setae at base. Propodal dorsal with dactylus almost to ventral proximal surface smooth, with ventral margin in¯ated junction with carpus. Carpus strongly pro- and rounded; oblique row of long plumose duced and rounded dorsodistally, dorsal mar- setae on distal margin of lateral surface; dis- gin smooth; lateral surface smooth, produced tal and ventral margins with long plumose area smooth, irregular, interrupted row of ru- setae; dorsolateral surface as narrow, gae and submarginal elevated ridge ventrally, oblique, ¯attened shelf, with short setae on rugae and ridge with long plumose setae; 310 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 margins with short plumose setae; mesial long plumose setae, dorsal margin with short surface smooth, with row of long plumose simple setae; mesial surface with dorsal de- setae distally and subdorsally. Merus with calci®ed region, demarcated ventrally by lon- large median decalci®ed window covering gitudinal elevated ridge with row of short se- nearly all of lateral surface, with few scat- tae; with setose punctations ventral to decal- tered long plumose setae on surface and mar- ci®ed window. Propodus expanded dorsally gins; mesial surface nearly smooth, with two and ventrally; ventral expansion reaching long rows of setae. Basis-ischium incom- ventral margin of dactylus, ventral margin pletely fused and unarmed. Coxa unarmed. with long plumose setae; dorsal expansion Pereopod III (®g. 99C) dactylus with base with row of long plumose setae dorsally, to heel concave, heel narrow and acutely pro- oblique area with mat of short simple setae; duced, heel to tip with broadly concave in- lateral surface smooth, mesial surface dent and small concave region at midpoint of smooth, with distoventral area of few patches proximal margin, tip acute, tip to base of long plumose setae. Carpus slightly pro- smoothly convex; lateral surface smooth, duced dorsodistally; ventral four-®fths of lat- with several small tufts of short setae in eral surface and mesial surface smooth, dor- roughly straight line across medioproximal sodistal ®fth of lateral surface with mat of surface, dorsodistal margin with tufts of short short setae; dorsal margin with short simple setae; ventral margin with long plumose se- and long plumose setae; ventral margin with tae, dorsal margin with short simple and plu- short simple setae; mesial surface decalci®ed mose setae; mesial surface smooth, with plu- medially. Merus with scattered, short, trans- mose setae proximally at junction with pro- verse rows of setae on lateral surface, dorsal podus. Propodus not in¯ated dorsoventrally; and ventrodistal margins with long plumose lateral surface smooth, with long plumose se- setae; proximoventral half of mesial surface tae in oblique row, simple setae on dorsal with large decalci®ed window. Basis-ischium margin, plumose setae on ventral margin; incompletely fused and unarmed. Coxa un- dorsolateral surface narrow, oblique, ¯at- armed. tened, with setose mat; mesial surface Abdomen (®g. 99E) somite I wider than smooth. Carpus produced dorsodistally, ex- long, widest posteriorly; dorsal surface with ceeding proximal margin of propodus by anterior margin straight; posterior margin one-third length of propodus; tip rounded, curved, with elevated submarginal row of dorsolateral margin unarmed; lateral surface short setae; small transverse decalci®ed win- slightly rugose dorsodistally, with mat of dows laterad of segment median. Somite II short setae and two interrupted rows of setae dorsal surface with irregular submarginal ventrally; mesial surface smooth, with long transverse ridge anteriorly; with small trans- plumose setae on distal margin and in verse decalci®ed windows laterad of segment oblique row on surface. Merus smooth, with median just anterior to submarginal ridge; large decalci®ed window covering nearly pleura expanded and directed anterolaterally; half of lateral surface medially; dorsal and anterolateral margins angled, anterior and ventral margins unarmed, with long plumose lateral margins with long plumose setae, pos- setae; distolateral margin with long plumose terolateral angle rounded, posterior margin setae; mesial surface smooth. Basis-ischium with short setae; posteromesial angle with incompletely fused and unarmed. Coxa un- mat of short simple setae. Somite III similar armed. Female with large gonopore on an- to somite II, but narrower, shorter, anterior terior mesial margin of coxa, surrounded submarginal windows present; pleura thinner with short plumose setae; male with tiny pore and shorter than on somite II, directed an- located more mesially. terolaterally, with setae as in somite II; an- Pereopod IV (®g. 99D) dactylus with base terolateral angle subacute; dorsal surface to tip convex proximally, with broadly obliquely ¯attened anterolaterally, with sub- rounded, strongly concave indent and almost marginal row of short setae. Somite IV sim- straight from indent to tip, tip acute, tip to ilar to somite III, but thinner and shorter, an- base concave distally to convex proximally; terior submarginal windows present; dorsal lateral surface smooth, ventral margin with surface with few short setae anterolaterally; 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 311 pleura thinner and shorter than on somite III, ology'' (ICZN, 1958) and is the type of the directed anterolaterally; dorsal surface genus Albunea by subsequent designation of obliquely ¯attened anterolaterally; margins Holthuis (1956). This species is called ``Jak- with long plumose setae. Somite V wider alan quai'' or ``buffalo cicada'' in Thailand, than somite IV, anterior submarginal win- terms also used for the genus Hippa (Boon- dows present; lateral margins with plumose ruang and Phasuk, 1975). setae; pleura absent. Somite VI slightly Although Linnaeus (1758, 1767) gave the broader than somite V, anterior submarginal type locality as ``Asia,'' it should be noted windows present; dorsal surface with short that he also (1764) cited India as the ``habi- transverse rows of setae laterad of midline; tat'' for this species, which is most likely pleura absent. where his specimens came from. Conse- Females with uniramous, paired pleopods quently, the neotype specimen (ZMO on somites II±V; males without pleopods. F17538) is selected from an Indian locality. Telson of male (®g. 99F) broadly trian- A neotype is needed to stabilize the identity gular, laterally produced, length subequal to of this species, as there have been numerous width, with broadly rounded tip; thickly cal- instances of confusion between other albu- ci®ed medially, in¯ated dorsally; distal two- neid taxa and A. symmysta (see synonymy thirds with lateral decalci®ed region; median list above). The so-called holotype of Cancer longitudinal groove extending one-half dorsipes ``Herbst'' (ZMB Herbst 2231), as length, row of long simple setae of either cited by K. Sakai (1999), is merely Herbst's side of median groove beginning at median (1804) specimen of A. symmysta. The species and continuing almost to distal margin of cal- dorsipes was established in the genus Hippa ci®ed area; proximolateral angles with few by Fabricius in 1793. long simple setae; margins with long simple The misspelling ``symnista'' was intro- setae. Telson of female (®g. 99G) ¯attened, duced by Linnaeus (1767) and unfortunately ovate, and evenly calci®ed with slightly pro- became the spelling used by most authors up duced tip; median groove similar to male, se- to the present. It is hoped that the correct tal row from midpoint of median groove to original spelling of this species will now be near distal margin of telson with simple setae universally adopted. subequal to those of male; proximolateral an- Albunea symmysta is a model animal that gle with few short setae, margins with long has been used to study endocrinology, sper- simple setae. matophore formation, and breeding patterns DISTRIBUTION: From the east coast of India (Panneerselvam and Subramoniam, 1983; throughout southeast Asia to the Philippines Subramoniam, 1984; Subramoniam and Pan- and Indonesia as far east as Java, in up to neerselvam, 1985). Because of this, more is 151.5 m depth. Also Queensland and Lord known about the biology of this species than Howe Island, Australia. any other albuneid, but not all published MAXIMUM SIZE: Males: 17.4 mm cl; fe- statements are accurate. As discussed for the males: 21.9 mm cl. family and genus, this species is neither blind TYPE SPECIMENS: The type or types of nor a ®lter-feeder, as suggested by Subra- Cancer symmysta are not extant and a neo- moniam and Panneerselvam (1985). type ((, 14.8 mm cl) is herein designated as This species breeds continuously along the ZMO F17538. BMNH 1912.11.22.106 (ho- Madras coast, with peaks in January and lotype of A. edsoni). June, but the maximum percentage of ovigers TYPE LOCALITIES: ``Asia'' was the only lo- is only 25% (Subramoniam and Panneersel- cality information given by Linnaeus (1758) vam, 1985). Sexual maturity of females was for A. symmysta; the neotype deisgnated reported as 17 mm cl, and maximum size as herein (ZMO F17538) was collected from 22 m cl (Subramoniam and Panneerselvam, Madras, India, which becomes the type lo- 1985). However, an ovigerous & of 12.9 mm cality of A. symmysta; Lord Howe Island, cl is known from the Philippines (AMNH New South Wales, Australia (A. edsoni). 10393). The larval stages were described by REMARKS: Albunea symmysta is no. 1565 Menon (1937) from Indian plankton samples, on the ``Of®cial list of speci®c names in zo- and there appear to be ®ve zoeal stages. 312 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

There are 12 gills along each side in the stage are probably A. occultus, but need to be ex- IV zoea and 13 in stage V zoea vs. 10 in an amined for con®rmation. adult. Menon (1943) incorrectly suggested Thomassin (1969) reported this species that this showed evidence of recapitulation of from Madagascar, but he had no specimens ancestral characters in the larval stages. and relied on the personal communication of The androgenic gland of A. symmysta,as two individuals for his information. Howev- described by Sarojini (1962), is distinctive er, the species is otherwise not known to oc- and unlike that of the pagurid hermit crab he cur in Madagascar and probably does not oc- studied, but the apparent overall variability cur west of India. Because of this and nu- in this gland within a single species makes merous other misidenti®cations in the litera- its utility in phylogenetic studies question- ture, Thomassin's (1969: text-®g. 11) able. distribution map for this species is not ac- Albunea symmysta has been well studied curate. The citation of Richmond (1997) ap- with regards to its parasites. One 12.3 mm cl pears to be based on the same incorrect data & from Java (ZMUC 2713) has an attached from Thomassin (1969) and not on actual rhizocephalan parasite, Sacculina anceps specimens from east Africa. Boschma, 1931. This parasite is one of only The identities of the dry specimens cited a few species of ``Sacculina'' not found on by Fransen et al. (1997) were con®rmed by brachyurans and may not properly belong to examination of sketches kindly supplied by this genus (but see Boschma, 1955). Saccu- Dr. Fransen. lina anceps was described from six speci- Calado (1997a) described the species Al- mens found on specimens of A. symmysta bunea edsoni, based on a single & specimen (ZMUC 2713). A lectotype for S. anceps was from Australia, but this specimen is actually selected by Boschma (1937) as ``the best pre- a large example of A. symmysta. The main served of the two specimens of which lon- diagnostic feature given by Calado (1997a) gitudinal sections have been made''. The was ``17 teeth on the right frontal edge and specimen cited above is therefore a paralec- 16 on the left.'' As shown herein, the number totype of S. anceps. High levels of infection of teeth on the frontal margin of Albunea in Indian populations of A. symmysta by mi- species is only useful for separating species crophallid trematode metacercariae (Platy- groups such as the ``microps-group'' from helminthes) were reported by Anantaraman the ``symmysta-group.'' It is not a valid and Subramoniam (1976). The parasites were means of identifying species. Although not tentatively identi®ed as Microphallus sp. and explicitly stated by Calado (1997a), the prob- were found in the ovaries of female crabs, able reason she described this new species while male crabs were uninfected. was because the specimen was not conspeci®c As A. symmysta is the only species known with the single specimen of ``A. symmysta'' at present from either Sri Lanka or India, it she examined (Calado, 1995). I have exam- seems likely that this is the taxon Southwell ined that specimen (BMNH 1956.1.14.20) (1910: 184) collected from near Ceylon, in and, as can be seen from Calado's (1995: pl. spite of his having also listed A. symmysta as 4, ®g. i, pl. 5, ®g. h, pl. 21, ®gs. a, b, pl. 22, a separate taxon (Southwell, 1910: 183). ®gs. a±f) illustrations, it is actually a mis- It is dif®cult to know what species SereÁne identi®ed A. microps. Calado (1995, 1997a) and Umali (1965) were dealing with from the apparently uncritically accepted the identi®- Philippines, but at least some of their mate- cation of this specimen on the label as writ- rial was true A. symmysta. This can be seen ten, and she incorrectly concluded that the in their illustrations of the in¯ated distal pe- Lord Howe Island specimen represented an duncular segments (text-®g. 5a) and dactyli undescribed species. This demonstrates the (text-®g. 3), which match well with A. sym- importance of studying large series of spec- mysta. Some of their material is clearly not imens in order to correctly understand intra- A. symmysta, however, as can be seen in their speci®c variation. discussion of the variable shape of the dac- This species is closely related to A. occul- tylus of pereopod IV. These other specimens tus and these two species appear to be inter- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 313 mediate in form between the ``holthuisi- 05D06BT, northwest shelf, 19Њ29.7ЈS, group'' and the ``paretii-group.'' 