PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 117(3):368-376. 2004. A new species of (: Agostocarididae) from Acklins Island, Bahamas

Fernando Alvarez, Jose Luis Villalobos, and Thomas M. Iliffe (FDA, JLV) Colleccion Nacional de Crustaceos, Institute de Biologia, Universidad Nacional Autonoma de Mexico, Apartado Postal 70-153, Mexico 04510 D.F., Mexico, e-mail: [email protected]; (TMI) Department of Marine Biology, Texas A&M University at Galveston, Galveston, Texas 77553-1675, U.S.A.

Abstract.—The new bresilioid shrimp Agostocaris acklinsensis is described from an anchialine cave in Acklins Island, Bahamas. This is the third species described in the genus. The new species is characterized by having small ex- opods on the third and fourth pereiopods, one spine on the ischium of the fifth pereiopod, and an outer ramus of the uropods with one distolateral spine. A key to the species of Agostocaris is provided.

The family Agostocarididae Hart & range of variation in taxonomically impor­ Manning, 1986, was created to accommo­ tant characters increases, making it neces­ date Agostocaris williamsi, from Grand Ba­ sary to provide a new diagnosis for the ge­ hama and Turks and Caicos, a species that nus. appeared to be morphologically similar to some species in the De Haan, 1849, Materials and Methods and the Caiman, 1896, but had a distinct morphology of the propodus and Specimens of the new Agostocaris de­ dactylus of the first two pereiopods. Ken- scribed herein were collected during an ex­ sley (1988) described a second species from pedition to Crooked and Acklins Islands, Cozumel, Mexico, Agostocaris bozanici, Bahamas, in January 1999. The new spe­ which exhibits the same unique pereiopodal cies was captured in Jumby Hole Cave morphology, placing it also in the Agosto­ (22°29.275'N, 73°53.501'W), Snug Corner, carididae. Holthuis (1993) synonymized the Acklins Island, Bahamas, 11 January 1999 Agostocariidae with the Bresiliidae. How­ (Fig. 1). This cave is located about 250 m ever, Martin & Davis (2001) have proposed inland from the west side of the island fac­ to recognize the family Agostocarididae ing the shallow water Bight of Acklins. It within the superfamily Bresilioidea Cai­ is actually a complex of closely associated man, 1896, where a hetereogeneous assem­ caves that were mined for guano in the past. blage of forms are included in five families. More than 3 m of soil and guano were re­ At best, as pointed out by Kensley moved from pits within dry portions of the (1988), the relationships of Agostocaris are cave. One of these caves contains a 20 m unclear. The particular articulation of the diameter, shallow (30 to 50 cm deep) pool. proprodus of the first pair of legs, and the Sediments in the pool consist of a thick lay­ morphology of the chela of the second pair er of guano from a bat roost located directly of legs, are unique characters not shared by above. Tidal range in the pool appeared to any other genus in the Bresilioidea. With be about 30 cm. This pool is in total dark­ respect to the diagnosis of Agostocaris, ness but is close to 4 or more entrances on with the new species described herein, the all sides. Salinity was measured at 32.5%c VOLUME 117, NUMBER 3 369 T?^ T7i0° I

GULF OF MEXICO

Ackllns Island

Liza Bay Cave

Fig. 1. Map showing tiie location of the type locality of Agostocaris acklinsensis, Acklins Island in the Bahamas. 370 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