118Њ52.1ЈE, 38±39 m, Oct. 25, 1983, coll. CSIRO: 1 (, 12.9 mm cl, 3 &, 6.2±12.4 mm Albunea occultus, new species cl, paratypes (QM W22322); Sta. 05B07BT, Figures 100, 101 northwest shelf, 19Њ30.9ЈS, 118Њ49.2ЈE, 38± 39 m, Oct. 25, 1983, coll. CSIRO: 1 &, 17.7 Albunea symnista [sic]: Gordon, 1938: 187 mm cl, paratype (QM W22323); Sta. (part)*. ± SereÁne and Umali, 1965: 90±95 05D02BT, northwest shelf, 19Њ29.4ЈS, (part)*. ± Miyake, 1991: 157, pl. 53, ®g. 2.Ð 118Њ52.4ЈE, 37±38 m, Oct. 24, 1983, coll. Asakura, 1995: 376, ®g. 21±285 (not Albunea CSIRO: 3 , 6.1±14.0 mm cl, 3 , 7.5±9.2 symmysta (Linnaeus, 1758)). ( & Albunea symmista [sic]: Miyake, 1965: 651* (not mm cl, paratypes (QM W22326); Sta. Albunea symmysta (Linnaeus, 1758)). 04B09BT, northwest shelf, 19Њ28.4ЈS, 118Њ55.2ЈE, 39 m, Aug. 31, 1983, coll. CSI- MATERIAL EXAMINED: Japan: Tosa-Shi- RO: 1 &, 7.7 mm cl, paratype (QM mizu, Tosa Bay, Shikoku Island, 4±7 m, May W22329); Sta. 05B12BT, northwest shelf, 1959, coll. K. Kurohara: 2 &, 8.5±15.3 mm 19Њ03.6ЈS, 119Њ03.4ЈE, 82 m, Oct. 23, 1983, cl, paratypes (ZLKU 7040±7041); Tosa Bay, coll. CSIRO: 2 (, 6.3±10.3 mm cl, paratypes Shikoku Island, July 6, 1959, coll. S. Nakay- (QM W22338); Sta. 04B14S, northwest ama: 1 (, 16.7 mm cl, 1 &, 18.1 mm cl, shelf, 19Њ43.9ЈS, 117Њ54.5ЈE, 52±53 m, Sept. paratypes (ZLKU 7062±7063). 2, 1983, coll. CSIRO: 2 &, 5.6±6.2 mm cl, Taiwan: Taiwan Strait, south of Formosa paratypes (QM W22340); ``northwest Aus- Banks to the Penghu (Pescadores) Islands, tralia,'' coll. unknown: 1 &, 10.4 mm cl, 30±50 m, May 5, 1972, coll. commercial paratype (BMNH 1932.11.30.64); Northern ®shermen: 1 &, 23.0 m cl, paratype (CASIZ Territory: Western shore, Darwin Harbour, 109246); Kaohsiung, 15 m, May 10, 1991, Nov. 3, 1963, coll. unknown: 1 (, 18.6 mm coll. unknown: 1 &, 24.0 mm cl, paratype cl, paratype (WAM 23393); Waigait, Darwin, (NTOU); Kaohsiung, 20 m, April 3, 1988, Nov. 11, 1969, coll. O. J. Cameron: 2 (, coll. unknown: 1 &, 25.3 mm cl, paratype 15.1±17.5 mm cl, 1 &, 18.1 mm cl, paratype (NTOU); ``Taiwan,'' May 17, 1992, coll. un- (AM P19435); Mandora, Darwin, Nov. 22, known: 1 &, 22.3 mm cl, paratype (NTOU); 1972, coll. N. Coleman: 1 oviger, 23.7 mm ``southern Taiwan,'' 5 m, Jan. 24, 1992, coll. cl, paratype (AM P19436); Mandora, Dar- unknown: 1 (, 19.7 mm cl, paratype win, Nov. 21, 1972, coll. N. Coleman: 2 (, (NTOU). 14.0±17.5 mm cl, paratypes (AM P23809); Philippines: Mindoro, coll. P. de Mesa: 1 Dudley Point, Darwin, Sept. 17, 1970, coll. (, 10.1 mm cl, paratype (MCZ 10261); Mo- O. J. and J. Cameron: 1 &, 21.0 mm cl, para- rong, Bataan, Feb. 28, 1999, coll. J. D. Wil- type (AM P19993); sand bar no. 1, Darwin, liams: 1 juvenile, 4.0 mm cl, paratype Nov. 14, 1970, coll. O. J. and J. Cameron: 1 (AMNH 18094); Nasasa Bay, Zambales (, 19.7 mm cl, holotype (AM P20486); Province, 5±18 fms (ϭ 9.1±32.9 m), Jan. Queensland: Michaelmas Cay, Great Barrier 30&ndash&b. 8, 1960, coll. F. G. Dayrit and Reef, north of Cairns, Sept. 14, 1963, coll. J. E. Norton: 1 (, 6.7 mm cl, 1 &, 7.3 mm W. Goode: 1 &, 11.1 mm cl, paratype (WAM cl, paratypes (USNM 267779); south lagoon, 24510); Bowen, Feb. 27, 1934, coll. un- Sibutu, Sulu Archipelago, 04Њ31ЈN, 119Њ22ЈE, known: 1 (, 16.5 mm cl, paratype (QM 13 fms (ϭ 23.8 m), Feb. 25±26, 1964, coll. W25188); Mornington Island, Gulf of Car- B. R. Wilson on ``Pele'': 1 (, 8.6 mm cl, pentaria, 16Њ29ЈS, 139Њ34ЈE, coll. unknown: paratype (WAM 10411). 1 (, 9.1 mm cl, paratype (MOV J44733); off Australia: Western Australia: Coral Bay, Cape Moreton, April 1964, coll. B. Harris: 1 1989, coll. C. Simpson: 1 &, 20.2 mm cl, &, 26.7 mm cl, paratype (QM W2380); New paratype (WAM 20261); Sta. 3, northeast South Wales: Off Richmond River mouth, corner of North Pasco Island, 20Њ54ЈS, off Ballina, 28Њ52ЈS, 153Њ34ЈE, 16 m, Oct. 6, 115Њ19ЈE, Sept. 17, 1966, coll. WAM-USNM 1962, coll. unknown: 1 (, 17.3 mm cl, para- Barrow Island Expedition 1966: 1 (, 16.4 type (AM P15354); 3 mi off mouth of Clar- mm cl, paratype (WAM 23392); Sta. ence River, 15 fms (ϭ 27.4 m), late 1968, 314 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 coll. J. J. Toyer: 1 &, 22.2 mm cl, allotype ments. Mesogastric region smooth; CG2 pre- (AM P16778); within 2 km north of Clarence sent as one to three small medial elements; River bar, 8±9 fms (ϭ 14.6±16.5 m), June CG3 broken into two short medial and two 10, 1976, coll. A. E. Ford: 1 &, 25.0 mm cl, to four longer lateral elements between pos- paratype (AM P24477). terior lateral elements of CG1, medial ele- Limited Data: ``OceÂan Indien:'' 1 &, 20.6 ments anteriorly displaced; CG4 with one or mm cl (MNHN-Hi 89). two short, concave, anteriorly displaced me- DIAGNOSIS: Carapace wider than long, cov- dial elements between longer supralateral el- ered with lightly setose grooves. Anterior ements of CG4. Hepatic region smooth, with margin with 8±11 spines on either side of oblique setose groove at median of lateral ocular sinus. Setal ®eld with narrow lateral margin. Epibranchial region generally trian- elements and concave anterior margin. CG1 gular, smooth; posterolateral margin without with separate posterior lateral elements; CG4 rows of setae. Metagastric region smooth; with one or two short, concave, anteriorly CG5 present as two triangular elements. CG6 displaced medial elements between longer strongly crenulate, strongly anteriorly con- supralateral elements of CG4; CG5 of two cave medially and sloping out to anteriorly triangular elements; CG6 and CG7 separate; convex lateral thirds. CG7 oblique, not CG8 broken; CG11 present; posterior sub- reaching lateral margins of median segment marginal groove entire. Rostrum present, not of CG6. Cardiac region smooth; CG8 present reaching posterior margin of ocular plate. as one long or two short lateral elements. Ocular plate triangular. Distal peduncular CG9 present as two short lateral grooves segments dorsoventrally ¯attened and trian- with gap at midline. CG10 present as two gular, tapering at tip, approximated along short lateral elements. CG11 present as one mesial margins, lateral margins convex, me- long or two short medial elements. Post- sial margins straight. Cornea at tip. Dactylus CG11 element absent. Branchial region with of pereopod II with heel produced, tapered, numerous short, transverse rows of setae lat- and subacute. Dactylus of pereopod III with erally. Posterior margin deeply and evenly heel tapering, projecting, acute. Dactylus of convex, with submarginal groove reaching pereopod IV sinuous from base to tip, with three-fourths up margin of posterior concav- low rounded heel and shallow indent. Telson ity to entire. Branchiostegite with short an- of male broadly triangular, laterally pro- terior submarginal spine; anterior region with duced, length subequal to width, with broad- scattered, short, transverse lines ventral to ly rounded produced tip; thickly calci®ed linea anomurica; with many short rows of medially, in¯ated dorsally; distal two-thirds setae and sparsely covered with long plu- with lateral decalci®ed region, median row mose setae ventrally; posterior region mem- of thin setae. Telson of female ¯attened, branous, with numerous irregular fragments ovate, and evenly calci®ed with slightly pro- and sparsely covered with long plumose se- duced tip. tae. DESCRIPTION: Carapace (®g. 100A) slightly Ocular plate (®g. 100B) triangular with wider than long. Anterior margin slightly deep median indentation; median peduncular concave on either side of ocular sinus, be- segments present as small ovate calci®ed ar- coming convex laterally, with 8±11 large eas lateral to ocular plate. Distal peduncular spines (n ϭ 6) along length. Rostrum as segments elongate, subtriangular, with slight- small acute tooth, not reaching proximal ly convex lateral and straight mesial margins, margin of ocular plate. Ocular sinus smooth- cornea covering lateral portion of distal tip; ly concave, with one or two large spines and lateral margins with notch one-fourth distal two or three small spinules. Frontal region from base; mesial margins approximated at smooth; setal ®eld narrow anteriorly and pos- base, with long plumose setae; tuft of plu- teriorly; posterior lateral elements reduced to mose setae at proximolateral ventral angles narrow bands of setae. CG1 parallel to an- and ventromedial oblique row of plumose se- terior margin of carapace, sinuous, strongly tae extending from tuft to three-fourths crenulate, divided into medial fragment and length of segment. curved, posteriorly displaced lateral ele- Antennule (®g. 100C) with segment III 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 315

Fig. 100. Albunea occultus, n. sp.: A±J, &, 25.0 mm cl, AM P24477, paratype. A. Carapace, bran- chiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Right maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 3.3 mm (B, E, F), 4.4 mm (I), 6.7 mm (C, D, G, H, J), and 7.4 mm (A). 316 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 narrow proximally, expanding distally to 1.5 gin. Endopodal external lobe truncate distally times proximal width; with plumose setae on and curled under; internal lobe reduced with dorsal and ventral margins; dorsal exopodal six thick setae at distolateral margin. ¯agellum with 69±88 articles (n ϭ 6), long Maxilla (®g. 100G) exopod evenly round- plumose setae on dorsal and ventral margins; ed, with plumose setae along distal margin. ventral endopodal ¯agellum with two or Scaphognathite bluntly angled on posterior three articles (n ϭ 6), plumose setae on dor- lobe, with plumose setae. sal and ventral margins. Segment II medially Maxilliped I (®g. 100H) epipod with plu- in¯ated from dorsal view, with plumose setae mose setae on margins and mesial surface. on dorsal and ventral margins and scattered Endite tapered distally and subequal to ®rst on ventrolateral quarter of surface. Segment segment of exopod. Exopod with two seg- I longer than wide, unarmed; dorsal third of ments; proximal segment narrow, margins lateral surface rugose, with long plumose se- parallel, with plumose setae; distal segment tae; long plumose setae on dorsal and ventral spatulate, longer than wide, broadest medi- margins. ally, margins, proximolateral and mesiome- Antenna (®g. 100D) with segment V ap- dial surface with long plumose setae. Endo- proximately two times longer than wide, with pod ¯attened and elongate, reaching two- long plumose setae on dorsal and ventral thirds to distal end of proximal exopodal seg- margins and scattered on lateral surface; ¯a- ment; plumose setae on margins and median gellum with seven or eight articles (n ϭ 6; of lateral surface. but QM 2380 with 10 articles on one side Maxilliped II (®g. 100I) dactylus evenly and eight on other), long plumose setae on rounded, length slightly greater than width, dorsal, ventral, and distal margins. Segment with thick simple setae distally and on dis- IV expanded distally, with long plumose se- tolateral surface. Propodus two times wider tae on dorsal and ventral margins, and row than long, slightly produced at dorsodistal of short setae on dorsolateral surface. Seg- angle, with plumose setae on dorsal margin ment III with short plumose setae on dorsal and patch of long simple setae on dorsodistal margin, long plumose setae on ventral mar- and ventrodistal angles. Carpus not produced gin. Segment II short, widening distally, ru- dorsodistally, approximately two times lon- gose, with plumose setae on margins and ger than wide; long simple setae on dorsal scattered on lateral surface; antennal acicle and ventrodistal margins. Merus approxi- long, thin, and exceeding distal margin of mately three times longer than wide, margins segment IV by one-seventh length of seg- parallel; with simple and plumose setae on ment IV, with long plumose setae on dorsal margins and scattered on surface. Basis-is- margin. Segment I rounded proximally, ¯at- chium incompletely fused, with plumose se- tened ventrolaterally, with long plumose se- tae on margins. Exopod one-third longer than tae on margins, lobe, and in short row on merus, ¯agellum with one elongate article, surface rugae behind spine; lateral surface approximately as long as carpus. with acute spine dorsodistally, with low Maxilliped III (®g. 100J) dactylus oblong semicircular dorsolateral lobe ventrodistal to with rounded tip; long plumose setae on mar- spine; segment with ventromesial antennal gins and lateral surface. Propodus dorsodis- gland pore. tally in¯ated, with longitudinal median row Mandible (®g. 100E) incisor process with of plumose setae on lateral surface; margins three teeth; cutting edge smooth. Palp three- with plumose setae. Carpus produced onto segmented, with plumose setae on margins propodus almost one-third length of propo- and long, thick, simple setae arising from dus; lateral surface with two rows of plumose bend in second segment and on distal margin setae; long plumose setae on margins. Merus of terminal segment. in¯ated, unarmed, with plumose setae on Maxillule (®g. 100F) distal endite proxi- margins and scattered in short patches on lat- mally narrow, widening to in¯ated distal end, eral surface. Basis-ischium incompletely with thick simple setae on distal margin and fused, with weak crista dentata of three or thin simple setae on dorsal margin. Proximal four teeth. Exopod two-segmented: proximal endite with thick simple setae on distal mar- segment small; distal segment styliform, ta- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 317

Fig. 101. Albunea occultus, n. sp.: A±E, &, 25.0 mm cl, AM P24477, paratype; F, (, 17.5 mm cl, AM P23809, paratype; G, &, 21.0 mm cl, AM P19993, paratype. A. Left pereopod I, lateral view. B. Left pereopod II, lateral view. C. Left pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 3.3 mm (F, G), 6.7 mm (B±D), 7.4 mm (E), and 8.0 mm (A). pering, approximately one-third length of transverse rows of setose rugae; dorsal mar- merus; with plumose setae on margins; with- gin unarmed; ventral margin distally pro- out ¯agellum. duced into acute spine; cutting edge lacking Pereopod I (®g. 101A) dactylus curved teeth, lined with long plumose setae; dorsal and tapering; lateral and mesial surfaces margin with long plumose setae, ventral mar- smooth; dorsal margin with long plumose se- gin with short simple setae. Carpus with dor- tae; ventral margin with short simple setae. sodistal angle produced into strong corneous- Propodal lateral surface with numerous short, tipped spine; dorsal margin with short trans- 318 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 verse grooves behind spine; dorsal and distal completely fused and unarmed. Coxa un- margins with long plumose setae; lateral sur- armed. face with small distal rugose area, few trans- Pereopod III (®g. 101C) dactylus with verse setose ridges on distal half of surface; base to heel concave, heel narrow and acute- mesial surface smooth, with subdorsal and ly produced, heel to tip with concave area at medial transverse rows of long plumose se- base of heel and broadly concave indent, tip tae, margins with long plumose setae. Merus acute, tip to base smoothly convex; lateral unarmed; lateral surface with scattered trans- surface smooth, with several small tufts of verse rows of long plumose setae, proximo- short setae in generally straight line across dorsal margin with long plumose setae; me- medioproximal surface, dorsodistal margin sial surface with few scattered setae; fully with tufts of short setae; ventral margin with calci®ed. Basis-ischium incompletely fused, long plumose setae, dorsal margin with short unarmed. Coxa unarmed. simple and plumose setae; mesial surface Pereopod II (®g. 101B) dactylus smooth; smooth, with plumose setae proximally at base to heel slightly concave, heel produced, junction with propodus. Propodus not in¯at- broad and rounded at tip, heel to tip with ed dorsoventrally; lateral surface smooth, narrow, acute indent, tip acute, tip to base with long plumose setae in oblique row, sim- broadly convex; lateral surface smooth, with ple setae on dorsal margin, plumose setae on several small tufts of short setae in generally ventral margin; dorsolateral surface narrow, straight line across medioproximal surface, oblique, ¯attened, with setose mat; mesial several widely spaced submarginal tufts of surface smooth, with few scattered setae. short setae dorsodistally; mesial surface Carpus produced dorsodistally, exceeding smooth, ventral margin with long plumose proximal margin of propodus by one-third setae, dorsal margin with short simple setae, length of propodus; tip rounded, dorsolateral with patch of long plumose setae at base. margin unarmed; lateral surface slightly ru- gose dorsodistally, with mat of short setae Propodal dorsal surface smooth, with ventral and two interrupted rows of setae ventrally; margin in¯ated and rounded; oblique row of mesial surface smooth, with long plumose long plumose setae on distal margin of lateral setae on distal margin and in transverse row surface; distal and ventral margins with long on surface. Merus smooth, with large decal- plumose setae; dorsolateral surface as nar- ci®ed window covering nearly half of lateral row, oblique, ¯attened shelf, with short setae surface medially; dorsal and ventral margins on dorsal margin and long plumose setae on unarmed, with long plumose setae on disto- ventral margin; mesial surface with elevated, dorsal and mediolateral margins; mesial sur- curved, setose ridge from ventral junction face smooth. Basis-ischium incompletely with dactylus almost to ventral proximal fused and unarmed. Coxa unarmed. Female junction with carpus. Carpus strongly pro- with large gonopore on anterior mesial mar- duced and rounded dorsodistally, dorsal mar- gin of coxa, surrounded with short plumose gin smooth; lateral surface smooth, produced setae; male with tiny pore located more me- area smooth, irregular, interrupted row of ru- sially. gae and submarginal elevated ridge ventrally, Pereopod IV (®g. 101D) dactylus with rugae and ridge with long plumose setae; base to tip convex proximally, concave dis- dorsal margin with short plumose setae, ven- tally, tip acute, tip to base concave distally tral margin with long plumose setae; mesial to convex proximally; lateral surface smooth, surface dorsal margin with numerous low, ventral margin with long plumose setae, dor- rounded spines, surface smooth, with row of sal margin with short simple setae; mesial long plumose setae distally and in oblique surface with dorsal decalci®ed region, de- interrupted row. Merus with large median de- marcated ventrally by longitudinal elevated calci®ed window covering nearly all of lat- ridge with row of short setae; with setose eral surface, with few scattered tufts of long punctations ventral to decalci®ed window. plumose setae on surface and on distodorsal Propodus expanded dorsally and ventrally; margin; mesial surface nearly smooth, with ventral expansion reaching ventral margin of two long rows of setae. Basis-ischium in- dactylus, ventral margin with long plumose 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 319 setae; dorsal expansion with row of long plu- broader than somite V, anterior submarginal mose setae dorsally, oblique area with mat of windows present; dorsal surface with short short simple setae; lateral surface smooth, transverse rows of setae laterad of midline; mesial surface smooth, with distoventral area pleura absent. of few patches of long plumose setae. Carpus Females with uniramous, paired pleopods slightly produced and subacute dorsodistally; on somites II±V; males without pleopods. ventral four-®fths of lateral surface and me- Telson of male (®g. 101F) broadly trian- sial surface smooth, dorsodistal one-®fth of gular, laterally produced, length subequal to lateral surface with mat of short setae; dorsal width, with broadly rounded produced tip; margin with short simple and long plumose thickly calci®ed medially, in¯ated dorsally; setae; ventral margin with short simple setae; distal two-thirds with lateral decalci®ed re- mesial surface decalci®ed medially, distal gion; median longitudinal groove extending margin with long plumose setae. Merus with one-half of length, row of long simple setae scattered, short, transverse rows of short sim- of either side of median groove beginning at ple setae on lateral surface, dorsal and me- median and continuing almost to distal mar- dioventral margins with long plumose setae; gin of calci®ed area; proximolateral angles proximoventral half of mesial surface with with few long simple setae; margins with large decalci®ed window. Basis-ischium in- long simple setae. Telson of female (®g. completely fused and unarmed. Coxa un- 101G) ¯attened, ovate, and evenly calci®ed armed. with slightly produced rounded tip; median Abdomen (®g. 101E) somite I wider than groove similar to male, setal row from mid- long, widest posteriorly; dorsal surface with point of median groove to near distal margin anterior margin straight; posterior margin of telson with simple setae approximately curved with elevated submarginal row of one-half size of those of male; proximolater- short setae; small transverse decalci®ed win- al angle with few short setae, margins with dows laterad of segment median. Somite II long simple setae. dorsal surface with irregular submarginal DISTRIBUTION: From southern Japan south- transverse ridge anteriorly; with small trans- ward to the eastern and western coasts of verse decalci®ed windows laterad of segment Australia, in up to 82 m depth. median just anterior to submarginal ridge; MAXIMUM SIZE: Males: 19.7 mm cl; fe- pleura expanded and directed laterally; an- males: 26.7 mm cl. terolateral margins angled, anterior and lat- eral margins with long plumose setae, pos- TYPE SPECIMENS: AM P20486 (holotype), terolateral angle rounded, posterior margin AM P16778 (allotype), AM P15354 (para- with short setae; posteromesial angle with type), AM P19435 (3 paratypes), AM mat of short simple setae. Somite III similar P19436 (paratype), AM P19993 (paratype), to somite II, but narrower, shorter, anterior AM P23809 (2 paratypes), AM P24477 (para- submarginal windows present; pleura thinner type), AMNH 18094 (paratype), BMNH and shorter than on somite II, directed pos- 1932.11.30.64 (paratype), CASIZ 109246 terolaterally, with setae as in somite II; an- (paratype), MCZ 10261 (paratype), MOV terolateral angle subacute; dorsal surface J44733 (paratype), NTOU (paratype), NTOU obliquely ¯attened anterolaterally, with sub- (paratype), NTOU (paratype), NTOU (para- marginal row of short setae. Somite IV sim- type), QM W2380 (paratype), QM W22322 ilar to somite III, but thinner and shorter, an- (4 paratypes), QM W22323 (paratype), QM terior submarginal windows present; dorsal W22326 (6 paratypes), QM W22338 (2 para- surface with few short setae anterolaterally; types), QM W22329 (paratype), QM pleura thinner and shorter than on somite III, W22340 (2 paratypes), QM W25188 (para- directed posterolaterally; dorsal surface type), USNM 267779 (2 paratypes), WAM obliquely ¯attened anterolaterally; margins 10411 (paratype), WAM 20261 (paratype), with long plumose setae. Somite V wider WAM 23392 (paratype), WAM 23393 (para- than somite IV, anterior submarginal win- type), WAM 24510 (paratype), ZLKU 7040± dows present; lateral margins with plumose 7041 (2 paratypes), ZLKU 7062±7063 (2 setae; pleura absent. Somite VI slightly paratypes). 320 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

TYPE LOCALITY: Sand bar no. 1, Darwin, Verde Islands: La Praya, 10±30 m, 1883, Northern Territory, Australia. coll. ``Talisman'': 2 (, 12.3±13.1 mm cl, 4 ETYMOLOGY: The speci®c name of this tax- &, 13.4±18.4 mm cl, syntypes (MNHN-Hi on is derived from the Latin word meaning 7); Sta. 7.V06, 16Њ10ЈN, 22Њ57ЈW, west ``hidden,'' and refers to the confusion of this coast, Boa Vista, Aug. 27±28, 1986, coll. species with A. symmysta both in the litera- CANCAP: 3 &, 14.5±19.6 mm cl (RMNH ture and in museum collections. 38604), 2 &, 17.9±19.4 mm cl (AMNH REMARKS: The specimen illustrated by 18078 ex RMNH 38604); Plage de Mindolo, Asakura (1995) offers a rare example of a IÃle de SaÄo Vicente, May 15, 1950, coll. J. record that can be con®rmed based on a pho- Cadenat: 1 (, 12.7 mm cl (MNHN-Hi 12); tograph alone. Asakura's (1995) specimen Sta. 38, 7 m, coll.? ``Calypso'': 1 &, 12.5 clearly has an entire posterior submarginal mm cl (MNHN-Hi 181). groove on the carapace. Of the three species Senegal: Virage, Cape Vert, Dakar, April of Albunea known from Japan (A. symmysta, 12, 1982, coll. F. C. Roest: 5 (, 8.6±10.8 mm A. groeningi, A. occultus), only A. occultus cl, 2 &, 11.7±12.9 mm cl (RMNH 35926), 2 possesses this diagnostic character. (, 9.8 mm cl, 1 &, 13.8 mm cl (AMNH It is dif®cult to know what species SereÁne 18079 ex RMNH 35926). and Umali (1965) were dealing with from the Liberia: Sta. 289, Net 70B, 03Њ04Ј45ЉN, Philippines, but at least some of their mate- 16Њ62ЈW, 0±132 m, Aug. 23±24, 1927, coll. rial was probably A. occultus, based on their R/V ``Discovery'': 1 zoea? V larva, 6.0 mm comments regarding the shape of the dactyli (BMNH 1951.2.17.2355); Sta. 295, Net of pereopod IV. Their material also likely TYF, 05Њ30Ј30ЉN, 17Њ45ЈW, 2500±2700 m contained A. symmysta, but all of it needs to (see below), Aug. 25, 1927, coll. ``Discov- be examined for con®rmation. ery'': 1 zoea? V larva, 7.0 mm (BMNH This species is most closely related to A. 1951.2.17.2356). symmysta. Ghana: Manford, Gold Coast, coll. C. Hupfer: 1 (, 19.1 mm cl (ZMH K±5136). Albunea elegans Ivory Coast: Grand Bassam, 1901, coll. A. Milne Edwards and Bouvier, 1898 V. Mayet: 2 &, 14.5±18.9 mm cl (MNHN-Hi Figures 102, 103 13). Dahomey: Chenal de Cotonov, near Co- Albunea symnista [sic]: BrulleÂ, 1836±1844: 17. ± Ozorio, 1888: 186 (not Albunea symmysta (Lin- tonou, July 19, 1963, coll. H. Hoestlandt: 3 naeus, 1758)). &, 6.2±7.4 mm cl (RMNH 20957). Albunea elegans A. Milne Edwards and Bouvier, Nigeria: West Mole, Lagos, June 16, 1898: 236±238.ÐA. Milne Edwards and Bou- 1963, coll. A. R. Longhurst: 2 unsexable, un- vier, 1900: 273±275, pl. 28, ®gs. 22±25. ± Gor- measurable specimens (RMNH 20959); Sta. don, 1938: 187 (list). ± Bouvier, 1940: 181. ± 224, Lagos, May 9, 1965, coll. ``Pillsbury'': Balss, 1957: 1598. 1 (, 11.5 mm cl (RMNH 22664). Albunea carabus: Balss, 1916a: 37*. ± Monod, Gabon: Sta. 277, 01Њ44ЈS, 08Њ38ЈE, 0±88 1933: 473 (part) (not Albunea carabus (Linnae- m, Aug. 7, 1927, coll. R/V ``Discovery'': us, 1758)). 1 zoea? V larva, 5.9 mm (BMNH Albunea paretoi: Monod, 1956: 37±40, ®gs. 2±9. ± Buchanan, 1957: 56. ± Sourie, 1957: 13, 31, 1951.2.17.2354). 78. ± Forest, 1958: 147 (list). ± Gauld, 1960: Congo: Baie de Pointe-Noire, Jan. 1969, 66 (list). ± Rossignol, 1962: 124. ± Fransen, coll. unknown: 1 (, 11.9 mm cl (MNHN-Hi 1991: 56, 77 (unjusti®ed emendation) (not Al- 14). bunea paretii GueÂrin MeÂneville, 1853). Zaire: Sta. 276, Net 100B, 05Њ54ЈS, Albunea paretii: Rossignol, 1957: 97, ®g. 12. ± 11Њ19ЈE, 0±110 m, Aug. 5, 1927, coll. R/V Holthuis and Manning, 1970: 251±252*. ± Kae- ``Discovery'': 2 zoea? V larvae, 6.5±6.7 mm stner, 1980: 336 (part). ± TuÈrkay, 1982: 98, 101, (BMNH 1951.2.17.2353). 110. ± d'Udekem d'Acoz, 1999: 171 (part) (not DIAGNOSIS: Carapace wider than long, cov- Albunea paretii GueÂrin MeÂneville, 1853). ered with lightly setose grooves. Anterior ?Albunea sp. A Lebour, 1959: 129, ®g. 14. margin with 10±12 spines on either side of MATERIAL EXAMINED: Portugal: Cape ocular sinus. Setal ®eld with narrow lateral 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 321 elements and straight anterior margin. CG1 sloping out to anteriorly convex lateral with separate posterior lateral elements; CG4 thirds. CG7 oblique, reaching lateral margins with two to four short, anteriorly displaced of median segment of CG6. Cardiac region medial elements between longer supralateral smooth; CG8 present as two short medial and elements of CG4; CG5 of two oblique tri- two short lateral elements. CG9 present as angular elements; CG6 and CG7 fused; CG8 two short lateral grooves with gap at midline. broken; CG11 present. Rostrum present, not CG10 present as two short elements. CG11 reaching posterior margin of ocular plate. present as one or two short medial elements. Ocular plate triangular. Distal peduncular Post-CG11 element absent. Branchial region segments dorsoventrally ¯attened and trian- with numerous short, transverse rows of se- gular in shape, tapering at tip, approximated tae. Posterior margin deeply and evenly con- along mesial margins, lateral margins con- vex, with submarginal groove reaching two- vex, mesial margins straight. Cornea at tip. thirds up margin of posterior concavity. Dactylus of pereopod II with heel produced, Branchiostegite with strong anterior submar- tapered, and acute. Dactylus of pereopod III ginal spine; anterior region with scattered with heel broad, projecting, acute. Dactylus short, transverse lines ventral to linea ano- of pereopod IV sinuous from base to tip, with murica; with many short rows of setae and low rounded heel and shallow indent. Telson sparsely covered with long plumose setae of male broadly triangular, tip broadly round- ventrally; posterior region membranous, with ed, thickly calci®ed medially, in¯ated dor- numerous irregular fragments and sparsely sally, distal two-thirds with lateral decalci®ed covered with long plumose setae. region, median row of thin setae. Telson of Ocular plate (®g. 102B) triangular with female ¯attened, ovate, and evenly calci®ed shallow median indentation; median pedun- with slightly produced tip. cular segments present as small ovate calci- DESCRIPTION: Carapace (®g. 102A) slightly ®ed areas lateral to ocular plate. Distal pe- wider than long. Anterior margin slightly duncular segments elongate, subtriangular, concave on either side of ocular sinus, be- 0.27±0.32 length of carapace, with convex coming convex laterally with 10±12 large lateral and straight mesial margins, cornea spines (n ϭ 6) along length. Rostrum as covering distal tip; lateral margins with notch small acute tooth, not reaching proximal one-third distal from base; mesial margins margin of ocular plate. Ocular sinus smooth- approximated at base; mesial and lateral mar- ly concave, surface smooth or with one small gins with long plumose setae; tuft of plumose spinule. Frontal region smooth; setal ®eld setae at proximolateral ventral angle and narrow anteriorly, widening posteriorly; pos- ventromedial row of plumose setae extending terior lateral elements reduced to narrow from tuft to base of cornea. bands of setae. CG1 parallel to anterior mar- Antennule (®g. 102C) with segment III gin of carapace, sinuous, strongly crenulate, narrow proximally, expanding distally to divided into medial fragment and curved, twice proximal width; with plumose setae on posteriorly displaced lateral elements. Me- dorsal and ventral margins and sparsely scat- sogastric region smooth; CG2 present as two tered on lateral surface; dorsal exopodal ¯a- short medial elements; CG3 broken into six gellum with 100±120 articles (n ϭ 6), long to eight short elements between posterior lat- plumose setae on dorsal and ventral margins; eral elements of CG1; CG4 with two to four ventral endopodal ¯agellum short with two short, anteriorly displaced medial elements articles (n ϭ 6) and plumose setae on dorsal between longer supralateral elements of and ventral margins. Segment II medially in- CG4. Hepatic region smooth, with oblique ¯ated in dorsal view, with plumose setae on setose groove at median of lateral margin. dorsal and ventral margins and scattered on Epibranchial region generally triangular, ventrolateral third of surface. Segment I wid- smooth; posterolateral margin with two or er than long, with small dorsodistal spine; three tiny rows of setae. Metagastric region dorsal third of lateral surface rugose, with smooth; CG5 present as two oblique, trian- long plumose setae; long plumose setae on gular elements. CG6 strongly crenulate, dorsal and ventral margins. strongly anteriorly concave medially and Antenna (®g. 102D) with segment V ap- 322 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Fig. 102. Albunea elegans A. Milne Edwards and Bouvier, 1898: A±J, (, 10.4 mm cl, AMNH 18079. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 1.6 mm (B, E, F), 2.2 mm (I), and 3.3 mm (A, C, D, G, H, J). 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 323 proximately two times longer than wide, with plumose setae on margins and median of lat- long plumose setae on dorsal and ventral eral surface. margins and scattered on lateral surface; ¯a- Maxilliped II (®g. 102I) dactylus evenly gellum with seven articles (n ϭ 6), long plu- rounded, length slightly greater than width, mose setae on dorsal, ventral, and distal mar- with thick simple setae distally and on dis- gins. Segment IV expanded distally, with tolateral surface. Propodus 1.5 times wider long plumose setae on dorsal, ventral, and than long, slightly produced at dorsodistal distal margins, and row of setae on dorsolat- angle, with plumose setae on dorsal margin eral surface. Segment III with long plumose and patch of long simple setae on lateral sur- setae on dorsal and ventral margin and in face and ventrolateral angle. Carpus not pro- short row on surface. Segment II short, wid- duced dorsodistally, approximately two times ening distally, rugose, with plumose setae on longer than wide; long simple setae on dorsal margins and scattered on lateral surface; an- margin. Merus approximately three times tennal acicle long, thin, and exceeding distal longer than wide, margins parallel; with sim- margin of segment IV by one-third length of ple and plumose setae on margins and scat- segment IV, with long plumose setae on dor- tered on surface. Basis-ischium incompletely sal margin. Segment I rounded proximally, fused with plumose setae on margins. Exo- ¯attened ventrolaterally, with long plumose pod one-half longer than merus, ¯agellum setae on margins and scattered in line of sur- with one elongate article, approximately as face rugae behind spine; lateral surface with long as carpus is wide. acute spine dorsodistally, with low semicir- Maxilliped III (®g. 102J) dactylus with cular dorsolateral lobe ventrodistal to spine; rounded tip; long plumose setae on margins segment with ventromesial antennal gland and lateral surface. Propodus dorsodistally pore. in¯ated, with longitudinal median row of Mandible (®g. 102E) incisor process with plumose setae on lateral surface; margins two teeth; cutting edge with one tooth. Palp with plumose setae. Carpus produced onto three-segmented, with plumose setae on mar- propodus almost one-fourth length of pro- gins and long, thick, simple setae arising podus; lateral surface with two rows of plu- from bend in second segment and on distal mose setae on surface, plumose setae on margin of terminal segment. margins. Merus in¯ated, unarmed, with plu- Maxillule (®g. 102F) distal endite proxi- mose setae on margins and scattered on lat- mally narrow, widening to in¯ated distal end, eral surface. Basis-ischium incompletely with thick simple setae on distal margin and fused, with weak crista dentata of two teeth. thin simple setae on dorsal margin. Proximal Exopod two-segmented: proximal segment endite with thick simple setae on distal mar- small; distal segment styliform, tapering, ap- gin. Endopodal external lobe truncate distally proximately one-third length of merus; with and curled under; internal lobe reduced with plumose setae on margins; without ¯agellum. three thick setae at distolateral margin. Pereopod I (®g. 103A) dactylus curved Maxilla (®g. 102G) exopod evenly round- and tapering; lateral and mesial surfaces ed, with plumose setae along distal margin. smooth; dorsal margin with long plumose Scaphognathite bluntly angled on posterior and short simple setae; ventral margin with lobe, with plumose setae. short simple setae. Propodal lateral surface Maxilliped I (®g. 102H) epipod with plu- with numerous short, transverse rows of se- mose setae on margins, distolateral surface tose rugae; dorsal margin unarmed; ventral and mesial surface. Endite tapered distally margin distally produced into acute spine; and subequal to ®rst segment of exopod. Ex- cutting edge lacking teeth, lined with long opod with two segments; proximal segment plumose setae; dorsal margin with long plu- narrow, margins parallel with plumose setae; mose setae, ventral margin with short simple distal segment spatulate, longer than wide, setae. Carpus with dorsodistal angle pro- broadest medially, margins and mesioventral duced into strong corneous-tipped spine; dor- surface with long plumose setae. Endopod sal margin with few short, transverse grooves ¯attened and elongate, reaching two-thirds to behind spine; dorsal and distal margins with distal end of proximal exopodal segment; long plumose setae; lateral surface with 324 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272

Fig. 103. Albunea elegans A. Milne Edwards and Bouvier, 1898: A±F, (, 10.4 mm cl, AMNH 18079; G, &, 13.8 mm cl, AMNH 18079. A. Right pereopod I, lateral view. B. Left pereopod II, lateral view. C. Right pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I± VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 2.2 mm (F, G), 3.3 mm (B, D, E), and 4.4 mm (A, C). 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 325 small distal rugose area, few transverse se- sinuous proximal half of margin, tip acute, tose ridges on distal half of surface; mesial tip to base smoothly convex; lateral surface surface smooth, with medial transverse row smooth, with several small tufts of short se- of setae, margins with long plumose setae. tae in generally straight line across medi- Merus unarmed; lateral surface with scat- oproximal surface, dorsodistal margin with tered transverse rows of long plumose setae, tufts of short setae; ventral margin with long margins with long plumose setae; mesial sur- plumose setae, dorsal margin with short sim- face with few scattered setae; fully calci®ed. ple and plumose setae; mesial surface Basis-ischium incompletely fused, unarmed. smooth with plumose setae proximally at Coxa of male with anterior spine; coxa of junction with propodus. Propodus not in¯at- female unarmed. ed dorsoventrally; lateral surface smooth, Pereopod II (®g. 103B) dactylus smooth; with long plumose setae in oblique row, sim- base to heel concave, heel produced and ple setae on dorsal margin; dorsolateral sur- acute, heel to tip with narrow, acute indent, face narrow, oblique, ¯attened, with setose tip acute, tip to base broadly convex; lateral mat; mesial surface smooth. Carpus pro- surface smooth, with several small tufts of duced dorsodistally, only slightly exceeding short setae in generally straight line across proximal margin of propodus; dorsolateral medioproximal surface, several widely margin unarmed; lateral surface slightly ru- spaced submarginal tufts of short setae dor- gose dorsodistally, with thin mat of short se- sodistally; mesial surface smooth, ventral tae and two interrupted rows of setae ven- margin with long plumose setae, dorsal mar- trally; mesial surface smooth, with long plu- gin with short simple setae and patch of long mose setae on distal margin and in oblique plumose setae at base. Propodal dorsal sur- row on surface. Merus smooth, with large face smooth, with ventral margin in¯ated and decalci®ed window covering nearly half of rounded; oblique row of long plumose setae lateral surface medially; dorsal and ventral on distal margin of lateral surface; distal and margins unarmed but with long plumose se- ventral margins with long plumose setae; tae; distolateral margin with long plumose dorsolateral surface as narrow, oblique, ¯at- setae; mesial surface smooth. Basis-ischium tened shelf, with short setae on dorsal margin incompletely fused and unarmed. Male coxa and long plumose setae on ventral margin; with spine on anterior margin; female lack- mesial surface with elevated, curved setose ing spine. Female with large gonopore on an- ridge from ventral junction with dactylus al- terior mesial margin of coxa, surrounded most to ventral proximal junction with car- with short plumose setae; male with small pus. Carpus strongly produced and rounded pore. dorsodistally, dorsal margin smooth; lateral Pereopod IV (®g. 103D) dactylus with surface smooth, with small setose mat at tip base to tip convex proximally, concave dis- of produced area and irregular, interrupted tally, indent smoothly joined with margin, tip row of rugae and submarginal elevated ridge acute, tip to base concave distally to convex ventrally, rugae and ridge with long plumose proximally; lateral surface smooth, ventral setae; margins with short plumose setae; me- margin with long plumose setae, dorsal mar- sial surface smooth with row of long plu- gin with short simple setae; mesial surface mose setae distally and subdorsally. Merus with dorsal decalci®ed region, demarcated with large median decalci®ed window cov- ventrally by longitudinal elevated ridge with ering nearly all of lateral surface, with few row of short setae; with setose punctations scattered long plumose setae on surface and ventral to decalci®ed window. Propodus ex- margins; mesial surface nearly smooth, with panded dorsally and ventrally; ventral expan- two long rows of setae. Basis-ischium in- sion not reaching ventral margin of dactylus, completely fused and unarmed. Male coxa margin with long plumose setae; dorsal ex- with anterior margin spine; female coxa pansion with row of long plumose setae dor- without spine. sally, oblique area with mat of short simple Pereopod III (®g. 103C) dactylus with setae; lateral and mesial surfaces smooth. base to heel concave, heel acutely produced, Carpus slightly produced dorsodistally; ven- heel to tip with broadly concave indent and tral three-fourth of lateral surface and mesial 326 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 surface smooth, dorsodistal quarter of lateral ing almost to distal margin of telson; proxi- surface with mat of short setae; dorsal mar- molateral angles with patch of long simple gin with short simple and long plumose se- setae; margins with long simple setae. Telson tae; ventral margin with short simple setae; of female (®g. 103G) ¯attened, ovate, and mesial surface decalci®ed medially. Merus evenly calci®ed with slightly produced tip; with scattered short transverse rows of setae median groove similar to male, setal row on lateral surface, dorsal and ventrodistal from midpoint of median groove to near dis- margins with long plumose setae; proximov- tal margin of telson with simple setae ap- entral half of mesial surface with large de- proximately one-fourth length of those on calci®ed window. Basis-ischium incomplete- male; proximolateral angle with patch of se- ly fused and unarmed. Male coxa with spine tae, margins with long simple setae. on anterior margin; female coxa unarmed. DISTRIBUTION: Cape Verde Islands and Se- Abdomen (®g. 103E) with somite I length negal southward to Pointe-Noire, Congo, in and width subequal, widest posteriorly; dor- up to 30 m depth. sal surface with anterior margin straight; pos- MAXIMUM SIZE: Males: 19.1 mm cl; fe- terior margin curved with elevated submar- males: 19.6 mm cl. ginal row of short setae; small transverse, de- TYPE SPECIMENS: MNHN-Hi 7 (6 syn- calci®ed windows laterad of segment medi- types). an. Somite II dorsal surface with submarginal TYPE LOCALITY: La Praya, Cape Verde Is- transverse ridge anteriorly; with small trans- lands, Portugal, 10±30 m depth. verse, decalci®ed windows laterad of seg- REMARKS: A. Milne Edwards and Bouvier ment median just anterior to submarginal (1898) erroneously cited this species as hav- ridge; pleura expanded and directed antero- ing a maxilliped III with one pleurobranch, laterally; lateral margins rounded, anterior two arthrobranchs, and lacking an epipod, and lateral margins with long plumose setae, and also pereopod V having two arthro- posterior margin with short setae; postero- branchs. These authors subsequently (1900) mesial angle with mat of short simple setae. correctly stated that there are no arthro- Somite III similar to somite II, but narrower, branchs on pereopod V, but they repeated the shorter; pleura thinner and shorter than on inaccuracies regarding maxilliped III. This somite II, directed anterolaterally, with setae error became propagated through time, as as in somite II; anterolateral angle subacute; Balss (1957) repeated A. Milne Edwards and dorsal surface obliquely ¯attened anterolat- Bouvier's (1900) incorrect statements and erally. Somite IV similar to somite III, but applied them to the genus as a whole in his thinner and shorter; dorsal surface with few monumental treatise on the Decapoda. short setae anterolaterally; pleura thinner and The west African larvae reported by Le- shorter than on somite III, directed postero- bour (1959) as ``Albunea sp. A'' strongly re- laterally; dorsal surface obliquely ¯attened semble the late stage (V or VI) zoeae of A. anterolaterally; margins with long plumose catherinae, n. sp., a closely related species setae. Somite V wider than somite IV; lateral (Kurata, 1970). This suggests that these lar- margins with plumose setae; pleura absent. vae are A. elegans, rather than the sympatric, Somite VI slightly broader than somite V; but more distantly related, A. carabus. Sev- dorsal surface with short transverse rows of eral lots of larval specimens, probably stage setae laterad of midline; pleura absent. V zoeae collected by R/V ``Discovery'' from Females with uniramous, paired pleopods east African localities, are referred to this on somites II±V; males without pleopods. species as they are extremely similar to the Telson of male (®g. 103F) broadly trian- described larvae of A. paretii, another taxon gular, slightly longer than wide, with broadly closely related to A. elegans. The collection rounded tip; thickly calci®ed medially, in- depth of 2500±2700 m for a putative larvae ¯ated dorsally; distal two-thirds with lateral of this species from off Liberia (BMNH decalci®ed region; median longitudinal 1951.2.17.2356) is a suspect depth for any groove extending one-half length, row of albuneid larvae, as the deepest living adults long simple setae of either side of median are known in no more that 225 m of water. groove beginning at distal end and continu- Although this species is similar to A. par- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 327 etii, and for many years was confused with Albunea gibbesii: Benedict, 1901: 139* (not Al- that western Atlantic species, it is not as bunea gibbesii Stimpson, 1859). closely related to A. paretii, as is the Paci®c Albunea axyopthalma [sic]: Moreira, 1901: 88 A. lucasia. Albunea elegans is easily distin- (list). Albunea oxycephala [sic]: Verrill, 1901: 18±19* guished from A. catherinae, n. sp. by the (error for Albunea oxyophthalma Miers, 1878). shapes of the dactyli of pereopods III and IV. Albunea paretoi [sic]: Castro, 1967: 2. It can be separated from A. paretii by the Albunea sp. Abele, 1976: 266±267*. longer length of the branchiostegite spine, Albunea sp. Bowen et al., 1979: 253. broken CG8, and the different shape of the Albunea paretti [sic]: Rodriguez, 1980: 239 (list). dactylus of pereopod III. ?Albunea Paretii: Boas, 1880: 140, ®gs. 34, 61, 91, 119, 144, 195. ?Albunea sp. Gurney, 1924: 187, ®g. 73*. Albunea paretii GueÂrin MeÂneville, 1853 ?Albunea sp. A Gurney, 1942: 263±266, ®g. Figures 104, 105 110e±h. not Albunea paretii: Kingsley, 1880: 409±410. ± Albunea oxyophthalmus White, 1847: 57 (nomen Williams, 1965: 137±138 (part), ®gs. 112, 113*. nudum)*. ± Kurata, 1970: 182, pls. 52, 53. ± CoeÃlho and Albunea symnista [sic]: Gibbes, 1850a: 24* (not Ramos, 1972: 176 (part). ± DoÈrges, 1977: 416. Albunea symmysta (Linnaeus, 1758)). ± Young, 1978: 177. ± Kaestner, 1980: 336 Albunea Paretii GueÂrin MeÂneville, 1853: 47±48, (part). ± Wenner and Read, 1982: 188. ± Wil- pl. 1, ®g. 10*. ± Bolivar, 1875: 21. ± Seridji, liams, 1984: 249±250 (part), ®gs. 182, 183*. ± 1988: 1298. Fox and Ruppert, 1985: 259 (list). ± Martin and Albunaea [sic] oxyophthalma: Stimpson, 1858: Abele, 1986: 611, ®gs. 1b, 2d, 3b, 4b, 5b, 6b, 230 (list). 8b, 9b, c, 10b, 12b, 14b, 15b, 17b. ± Manning, Albunaea [sic] Paretii: Stimpson, 1858: 230. ± 1988: 626±628*. ± Ruppert and Fox, 1988: Stimpson, 1859: 78 (list). 250, 404, ®g. 227. ± Williams et al., 1989: 35. Albunea oxyophthalma Miers, 1878: 329±330, pl. ± Calado et al., 1990: 747 (part), ®g. 2a, b* (ϭ 5, ®gs. 14, 15*. ± Rathbun, 1897: 42*. ± Ben- Albunea catherinae, n. sp.). edict, 1901: 139. ± Verrill, 1901: 61±62, pl. 8, not Albunea oxyopthalma [sic]: Benedict. 1904: ®g. 1*. ± Verrill, 1908: 438, pl. 28, ®g. 1*. ± 625, ®g. 5* (ϭ Albunea catherinae, n. sp.). Gordon, 1938: 187 (part)*. ± Monod, 1939: not Albunea oxyophthalma: Southwell, 1910: 184 560±561, ®g. 5*. ± Bouvier, 1940: 181. ± Snod- (ϭ Albunea? symmysta (Linnaeus, 1758)). grass, 1952: ®g. 11a, d. not Albunea oxyophthalma: Gordon, 1938: 187 Albunea paretii: Miers, 1878: 330. ± Rodrigues (part), ®gs. 3d, 4d* (ϭ Albunea catherinae,n. da Costa, 1962: 6±7, pl. 1, ®gs. 5±7, pl. 3, ®gs. sp.). 1±3. ± Williams, 1965: 137±138 (part)*. ± not Albunea paretoi: Monod, 1956: 37±40, ®gs. CoeÃlho, 1966: 244. ± Fausto Filho, 1967: 12± 2±9. ± Buchanan, 1957: 56. ± Sourie, 1957: 13, 13. ± CoeÃlho and Ramos, 1972: 176 (part). ± 31, 78. ± Forest, 1958: 147 (list). ± Gauld, Gomes CorreÃa and da Silva Brum, 1980: 60. ± 1960: 66 (list). ± Rossignol, 1962: 124. ± Fran- Kaestner, 1980: 336 (part). ± Markham and sen, 1991: 56, 77 (unjusti®ed emendation) (ϭ McDermott, 1981: 1271. ± Williams, 1984: Albunea elegans A. Milne Edwards and Bou- 249±250 (part)*. ± Chace et al., 1986: 338±339, vier, 1898). ®g. 112. ± Calado, 1987: 106±118, pls. 9±12*. not Albunea paretii: Rossignol, 1957: 97, ®g. 12. ± CoeÃlho and Calado, 1987: 42±43, table 1. ± ± Holthuis and Manning, 1970: 251±252*. ± Markham, 1988: 30. ± Calado et al., 1990: 747 Kaestner, 1980: 336 (part). ± TuÈrkay, 1982: 98, (part), ®g. 2a, b*. ± HernaÂndez and BolanÄos, 101, 110. ± d'Udekem d'Acoz, 1999: 171 (part) 1995: 77. ± Calado, 1995: 51±55, pl. 4, ®g. f, (ϭ Albunea elegans A. Milne Edwards and pl. 5, ®g. e, pl. 13, ®g. a, pl. 14, ®gs. a, b, pl. Bouvier, 1898). 15, ®gs. a±e, pl. 16, ®gs. a±c*. ± Calado, 1997a: not Albunea paretti [sic]: Kurata, 1970: 180±182 17. ± Spivak, 1997: 74 (list). ± Calado, 1998: (ϭ Albunea catherinae, n. sp.). 407. ± Markham and Boyko, 1999: 4*. ± d'Udekem d'Acoz, 1999: 171 (part). ± Amaral MATERIAL EXAMINED: Bermuda: Hungry et al. in Nucci et al., 2001: 479. ± Morgado et Bay, July±Sept. 1901, coll. F. G. Gosling: 1 al. in Nucci et al., 2001: 479. (, 16.4 mm cl, 3 &, 15.9±16.8 mm cl Albunea pareti [sic]: Ortmann, 1896: 225 (list). ± (USNM 42197), 1 &, 14.5 mm cl (RMNH Ortmann, 1901: 1275. ± Moreira, 1901: 30, 88 14647 ex USNM 42197); Castle Island, Aug. (list). ± Boschi, 1981: 715, 740. 15, 1975, coll. M. L. Jones: 1 oviger, 13.3 328 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 mm cl (USNM 264760); outside Hungry coll. P. A. del Valle: 1 &, 13.9 mm cl (USNM Bay, 15±20 ft (ϭ 4.5±6.1 m), April 25, 1970, 84351). coll. J. Lightbourn: 1 (, 13.6 mm cl, 2 &, U.S. Virgin Islands: Magens Bay, St. 18.8±21.0 mm cl (USNM 267776); Paget Thomas, Aug. 8, 1971, coll. W. E. Rainey: 1 Beach, July 1901, coll. T. G. Gosling: 1 &, &, 22.4 mm cl (USNM 154546); St. Thomas, 20.0 mm cl (MCZ 19595); sandy beach be- coll. A. H. Riise: 2 &, 20.4±21.8 mm cl tween tides, Hungry Bay, Paget Parish, 1901, (ZMUC 2711); St. Thomas, coll. A. H. Riise: coll. T. G. Gosling: 1 &, 18.8 mm cl (YPM 2 &, 14.9±15.5 mm cl (ZMUC 2714); Drift 21138). Bay, Water Island, St. Thomas, July 13, USA: Florida: ``Florida,'' July 1859, coll. 1915, coll. C. R. Shoemaker: 1 (, 14.1 mm Capt. Woodbury: 1 &, 18.0 mm cl (MCZ cl (USNM 68611). 13243); south of Key West, 20 fms (ϭ 36.6 St. Maarten: Great Bay, June 24, 1955, m), April 12, 1940, coll. J. S. Schwengel: 1 coll. P. W. Hummelinck: 1 (, 17.2 mm cl (, 15.3 mm cl (ANSP 4327). (RMNH 23628). Bahamas: Sta. 62, Long Bay, April 20, St. Eustatius: Orange Baai, July 28, 1957, 1937, coll. Smithsonian-Hartford Expedition: coll. P. A. Van den Heuvel: 1 &, 24.0 mm cl 1 (, 15.1 mm cl (USNM 104655); at and (RMNH 11239); Orange Baai, Sept. 24, around tip of Pitts Town Point, northwestern 1957, coll. P. A. Van den Heuvel: 1 &, 24.9 tip of island, Crooked Island, June 1, 1962, mm cl (RMNH 11437); Zeelandia Baai, coll. J. Tyler: 1 (, 14.7 mm cl, 1 &, 16.7 Sept. 17, 1957, coll. P. A. Van den Heuvel: mm cl (USNM 260819); Andros Island, 3 &, 19.8±25.5 mm cl (RMNH 11438); Zee- March±April 1908, coll. B. E. Dahlgren and landia Baai, Jan. 4, 1958, coll. P. A. Van den H. MuÈller: 1 &, 10.8 mm cl (AMNH 5464). Heuvel: 1 &, 21.8 mm cl (RMNH 12209); Cuba: Sta. R55, Corrientes Bay, April 8, 1957, coll. P. A. Van den Heuvel: 1 (, 16.9 1937, coll. Smithsonian-Roebling Expedition mm cl, 2 &, 23.2 mm cl, 1 &, unmeasurable to Cuba: 1 (, 9.9 mm cl (USNM 260945); (RMNH 12210). Matanzas Bay, 2 m, coll. M. Ward: 2 (, 9.3± St. Kitts-Nevis: Sta. 67±58, Charlestown, 9.7 mm cl, 1 oviger, 15.2 mm cl (AM Nevis, 2±3 fms (ϭ 3.7±5.5 m), April 16, P45217). 1958, coll. Smithsonian-Bredin Caribbean Cayman Islands: 1 mi north of low point, Expedition: 1 (, 9.6 mm cl (USNM west beach, Grand Cayman, coll. unknown: 260860). 1 (, 5.9 mm cl, 1 &, 8.3 mm cl (ANSP Antigua and Barbuda: Sta. 92±58, be- uncataloged). tween Bird and Green Islands, NonSuch Bay, Jamaica: Kingston Harbor, 1893, coll. R. Antigua, 3±4 fms (ϭ 5.5±7.3 m), April 23± P. Bigelow: 3 &, 17.7±19.5 mm cl (USNM 24, 1958, coll. Smithsonian-Bredin Caribbe- 17990), 1 &, 17.9 mm cl (BMNH 1937.6.1.3 an Expedition: 1 &, 8.9 mm cl (USNM ex USNM 17990); Rio Bueno Bay, Aug. 260944). 1972, coll. E. A. Norse: 1 intersex, 16.8 mm Guadeloupe: Entre Basse Terre et la Ri- cl (LACM-AHF 1653±01). vieÁre des PeÁres, 15±20 m, Feb. 11, 1936, Dominican Republic: Barahona Bay, coll. R. Lami: 1 &, 5.0 mm cl (MNHN-Hi Santo Domingo, 1932±1933, coll. J. C. 198); Basse Terre, vers Vieux Fort, Feb. 20, Armstrong: 1 (, 11.0 mm cl (AMNH 1936, coll. R. Lami: 1 (, 6.6 mm cl 10356); Porto Plata, Santo Domingo, 27±30 (MNHN-Hi 199). ft (ϭ 8.2±9.1 m), July 1937, coll. W. J. St. Lucia: ``St. Lucia,'' June 16, 1880, Clench: 1 &, 11.9 mm cl (MCZ 9868). coll. J. Semper: 1 &, 24.1 mm cl (MCZ Puerto Rico: 1 km north of Mayaguez, 13261). July 26, 1974, coll. A. Williams: 2 (, 15.1± Barbados: ``Barbados,'' coll. C. T. 15.3 mm cl, 1 &, 19.7 mm cl (USNM Trenchman: 1 (, 12.6 mm cl (BMNH 267777); Sta. 6053, off Moon Castle, San 1921.6.9.5). Juan Harbor, 4±7.5 fms (ϭ 7.3±13.7 m), Jan. Trinidad and Tobago: Sta. 4019, Gulf of 15, 1899, coll. ``Fish Hawk'': 2 unmeasur- Apria, near San Fernando, Trinidad, 4.5 m, able juveniles (USNM 29011); San Juan, May 2, 1952, coll. Shell Expedition to the 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 329

Gulf of Paria: 1 juvenile, 2.4 mm cl, 1 un- 47Њ05ЈW, 21 fms (ϭ 38.4 m), Nov. 19, 1957, sexable specimen, 7.6 mm cl (RMNH 9097). coll. R/V ``Oregon'': 1 (, 18.9 mm cl Netherlands Antilles: Sint Michiels Baai, (USNM 101666); 00Њ08ЈS, 46Њ14ЈW, 24 m, CuracËao, 4 m, Jan. 1957, coll. L. B. Holthuis: March 2, 1963, coll. R/V ``Oregon'': 2 (, 1 (, 11.8 mm cl, 3 &, 6.1±11.7 mm cl. 1 &, 15.3±15.9 mm cl, 1 &, 16.5 mm cl (RMNH unmeasurable (RMNH 14646), 1 (, 11.5 23336); MaranhaÄo: Cruise 84, Sta. 23255, mm cl, 1 &, 11.5 mm cl (AMNH 18089 ex 00Њ34ЈN, 46Њ40ЈW, 45.7 m, Nov. 28, 1977, RMNH 14646). coll. D. Sutherland on R/V ``Oregon'': 1 &, Honduras: Ved Roman River, 1875, coll. 19.8 mm cl (HBOM 089:05495); Sta. 4220, P. Andersen: 1 (, 7.7 mm cl (ZMO F17537). 00Њ01ЈN, 45Њ48ЈW, 30 fms (ϭ 54.9 m), Panama: Sta. 258±7, Devil's Beach, July March 8, 1963, coll. R/V ``Oregon'': 1 (, 15, 1978, coll. M. L. Jones: 1 oviger, 16.4 15.7 mm cl, 1 &, 19.2 mm cl (USNM mm cl (USNM 260978); Shimmey Beach, 260814); Sta. 2091, 00Њ33ЈN, 47Њ03ЈW, 2 0 Ft. Sherman, Jan. 23, 1971, coll. L. G. Abele: fms (ϭ 36.6 m), Nov. 18, 1957, coll. R/V 2 (, 12.0±13.5 mm cl (USNM acc. 300691). ``Oregon'': 1 (, 16.4 mm cl (USNM Colombia: Sta. 103, Sample 405, Hum- 260815); Cruise 58, Sta. BBC 1617 (17700), boldt Bay, May 18, 1941, coll. J. C. Arms- 01Њ21ЈN, 47Њ32ЈW, 31±33 fms (ϭ 56.7±60.4 trong (``Askoy'' Expedition): 1 &, 18.2 mm m), May 12, 1975, coll. R/V ``Oregon II'': 1 cl (AMNH 18091); west coast of Santa Cat- (, 12.9 mm cl (USNM 260859); Cruise 58, alina Island, north of Fort Aury, Providencia Sta. BBC 1628, 00Њ54ЈN, 46Њ42ЈW, 38±43 Island Group, Aug. 12, 1969, coll. C. R. Gil- fms (ϭ 69.5±78.6 m), May 15, 1975, coll. R/ bert, J. C. Tyler, and S. Anderson: 1 &, 17.5 V ``Oregon II'': 1 (, 12.6 mm cl (USNM mm cl (USNM 260942); beach in front of 260861); EspõÂrito Santo: Praia dos Castel- Santa Mar Hotel, Santa Marta, Oct. 14, 1977, hamos, Anchieta, Nov. 21, 1971, coll. A. L. coll. M. L. Jones: 1 juvenile, 3.2 mm cl Castro: 1 &, 23.1 mm cl (MNRJ 1534); Rio (USNM 260946); 5 mi northwest of Galena de Janeiro: Prainha, Arraial de Cabo, Cabo Point Light, 12 fms (ϭ 21.9 m), April 25, Frio, Feb. 1985, coll. G. Nunan: 1 (, 18.4 1939, coll. R/V ``Velero III'': 1 (, 3.8 mm mm cl (MNRJ 1530); Itaipu, June 6, 1959, cl (LACM-AHF A48±39). coll. A. CoeÃlho, R. ArleÂ, and J. Becker: 1 &, Venezuela: Sta. 9, Isla de Margarita, Feb. 25.8 mm cl (MNRJ 1531); Praia dos Anjos, 16, 1977, coll. unknown: 1 (, 8.7 mm cl, 2 Arraial do Cabo, Feb. 1967, coll. J. Jurberg: &, 7.0±8.7 mm cl, 1 megalopa, 3.3 mm cl, 1 &, 22.0 mm cl (MNRJ 1532); Rio de Ja- 4 ®rst stage crabs, 2.6±3.3 mm cl (USNM neiro, coll. unknown: 1 (, 19.8 mm cl, 1 &, 260941); Sta. M±10±1, 1.5 m offshore, Isla 9.1 mm cl (MNRJ 1533); Sta. 39, 6 mi off Cubagua, west of Punta Brazil, Isla de Mar- mouth of Rio de Janeiro River, coll. ``Terra garita, Feb. 16. 1977, coll. M. L. Jones: 1 Nova'': 2 larvae, 4.7±4.8 mm (BMNH juvenile, 2.8 mm cl (USNM 260943); Sta. 1926.5.31.354±356); Rio De Janeiro, coll. S. M±4±1, 1.5 m offshore, approximately 5 km Krùyer: 1 &, 18.2 mm cl (ZMUC 2716); An- west of Guayacancito, Isla de Margarita, Feb. gra dos Reis, July 20, 1966, coll. unknown: 13, 1977, coll. M. L. Jones: 1 juvenile, 3.1 1 (, 7.3 mm cl (MNRJ 3855); Angra dos mm cl (USNM 260947). Reis, July 15, 1966, coll. unknown: 1 &, 7.1 French Guiana: Cayenne, coll. unknown: mm cl (MNRJ 3859); Angra dos Reis, July 1 (, 15.6 mm cl, syntype of A. oxyophthal- 7, 1966, coll. unknown: 1 (, 7.9 mm cl, 2 ma (BMNH 57.45). &, 4.9±8.3 mm cl (MNRJ 3861); Angra dos Brazil: ``Brazil,'' coll. unknown: 1 &, Reis, July 19, 1966, coll. unknown: 3 &, 6.1± 19.9 mm cl, syntype of A. oxyophthalma 7.7 mm cl (MNRJ 3862); Praia do Forte, (BMNH 64.7); ``Brazil,'' coll. unknown: 1 Cabo Frio, Aug. 24, 1986, coll. C. E. Ribei- (, 14.4 mm cl (UFES 1307); ``Brazil,'' coll. ro: 1 (, 22.3 mm cl (MNRJ 3864); SaÄo Pau- Banco Calipso: 1 &, 4.5 mm cl (MNRJ lo:SaÄo SebastiaÄo, Dec. 12, 1989, coll. un- 8164); AmapaÂ: Pesca Norte Island, 04Њ03ЈN, known: 1 (, 9.8 mm cl (AMNH 18090); Fla- 49Њ32.6ЈW, coll. S. Buitone: 1 juvenile, 3.9 mengo Cove, Ubatuba, Jan. 24, 1962, coll. mm cl (MNRJ 5408); ParaÂ: Sta. 2092, off L. Forneris: 1 &, 3.9 mm cl (RMNH 20958). the mouth of the Amazon River, 00Њ23ЈN, Limited Data: ``St. Croix'' (locality added 330 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 at later date): 1 &, 20.7 mm cl, holotype of sogastric region smooth; CG2 present as A. paretii (ANSP 4101a); ``Antilles'' (local- short medial element; CG3 broken into one ity may be inferred): 1 &, 21.9 mm cl (ANSP to three short elements and two long ele- 4101b); ``West Indies,'' 1 &, 12.1 mm cl ments between posterior lateral elements of (BMNH 1999.891). CG1; CG4 with one long (rarely two short), No Data:1(, 16.2 mm cl (ANSP 4763); anteriorly displaced, medial element between 1 &, 16.5 mm cl (USNM 42199); 1 &, 23.0 longer supralateral elements of CG4. Hepatic mm cl (BMNH 1998.92). region smooth, with oblique setose groove at DIAGNOSIS: Carapace wider than long, cov- median of lateral margin. Epibranchial re- ered with lightly setose grooves. Anterior gion generally triangular, smooth; posterolat- margin with 10±13 spines on either side of eral margin with three short rows of setae. ocular sinus. Setal ®eld with narrow lateral Metagastric region smooth; CG5 present as elements and concave anterior margin. CG1 two triangular elements. CG6 strongly cren- with separate posterior lateral elements; CG4 ulate, strongly anteriorly concave medially with one long (rarely two short), anteriorly and sloping out to anteriorly convex lateral displaced, medial element between longer thirds. CG7 oblique, almost reaching lateral supralateral elements of CG4; CG5 of two margins of median segment of CG6. Cardiac triangular elements; CG6 and CG7 separate; region smooth; CG8 present as three long el- CG8 entire; CG11 present. Rostrum present, ements; posterior element displaced slightly not reaching posterior margin of ocular plate. posteriorly. CG9 present as two short lateral Ocular plate triangular. Distal peduncular grooves with gap at midline. CG10 present segments dorsoventrally ¯attened and trian- as two long lateral elements, with gap be- gular in shape, tapering at tip, approximated tween fragments. CG11 present as short me- along mesial margins, lateral margins slight- dial element. Post-CG11 element present. ly convex, mesial margins straight. Cornea at Branchial region with numerous short, trans- tip. Dactylus of pereopod II with heel pro- verse rows of setae. Posterior margin deeply duced, tapered, and subacute. Dactylus of pe- and evenly convex, with submarginal groove reopod III with heel broad, projecting, acute. reaching two-thirds to three-fourths up mar- Dactylus of pereopod IV sinuous from base gin of posterior concavity. Branchiostegite to tip, with low rounded heel and shallow with short anterior submarginal spine; ante- indent. Telson of male ovate, slightly longer rior region with scattered short, transverse than wide, with broadly rounded tip, thickly lines ventral to linea anomurica; with many calci®ed medially, in¯ated dorsally, distal short rows of setae and sparsely covered with two-thirds with lateral decalci®ed region, long plumose setae ventrally; posterior re- median row of thin setae. Telson of female gion membranous, with numerous irregular ¯attened, ovate, and evenly calci®ed with fragments and sparsely covered with long slightly produced tip. plumose setae. DESCRIPTION: Carapace (®g. 104A) slightly Ocular plate (®g. 104B) triangular, with wider than long. Anterior margin slightly shallow median indentation; median pedun- concave on either side of ocular sinus, be- cular segments present as small ovate calci- coming convex laterally, with 10±13 large ®ed areas lateral to ocular plate. Distal pe- spines (n ϭ 6) along length. Rostrum as duncular segments elongate, subtriangular, small acute tooth, reaching to proximal mar- 0.20±0.36 length of carapace, with straight gin of ocular plate. Ocular sinus smoothly lateral and mesial margins, cornea covering concave and unarmed except in largest spec- distal tip; mesial margins approximated imens where lateral margins possess few ad- along proximal two-thirds; mesial and lateral ditional small spines. Frontal region smooth; margins with long plumose setae; tuft of plu- setal ®eld narrow anteriorly and posteriorly; mose setae at proximolateral ventral angle posterior lateral elements reduced to narrow and medial row of plumose setae extending bands of setae. CG1 parallel to anterior mar- from tuft to base of cornea. gin of carapace, sinuous, strongly crenulate, Antennule (®g. 104C) with segment III divided into medial fragment and curved, narrow proximally, expanding distally to two posteriorly displaced lateral elements. Me- times proximal width; with plumose setae on 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 331

Fig. 104. Albunea paretii GueÂrin MeÂneville, 1853: A±D, H±J, &, 11.5 mm cl, AMNH 18089; E± G, &, 7.0 mm cl, USNM 260941. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 1.6 mm (B, E, F), 2.2 mm (G, I), 3.3 mm (D, H, J), and 4.4 mm (A, C). dorsal and ventral margins and sparsely scat- or four articles (n ϭ 6) and plumose setae on tered on lateral surface; dorsal exopodal ¯a- dorsal and ventral margins. Segment II me- gellum with 78±97 articles (n ϭ 6), long plu- dially in¯ated in dorsal view, with plumose mose setae on dorsal and ventral margins; setae on dorsal and ventral margins and scat- ventral endopodal ¯agellum short with three tered on ventrolateral third of surface. Seg- 332 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 ment I wider than long, unarmed; dorsal third broadest medially, margins and mesioventral of lateral surface rugose, with long plumose surface with long plumose setae. Endopod setae; long plumose setae on dorsal and ven- ¯attened and elongate, reaching two-thirds to tral margins. distal end of proximal exopodal segment; Antenna (®g. 104D) with segment V ap- plumose setae on margins and median of lat- proximately three times longer than wide, eral surface. with long plumose setae on dorsal margin Maxilliped II (®g. 104I) dactylus evenly and scattered on lateral surface; ¯agellum rounded, length equal to width, with thick with seven articles (n ϭ 6), long plumose simple setae distally and on distolateral sur- setae on dorsal, ventral, and distal margins. face. Propodus 1.5 times wider than long, Segment IV expanded distally, with long plu- slightly produced at dorsodistal angle, with mose setae on dorsal, ventral, and distal mar- plumose setae on dorsal margin and patch of gins, and row of setae on dorsolateral sur- long simple setae on lateral surface and ven- face. Segment III with long plumose setae on trolateral angle. Carpus not produced dorso- dorsal and ventral margin and in short row distally, approximately two times longer than on surface. Segment II short, widening dis- wide; long simple setae on dorsal margin. tally, rugose, with plumose setae on margins Merus approximately three times longer than and scattered on lateral surface; antennal aci- wide, margins parallel; with simple and plu- cle long, thin, and exceeding distal margin of mose setae on margins and scattered on sur- segment IV by one-®fth length of segment face. Basis-ischium incompletely fused, with IV, with long plumose setae on dorsal mar- plumose setae on margins. Exopod one- gin. Segment I rounded proximally, ¯attened fourth longer than merus, ¯agellum with one ventrolaterally, with long plumose setae on elongate article. margins and scattered on surface rugae be- Maxilliped III (®g. 104J) dactylus with hind spine; lateral surface with acute spine rounded tip; long plumose setae on margins dorsodistally, with low semicircular dorso- and lateral surface. Propodus dorsally in¯at- lateral lobe ventrodistal to spine; segment ed, with longitudinal median row of plumose with ventromesial antennal gland pore. setae on lateral surface; margins with plu- Mandible (®g. 104E) incisor process with mose setae. Carpus produced onto propodus two teeth; cutting edge smooth. Palp three- almost one-fourth length of propodus; lateral segmented, with plumose setae on margins surface with two rows of plumose setae on and long, thick, simple setae arising from surface; plumose setae on margins. Merus in- bend in second segment and on distal margin ¯ated, unarmed, with plumose setae on mar- of terminal segment. gins and scattered on lateral surface. Basis- Maxillule (®g. 104F) distal endite proxi- ischium incompletely fused, with weak crista mally narrow, widening to in¯ated distal end, dentata of two or three teeth. Exopod two- with thick simple setae on distal margin and segmented: proximal segment small; distal thin simple setae on dorsal margin. Proximal segment styliform, tapering, approximately endite with thick simple setae on distal mar- one-third length of merus; with plumose se- gin. Endopodal external lobe truncate distally tae on margins; without ¯agellum. and curled under; internal lobe reduced with Pereopod I (®g. 105A) dactylus curved three thick setae at distolateral margin. and tapering; lateral and mesial surfaces Maxilla (®g. 104G) exopod evenly round- smooth; dorsal margin with long plumose ed, with plumose setae along distal margin. and short simple setae; ventral margin with Scaphognathite bluntly angled on posterior short simple setae. Propodal lateral surface lobe, with plumose setae. with numerous short, transverse rows of se- Maxilliped I (®g. 104H) epipod with plu- tose rugae; dorsal margin unarmed; ventral mose setae on margins, distolateral surface, margin distally produced into acute spine; and mesial surface. Endite tapered distally cutting edge lacking teeth, lined with long and subequal to ®rst segment of exopod. Ex- plumose setae; dorsal margin with long plu- opod with two segments; proximal segment mose setae, ventral margin with short simple narrow, margins parallel with plumose setae; setae. Carpus with dorsodistal angle pro- distal segment spatulate, longer than wide, duced into strong corneous-tipped spine; dor- 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 333

Fig. 105. Albunea paretii GueÂrin MeÂneville, 1853: A±E, G, &, 11.5 mm cl, AMNH 18089; F, (, 11.5 mm cl, AMNH 18089. A. Left pereopod I, lateral view. B. Left pereopod II, lateral view. C. Left pereopod III, lateral view. D. Left pereopod IV, lateral view. E. Abdominal somites I±VI, dorsal view. F. Telson of (, dorsal view. G. Telson of &, dorsal view. Scale ϭ 3.0 mm (F, G), 3.3 mm (A), and 4.4 mm (B±E). sal margin with short transverse grooves be- Basis-ischium incompletely fused, unarmed. hind spine; dorsal and distal margins with Coxa unarmed. long plumose setae; lateral surface with Pereopod II (®g. 105B) dactylus smooth; small distal rugose area, with few transverse base to heel concave, heel produced but sub- setose ridges on distal half of surface; mesial acute, heel to tip with wide, acute indent, tip surface smooth, with medial transverse row acute, tip to base broadly convex; lateral sur- of setae, margins with long plumose setae. face smooth, with several small tufts of short Merus unarmed; lateral surface with scat- setae in generally straight line across medi- tered transverse rows of long plumose setae, oproximal surface, several widely spaced margins with long plumose setae; mesial sur- submarginal tufts of short setae dorsodistally; face with few scattered setae; fully calci®ed. mesial surface smooth, ventral margin with 334 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 long plumose setae, dorsal margin with short and two interrupted rows of setae ventrally; simple setae and patch of long plumose setae mesial surface smooth, with long plumose at base. Propodal dorsal surface smooth, with setae on margins. Merus smooth, with large ventral margin in¯ated and rounded; oblique decalci®ed window covering nearly half of row of long plumose setae on distal margin lateral surface medially; dorsal and ventral of lateral surface; distal and ventral margins margins unarmed, with long plumose setae; with long plumose setae; dorsolateral surface distolateral margin with long plumose setae; as narrow, oblique, ¯attened shelf, with short mesial surface smooth. Basis-ischium incom- setae on dorsal margin and long plumose se- pletely fused and unarmed. Coxa with low tae on ventral margin; mesial surface with tubercle on anterior margin. Female with elevated, curved, setose ridge from ventral large gonopore on anterior mesial margin of junction with dactylus almost to ventral coxa, surrounded with short plumose setae; proximal junction with carpus. Carpus slight- male with small depression in analogous po- ly produced and gently rounded dorsodistal- sition (not true pore). ly, dorsal margin with few low spines on dis- Pereopod IV (®g. 105D) dactylus with tal two-thirds; lateral surface smooth, with base to tip convex proximally to concave dis- setose mat at tip of produced area and irreg- tally, tip acute, tip to base concave distally ular, interrupted row of rugae and submar- to convex proximally; lateral surface smooth, ginal elevated ridge ventrally, rugae and ventral margin with long plumose setae, dor- ridge with long plumose setae; margins with sal margin with short simple setae; mesial long plumose setae; mesial surface smooth, surface with dorsal decalci®ed region, de- with row of long plumose setae subdorsally. marcated ventrally by longitudinal elevated Merus with large median decalci®ed window ridge with row of short setae; with setose covering nearly all of lateral surface, with punctations ventral to decalci®ed window. few scattered long plumose setae on surface Propodus expanded dorsally and ventrally; and margins; mesial surface nearly smooth, ventral expansion reaching ventral margin of with two long rows of setae. Basis-ischium dactylus, margin with long plumose setae; incompletely fused and unarmed. Coxa with dorsal expansion with row of long plumose small low tubercle on anterior margin. setae dorsally, oblique area with mat of short Pereopod III (®g. 105C) dactylus with simple setae; lateral and mesial surfaces base to heel concave, heel acutely produced, smooth. Carpus slightly produced dorsodis- heel to tip with broadly concave indent and tally; ventral three-fourth of lateral surface small concave region at midpoint of proxi- and mesial surface smooth, dorsodistal quar- mal margin, tip acute, tip to base smoothly ter of lateral surface with mat of short setae; convex; lateral surface smooth, with several dorsal margin with short simple and long small tufts of short setae in generally straight plumose setae; ventral margin with short line across medioproximal surface, dorsodis- simple setae; mesial surface faintly decalci- tal margin with tufts of short setae; ventral ®ed medially. Merus with scattered, short, margin with long plumose setae, dorsal mar- transverse rows of setae on lateral surface, gin with short simple and plumose setae; me- dorsal and ventrodistal margins with long sial surface smooth, with plumose setae plumose setae; proximoventral half of mesial proximally at junction with propodus. Pro- surface with large decalci®ed window. Basis- podus not in¯ated dorsoventrally; lateral sur- ischium incompletely fused and unarmed. face smooth, with long plumose setae in Coxa unarmed. oblique row, simple setae on dorsal margin; Abdomen (®g. 105E) with somite I length dorsolateral surface narrow, oblique, ¯at- and width subequal, widest posteriorly; dor- tened, with setose mat; mesial surface with sal surface with anterior margin slightly con- scattered long setae on and near distal margin cave; posterior margin curved, with elevated and in oblique row on surface. Carpus pro- submarginal row of short setae; small trans- duced dorsodistally, only slightly exceeding verse decalci®ed windows laterad of segment proximal margin of propodus; dorsolateral median. Somite II dorsal surface with sub- margin unarmed; lateral surface slightly ru- marginal transverse ridge anteriorly; with gose dorsodistally, with mat of short setae small transverse decalci®ed windows laterad 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 335 of segment median just anterior to submar- oxyophthalma); BMNH 64.7 (syntype of A. ginal ridge; pleura expanded and directed an- oxyophthalma); the repository of an addi- terolaterally; lateral margins rounded, ante- tional syntype of A. oxyophthalma cited by rior and lateral margins with long plumose both White (1847) and Miers (1878) from St. setae, posterior margin with short setae; pos- Christophers is unknown. teromesial angle with mat of short simple se- TYPE LOCALITIES: Albunea paretii: ``Mer tae. Somite III similar to somite II, but nar- de GeÃnes'' or ``Mediterranea?'' (incorrect lo- rower, shorter; pleura thinner and shorter calities). Albunea oxyophthalma: St. Chris- than on somite II, directed anterolaterally, tophers, Cayenne [French Guiana], and Bra- with setae as in somite II; anterolateral angle zil. subacute; dorsal surface obliquely ¯attened REMARKS: ANSP 4101a is clearly GueÂrin anterolaterally. Somite IV similar to somite MeÂneville's (1853) ®gured specimen and is III, but thinner and shorter; dorsal surface the holotype by monotypy (see also Boyko, with few short setae anterolaterally; pleura 2000b). Kingsley (1880) stated that GueÂrin thinner and shorter than on somite III, di- MeÂneville's type was not in the ANSP, but rected posterolaterally; dorsal surface he was likely misled by the poor labeling of obliquely ¯attened anterolaterally; margins the specimen. The locality data ``Antilles'' with long plumose setae. Somite V wider was added to ANSP 4101a at a later date and than somite IV; lateral margins with plumose may have come from Stimpson's (1858) lo- setae; pleura absent. Somite VI slightly cality data for the species as ``Antillarum?'' broader than somite V; dorsal surface with The name A. oxyophthalma is a nomen nu- short transverse rows of setae laterad of mid- dum from both White (1847, as A. oxy- line; pleura absent. ophthalmus) and Stimpson (1858). Females with uniramous, paired pleopods Much of the dif®culty in recognizing the on somites II±V; males without pleopods. identity of this species was caused by GueÂrin Telson of male (®g. 105F) broadly trian- MeÂneville's (1853) uncertainty as to the type gular, slightly longer than wide, with broadly locality. GueÂrin MeÂneville (1853) was told rounded tip; thickly calci®ed medially, in- that his specimen came from Genoa (Italy), ¯ated dorsally; distal half with lateral decal- which he thought likely, but he also was ci®ed region; median longitudinal groove ex- aware that the ship which brought him the tending one-half length, row of long simple specimen had been to the Americas, and he setae of either side of median groove begin- mentioned this possibility as well. The latter ning at distal end and continuing almost to is the correct locality, and the specimen was distal margin of telson; proximolateral angles likely collected in the Caribbean or South with patch of short simple setae; margins America. with long simple setae. Telson of female (®g. It has been asserted by some authors (e.g., 105G) ¯attened, ovate, and evenly calci®ed Monod, 1956) that the correct spelling of this with slightly produced tip; median groove speci®c name should be ``paretoi,'' as the similar to male, setal row from end of me- species was named after the Marquis Pareto. dian groove to near distal margin of telson However, GueÂrin MeÂneville (1853) gave the with simple setae approximately one-half spelling as ``paretii'' twice, so there is no length of those on male; proximolateral angle evidence of a lapsus calami or printer's error, with patch of setae, margins with long simple and A. paretii is therefore the incorrect orig- setae. inal spelling and the one which must be used DISTRIBUTION: Bermuda and the Florida for the species. Keys south throughout the Caribbean and Gurney's (1942) Bermuda larvae are more Central America to Rio Grande do Sul, Bra- likely to be A. paretii rather than A. gibbesii, zil (Calado, 1987), in 0±101 m depth (Ca- as the latter species has only just been con- lado, 1987). ®rmed from Bermuda (herein) and is evi- MAXIMUM SIZE: Males: 22.3 mm cl; fe- dently much rarer there. The same can be males: 25.8 mm cl. said of Gurney's (1924) larvae from off Rio TYPE SPECIMENS: ANSP 4101a (holotype de Janeiro, Brazil, where the common spe- of A. paretii); BMNH 57.45 (syntype of A. cies is also A. paretii. The larvae of ``Albu- 336 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 nea sp.'' reported by Bowen et al. (1979) unbroken CG8, and the different shape of the were also probably this species, rather then dactyli of pereopod III. the less common A. gibbesii. The specimen from Rio Bueno Bay, Ja- Albunea lucasia de Saussure, 1853 maica (LACM-AHF 1653±01) is the type Figures 106, 107 host of the bopyrid isopod Albunione inde- Albuminea [sic] Lucasia de Saussure, 1853: 367, cora (Markham, 1988), which occurs in the pl. 12, ®g. 4*. branchial chamber. This host specimen is a Albunea lucasii [sic]: Stimpson, 1857: 485*. ± true intersex/feminized male, with large ple- Miers, 1878: 330±331. ± Steinbeck and Rick- opods and well-developed gonopores on the etts, 1941: 459. ®fth pereopods. This parasite is also known Albunaea [sic] Lucasii [sic]: Stimpson, 1858: 230 from the same host species in the Bahamas (list). (ex AMNH 5464) (Markham and Boyko, Albunea lucasia: Ortmann, 1896: 225 (list). ± Ort- 1999). mann, 1901: 1275. ± Gordon, 1938: 187 (list). The drawings of this species given by Ca- ± Bott, 1955: 50±51, pl. 4, ®g. 4a, b. ± Schuster- lado (1987: ®gs. 9. 10a) are inaccurate in that Dieterichs, 1956: 29, 34, 37, 40, 51. ± Dexter, 1972: 455*. ± Abele, 1976: 266±267*. ± Haig, the carapace grooves are highly stylized and, 1980: 289, ®g. 19.5. ± Moran, 1984: 79, ®g. 6. moreover, very different between the two il- ± CoeÃlho and Calado, 1987: 42±43, table 1. ± lustrations. The branchiostegite was drawn in Lemaitre and Alvarez LeoÂn, 1993: 49 (list). ± Calado's ®gure 9 as if it were fused to the Hendrickx, 1992: 9 (list). ± Calado, 1995: 38± carapace and were spinose on the margins, 39, pl. 4, ®g. c, pl. 9, ®g. a. ± Calado, 1997a: neither of which is true. These illustrations 17. ± Hendrickx and Harvey, 1999: 366 (list). were repeated by Calado (1995). All of the Albunea lucasi [sic]: Prahl et al., 1979: 55. ± material listed by Calado (1995) was from Prahl, 1986: 97. Brazil and, although not examined, undoubt- Albumienea [sic] lucasia: Calado, 1995: 38. edly belongs to this species rather than to the not Albunea lucasia: Ramos and Rios, 1995: 103, ®g.5(ϭ Albunea galapagensis, n. sp.). northern A. catherinae, n. sp. Although previous authors (e.g., Monod, MATERIAL EXAMINED: Mexico: ``Mexico,'' 1956) indicated that morphology-based sep- 1 (, 13.1 mm cl (ANSP 4102a); ``Mexico,'' aration of the trans-Atlantic A. paretii com- 1926, coll. Sec. Agriculture y Fomento: 1 (, plex into component species was not possi- 19.2 mm cl (USNM 62387); ``Isla Grande,'' ble, it is now evident that this is not correct. 10 fms (ϭ 18.3 m), April 8, 1937, coll. W. The Caribbean and South American speci- Williams and F. E. Lewis on R/V ``Strang- mens are all true A. paretii, while those from er'': 1 (, 8.5 mm cl (USNM 260983); Sta. North America are almost all A. catherinae, 963±39, ``White Friars Rocks,'' May 7, n. sp. The African material is all A. elegans. 1939, coll. R/V ``Velero III'': 1 (, 10.5 mm It is not known how far south along the west- cl, 3 &, 5.9±7.1 mm cl (USNM 260982); ern Gulf of Mexico, if at all, A. catherinae, Baja California Norte: Punta Gorda, 14 fms n. sp. ranges. (ϭ 25.6 m), April 24, 1937, coll. W. Wil- This species typi®es the ``paretii-group'' liams and F. E. Lewis on R/V ``Stranger'': 2 of Albunea, which also includes A. lucasia, &, 9.8±15.6 mm cl (USNM 267775); Baja A. elegans, A. steinitzi, and A. catherinae,n. California Sur: Cabo San Lucas, south of sp. All these species share a similar mor- Santa Rosalita, March 24, 1940, coll. E. phology of the male telson, and all have Ricketts: 1 &, 5.5 mm cl (USNM 267772); acute heels on the dactyli of pereopod III. Bahia de la Ventana, 5±10 fms (ϭ 9.1±18.3 Albunea paretii is the Atlantic analogue of m), April 21, 1937, coll. W. Williams and F. A. lucasia. Albunea paretii is easily distin- E. Lewis on R/V ``Stranger'': 1 &, 6.7 mm guished from A. catherinae, n. sp. by the dis- cl (USNM 267773); Bahia Salinas, Isla Car- tal ocular peduncle/carapace length ratio and men, 20 fms (ϭ 36.6 m), Dec. 19, 1931, coll. by the shapes of the dactyli of pereopods III S. A. Glassell: 1 juvenile, 4.5 mm cl (USNM and IV (see under A. catherinae for specif- 260997); Sonora: Puerto PenÄasco, May 3, ics). It can be separated from A. elegans by 1935, coll. S. A. Glassell: 1 (, 8.1 mm cl the shortness of the branchiostegite spine, (USNM 267768); north TiburoÂn Island, 20 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 337 fms (ϭ 36.6 m), Jan. 1, 1932, coll. S. A. Costa Rica: Gulf of Nicoya, July 6, 1981, Glassell: 1 unsexable specimen, 5.1 mm cl coll. unknown: 1 (, 5.6 mm cl (USNM (USNM 267771); Guaymas, Feb. 1951, coll. 267767). unknown: 1 (, 18.1 mm cl, 1 &, 22.4 mm Panama: Chame Point, June±July 1912, cl (CASIZ 109241); Bahia Bocochibampo, coll. R. Tweedie: 1 (, 11.8 mm cl, 1 &, 14.2 Guaymas, Nov. 1952, coll. J. P. Strohbeen: 1 mm cl (USNM 66048); Sta. 183±6, Culebra &, 20.3 mm cl (CASIZ 109245); Bahia Sol- Island, March 10, 1974, coll. NMNH-STRI dado, Guaymas, Nov. 27, 1955, coll. L. O. Panama Survey: 1 (, 18.9 mm cl (USNM Miles: 1 (, 13.5 mm cl, 1 &, 11.8 mm cl 260986); Sta. 253±3±1, Culebra Beach, Feb. (CASIZ 109247); Punta Cholla, May 13, 7, 1978, coll. H. W. Kaufman: 1 juvenile, 2.3 1941, coll. S. A. Glassell: 1 &, 9.0 mm cl mm cl (USNM 260991); Sta. 183±6B, Cu- (USNM 260994); Sinaloa: Mazatlan, 1 &, lebra Island, March 10, 1974, coll. M. L. 19.7 mm cl, holotype (ANSP 4102); Jalisco: Jones and H. W. Kaufmann: 1 juvenile, 3.1 Bahia Tenacatita Bay, 8±10 fms (ϭ 14.6± mm cl (USNM 304299); Sta. 263±2, Whore- 18.3 m), Feb. 17, 1938, coll. S. A. Glassell: house Reef, 1 m, July 20, 1978, coll. M. L. 2 (, 4.4±4.5 mm cl, 1 &, 5.7 mm cl (USNM Jones: 3 (, 6.7±7.9 mm cl, 1 &, 8.8 mm cl 267770); Bahia Tenacatita, 19Њ18ЈN, (USNM 260999); Sta. 182±1, Pilot House 104Њ51ЈW, 7 fms (ϭ 12.8 m), April 11, 1937, Beach, 4 ft (ϭ 1.2 m), March 9, 1974, coll. coll. F. E. Lewis on R/V ``Stranger'': 1 (, M. L. Jones and H. W. Kaufmann: 1 juvenile, 10.8 mm cl, 4 &, 8.1±9.2 mm cl, 1 oviger, 3.6 mm cl (USNM 260981); Sta. 182±5, Pi- 12.2 mm cl (USNM 260998); Puerto Vallar- lot House Beach, 60 ft (ϭ 18.2 m), March 9, ta, 10 fms (ϭ 18.3 m), April 13, 1937, coll. 1974, coll. M. L. Jones and H. W. Kaufmann: W. Williams and F. E. Lewis on R/V 1 juvenile, 4.5 mm cl (USNM 260989); Ft. ``Stranger'': 1 &, 10.5 mm cl (USNM Amador, Naos Island, June 1969, coll. D. 304300); Puerto Vallarta, 10 fms (ϭ 18.3 m), Dexter: 1 (, unmeasurable, 1 &, 7.9 mm cl April 13, 1937, coll. W. Williams and F. E. (USNM 260988); Pilot House Beach, Naos Lewis on R/V ``Stranger'': 1 &, 12.9 mm cl Island, Jan. 28, 1971, coll. T. A. Biffar: 1 (, (USNM 260992); Bahia Chamela, 8 fms (ϭ 17.5 mm cl, 1 oviger, 16.1 mm (USNM 14.6 m), Feb. 17, 1938, coll. S. A. Glassell: 304306); Ft. Amador, Boy Scout Islands, be- 1 (, 6.7 mm cl (USNM 260990); Guerrero: yond Naos Island, July 2, 1969, coll. L. G. Zihuatanejo, 20 fms (ϭ 36.6 m), Feb. 24, Abele: 4 &, 6.2±15.5 mm cl (USNM 1938, coll. S. A. Glassell: 1 juvenile, 3.3 mm 304307); Isla Gobernadoro, north of Panama cl (USNM 267774); Zihuatanejo, 12 fms (ϭ City, 50 ft (ϭ 15.2 m), coll. unknown: 1 (, 21.9 m), Feb. 24, 1938, coll. S. A. Glassell: 12.1 mm cl, 1 carapace, 12.1 mm cl (Feld- 1 &, 7.2 mm cl (USNM 260984); Bahia Dul- mann Collection). ce, 20 fms (ϭ 36.6 m), April 5, 1937, coll. Ecuador: Sta. 15, La Libertad, 5±12 fms W. Williams and F. E. Lewis on R/V (ϭ 9.1±21.9 m), Feb. 21, 1939, coll. F. E. ``Stranger'': 2 (, 8.5±11.3 mm cl (USNM Lewis on R/V ``Stranger'': 2 (, 7.2±10.3 260985); Oaxaca: Guatulco, 50 fms (ϭ 91.5 mm cl, 2 &, 7.6±7.9 mm cl, 1 anterior half, m), March 7, 1938, coll. S. A. Glassell: 1 &, 8.4 mm cl (USNM 260987). 9.7 mm cl (USNM 267769); Sta. 22, Puerto No Data:1&, 21.5 mm cl (MNHN-Hi Guatulco, 3±30 fms (ϭ 5.5±54.9 m), March 188). 15, 1939, coll. F. E. Lewis on R/V ``Strang- DIAGNOSIS: Carapace longer than wide, er'': 1 &, 12.6 mm cl (USNM 260995); Sta. covered with lightly setose grooves. Anterior 21, Bahia Chipequa, Gulf of Tehuantepec, 7± margin with 9±13 spines on either side of 18 fms (ϭ 12.8±32.9 m), March 15, 1939, ocular sinus. Setal ®eld with narrow lateral coll. R/V ``Stranger'': 1 (, 9.1 mm cl, 1 &, elements and concave anterior margin. CG1 11.1 mm cl (USNM 260996). with separate posterior lateral elements; CG4 Guatemala: ``Guatemala,'' coll. Paessler: with two to four short, anteriorly displaced, 2 (, 7.9±8.0 mm cl, 2 &, 8.0±14.0 mm cl medial elements between longer supralateral (ZMH K±5131); San Jose, 10±12 fms (ϭ elements of CG4; CG5 of two convex ele- 18.3±21.9 m), Feb. 2, 1939, coll. F. E. Lewis: ments; CG6 and CG7 separate; CG8 broken; 1 &, 8.2 mm cl (USNM 260993). CG11 present. Rostrum present, not reaching 338 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 posterior margin of ocular plate. Ocular plate placed. CG9 present as two short lateral triangular. Distal peduncular segments dor- grooves with gap at midline. CG10 present soventrally ¯attened and triangular in shape, as two long curved lateral elements, with gap tapering at tip, approximated along mesial between fragments. CG11 present as long margins, lateral margins convex, mesial mar- medial element. Post-CG11 element absent. gins straight. Cornea at tip. Dactylus of pe- Branchial region with numerous short, trans- reopod II with heel produced and rounded. verse rows of setae. Posterior margin deeply Dactylus of pereopod III with heel broad, and evenly convex, with submarginal groove projecting, acute. Dactylus of pereopod IV reaching four-®fths up margin of posterior sinuous from base to tip, with low rounded concavity. Branchiostegite with strong ante- heel and shallow indent. Telson of male rior submarginal spine; anterior region with ovate, slightly longer than wide, with broadly scattered short, transverse lines ventral to lin- rounded tip, thickly calci®ed medially, in- ea anomurica; with many short rows of setae ¯ated dorsally, distal two-thirds with lateral and sparsely covered with long plumose se- decalci®ed region, median row of thin setae. tae ventrally; posterior region membranous, Telson of female ¯attened, ovate, and evenly with numerous irregular fragments and calci®ed with slightly produced tip. sparsely covered with long plumose setae. DESCRIPTION: Carapace (®g. 106A) slightly Ocular plate (®g. 106B) triangular, with longer than wide. Anterior margin slightly shallow median indentation; median pedun- concave on either side of ocular sinus, be- cular segments present as small ovate calci- coming convex laterally, with 9±13 large ®ed areas lateral to ocular plate. Distal pe- spines (n ϭ 6) along length. Rostrum as duncular segments elongate, subtriangular, small acute tooth, not reaching proximal with slightly convex lateral and straight me- margin of ocular plate. Ocular sinus smooth- sial margins, cornea covering distolateral tip; ly concave and unarmed, except in largest mesial margins approximated along length; specimens which possess one or two small mesial and lateral margins with short plu- spines. Frontal region smooth; setal ®eld nar- mose setae; tuft of plumose setae at proxi- row anteriorly and posteriorly; posterior lat- molateral ventral angle and medial row of eral elements reduced to narrow bands of se- plumose setae extending from tuft along tae. CG1 parallel to anterior margin of car- proximal two-thirds of segment. apace, sinuous, strongly crenulate, divided Antennule (®g. 106C) with segment III into medial fragment and curved, posteriorly narrow proximally, expanding distally to displaced lateral elements. Mesogastric re- twice proximal width; with plumose setae on gion smooth; CG2 present as one or two dorsal and ventral margins; dorsal exopodal short medial elements; CG3 broken into one ¯agellum with 101±139 articles (n ϭ 6), long or two long, anteriorly displaced, medial el- plumose setae on dorsal and ventral margins; ements and two long lateral elements be- ventral endopodal ¯agellum with two to four tween posterior lateral elements of CG1; articles (n ϭ 6), plumose setae on dorsal and CG4 with two to four short, anteriorly dis- ventral margins. Segment II medially in¯ated placed, medial elements between longer su- in dorsal view, with plumose setae on dorsal pralateral elements of CG4. Hepatic region and ventral margins and in transverse medial smooth, with oblique setose groove at me- row on lateral surface. Segment I as long as dian of lateral margin. Epibranchial region wide, with short acute spine on dorsal mar- generally triangular, smooth; posterolateral gin; dorsal third of lateral surface rugose, margin with two short rows of setae. Meta- with long plumose setae; long plumose setae gastric region smooth; CG5 present as two on dorsal and ventral margins. convex elements. CG6 strongly crenulate, Antenna (®g. 106D) with segment V ap- strongly anteriorly concave medially and proximately three times longer than wide, sloping out to anteriorly convex lateral with long plumose setae on dorsal margin thirds. CG7 oblique, not reaching lateral and scattered on lateral surface; ¯agellum margins of median segment of CG6. Cardiac with seven or eight articles (n ϭ 6), long plu- region smooth; CG8 present as one to three mose setae on dorsal, ventral, and distal mar- long elements, medial element anteriorly dis- gins. Segment IV expanded distally, with 2002 BOYKO: WORLDWIDE REVISION OF ALBUNEIDAE 339

Fig. 106. Albunea lucasia de Saussure, 1853: A, B, D±J, (, 17.5 mm cl, USNM 304306; C, oviger, 16.1 mm cl, USNM 304306. A. Carapace, branchiostegite, and ocular peduncles, dorsal view. B. Ocular peduncles, dorsal view. C. Left antennule, lateral view. D. Left antenna, lateral view. E. Left mandible, mesial view. F. Left maxillule, lateral view. G. Left maxilla, lateral view. H. Left maxilliped I, lateral view. I. Left maxilliped II, lateral view. J. Left maxilliped III, lateral view. Scale ϭ 2.2 mm (B, F), 3.3 mm (A, E, I), and 4.4 mm (C, D, G, H, J). 340 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY NO. 272 long plumose setae on dorsal, ventral, and slightly produced at dorsodistal angle, with distal margins, and short scattered setae on plumose setae on dorsal margin and patch of lateral surface. Segment III with long plu- long simple setae on lateral surface and ven- mose setae on ventral margin, short simple trolateral angle. Carpus not produced dorso- setae on dorsal margin and scattered on lat- distally, approximately two times longer than eral surface. Segment II short, widening dis- wide; long simple setae on dorsal margin. tally, rugose, with plumose setae on margins Merus approximately three times longer than and scattered on lateral surface; antennal aci- wide, margins parallel; with simple and plu- cle long, thin, and exceeding distal margin of mose setae on margins and scattered on sur- segment IV by one-fourth length of segment face. Basis-ischium incompletely fused, with IV, with long plumose setae on dorsal mar- plumose setae on margins. Exopod one-half gin. Segment I rounded proximally, ¯attened longer than merus, ¯agellum with one elon- ventrolaterally, with long plumose setae on gate article. margins and scattered on surface rugae be- Maxilliped III (®g. 106J) dactylus with hind spine; lateral surface with acute spine rounded tip; long plumose setae on margins dorsodistally, with low semicircular dorso- and lateral surface. Propodus dorsally in¯at- lateral lobe ventrodistal to spine; segment ed, with longitudinal median row of plumose with ventromesial antennal gland pore. setae on lateral surface; margins with plu- Mandible (®g. 106E) incisor process with mose setae. Carpus produced onto propodus two teeth; cutting edge smooth. Palp three- almost one-third length of propodus; lateral segmented, with plumose setae on margins surface with two rows of plumose setae on and long, thick, simple setae arising from surface; plumose setae on margins. Merus in- bend in second segment and on distal margin ¯ated, unarmed, with plumose setae on mar- of terminal segment. gins and scattered on lateral surface. Basis- Maxillule (®g. 106F) distal endite proxi- ischium incompletely fused, with weak crista mally narrow, widening to in¯ated distal end, dentata of three or four teeth. Exopod two- with thick simple setae on distal margin and segmented: proximal segment small; distal thin simple setae on dorsal margin. Proximal segment styliform, tapering, approximately endite with thick simple setae on distal mar- two-®fths length of merus; with plumose se- gin. Endopodal external lobe truncate distally tae on margins; without ¯agellum. and curled under; internal lobe reduced with Pereopod I (®g. 107A) dactylus curved three thick setae at distolateral margin. and tapering; lateral and mesial surfaces Maxilla (®g. 106G) exopod evenly round- smooth; dorsal margin with long plumose ed, with plumose setae along distal margin. and short simple setae; ventral margin with Scaphognathite bluntly angled on posterior short simple setae. Propodal lateral surface lobe, with plumose setae. with numerous short, transverse rows of se- Maxilliped I (®g. 106H) epipod with plu- tose rugae; dorsal margin unarmed; ventral mose setae on margins, distolateral surface, margin distally produced into acute spine; and mesial surface. Endite tapered distally cutting edge lacking teeth, lined with long and subequal to ®rst segment of exopod. Ex- plumose setae; dorsal margin with long plu- opod with two segments; proximal segment mose setae, ventral margin with short simple narrow, parallel margins with plumose setae; setae. Carpus with dorsodistal angle pro- distal segment spatulate, longer than wide, duced into strong corneous-tipped spine; dor- broadest medially, margins and mesioventral sal margin with short transverse grooves be- surface with long plumose setae. Endopod hind spine; dorsal and distal margins with ¯attened and elongate, reaching two-thirds to long plumose setae; lateral surface with distal end of proximal exopodal segment; small distal rugose area, with few transverse, plumose setae on margins and median of lat- setose ridges on distal half of surface; mesial eral surface. surface smooth, with medial transverse row Maxilliped II (®g. 106I) dactylus evenly of setae, margins with long plumose setae. rounded, length equal to width, with thick Merus unarmed; lateral surface with scat- simple setae distally and on distolateral sur- tered transverse rows of long plumose setae, face. Propodus 1.5 times wider than long, margins with long plumose setae; mesial sur-