with a refractometer and water temperature date and collector as holotype; CNCR was 25.5°C. Specimens of Agostocaris were 19602. observed walking across the surface of Description.—Carapace globose, smooth, rocks and the guano bottom in 50 cm depth. devoid of spines. Rostrum laterally com­ They were collected by hand using glass pressed, triangular, ending in sharp tip, vials. Other invertebrates collected from the reaching distal end of first antennular seg­ cave pools included copepods, archiannelid ment; without teeth in mature individuals, and other polychaetes, mites and the shrimp with three dorsal teeth with alternating setae Barbouria cubensis (von Martens, 1872) in juveniles (Fig. 2a, b). Carapace without (Hippolytidae). grooves, inferior margin of orbit and pter- The specimens representing the new spe­ ygostomian angle slightly produced (Fig. cies are deposited in the Coleccion Nacion- 2a), pterygostomian regions produced lat­ al de Crustaceos (CNCR), Institute de Biol- erally (Fig. 2b). ogia, Universidad Nacional Autonoma de Abdomen smooth, somites 1-2 with Mexico. Other abbreviations used are: cl, rounded pleura, somites 3-5 with posterior postorbital carapace length, and tl, total angle of pleura subacute, sixth somite with length. posterior margin sinuous at insertions of telson and uropods. Telson 2.5 times as Results long as its basal width, tapering distally, Agostocaris Hart & Manning, 1986 distal width less than half of basal width; bearing four pairs of movable spines on Diagnosis.—Rostrum well developed, dorsal surface, spines located on distal two with or without dorsal teeth. Carapace lack­ thirds of dorsal surface; posterior margin ing spines and grooves. Eyes reduced, rounded, bearing 9 spines, second pair from fused, without pigment or weakly pigment­ external one longest (Fig. 4g). ed. Antennal scale with lateral spine. First Eyes pigmented, fused, forming part of a maxilliped with lash on exopod. Second single plate, peduncle and cornea not dis­ maxilliped with terminal segments serial. cernible, projected dorsally (Fig. 2c). An- Pleurobranchs on all pereiopods or on pe- tennule with first segment as long as seg­ reiopods 2-5. First and second pereiopods ments 2 and 3 combined; stylocerite acute, chelate, first pair heavier than second one. reaching distal margin of first segment (Fig. First pereiopod with propodus articulating 4e). Antennal scale 1.8 times as long as with carpus at one third of its length. Sec­ wide, laterodistal tooth short not exceeding ond pereiopod with carpus undivided; dac- distal margin of blade (Fig. 4f), flagellum tylus digitiform, heavier and longer than 1.25 times total length (Fig. 2a). propodus, both fingers without teeth or Mandible with stout 2-segmented palp, spines. Telson with 4-5 pairs of dorsal incisor process with six distal teeth, molar spines, posterior margin with variable num­ process conical, sharp distal end (Fig. 2d). ber of spines. Both mandibles approximately symmetri­ cal. First maxilla with distal lacinia oval Agostocaris acklinsensis, new species shaped, bearing three rows of short, thick Figs. 2-4 setae on mesial surface; proximal lacinia with single row of short, thick setae on dis- Material examined.—Holotype, female, tomesial margin; palp bearing one distal, cl 7.3 mm, tl 21.5 mm; 11 January 1999; long setae and two subdistal short ones on Jumby Hole Cave, Snug Corner, Acklins Is­ internal margin (Fig. 2e). Second maxilla land, Bahamas; collected by T. M. Iliffe; with scaphognathite approximately rectan­ CNCR 19601. Paratypes, 8 females, cl 4.0- gular distally, subtriangular proximally; dis­ 8.0 mm, tl 13.6-21.7 mm; same locality, tal margin with long, plumose setae; lateral VOLUME 117, NUMBER 3 371

Fig. 2. Agostocaris acklinsensis, new species, a female holotype, b-f female paratype: a, total lateral view; b, carapace, dorsal view; c, dorsal view of eyes, carapace removed; d, mandible; e, first maxilla; f, first maxil- liped. Scale bar represent: a-c, f, 1 mm; d-e, 0.5 mm. 372 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Fig. 3. Agostocaris acklinsensis, new species, female paratype: a, second maxilla; b, second maxilliped; c, third maxilliped; d, first pereiopod; e, detail of propodus and dactylus of first pereiopod; f, second pereiopod. Scale bars represent: a-d, f, 1 mm; e, 0.5 mm. VOLUME 117, NUMBER 3 373

Fig. 4. Agostocaris acklinsensis, new species, female paratype: a, third pereiopod; b, detail of proximal segments of third pereiopod; c, fourth pereiopod; d, fifth pereiopod; e, antennule; f, antenna; g; telson and uropods, left side omitted; h, first pleopod; i, second pleopod. Scale bars represent 1 mm. 374 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

margin with short plumose setae; internal dactylus with corneous sharp tip and four margin with long simple setae, increasing smaller teeth on internal surface, arthro­ in length distally, almost as long as sca- branch and pleurobranch present, finger­ phognathite; palp digitiform, devoid of se­ like exopod arising from basis (Fig. 4a, b). tae; distal endite trapezoidal, middle and Fourth pereiopod with ischium with two proximal endites approximately rectangular, spines, merus the longest segment, carpus all three bearing simple setae on distal mar­ and propodus of about the same length, gins (Fig. 3a). dactylus with corneous sharp tip and three First maxilliped with triangular endite smaller teeth on internal surface, arthro­ bearing marginal setae; palp digitiform, branch and pleurobranch present, finger­ with apical tuft of setae; exopod elongated, like exopod arising from basis (Fig. 4c). bearing long, simple setae distally; caridean Fifth pereiopod with ischium with one lobe broadly rounded, with submarginal spine, propodus the longest segment, dac­ row of short setae and long plumose setae tylus with corneous sharp tip and eight along margin; epipod bilobed, both lobes smaller teeth on internal surface, arthro­ trapezoidal, distal one smaller, devoid of se­ branch and pleurobranch present (Fig. 4d). tae (Fig. 2f). Second maxilliped with en- First pleopod with exopod setose, endo­ dopod pediform, 4-segmented, with contin­ pod devoid of setae, one third the length of uous row of setae along margin; exopod exopod (Fig. 4h). Second pleopod with en­ slender, bearing long simple setae distally; dopod and exopod setose, appendix interna epipod simple, flat, rounded (Fig. 3b). Third slender more than half the length of endo­ maxilliped with endopod 4-segmented, pod (Fig. 4i). bearing setae on mesial margin; exopod as Uropods with external ramus bearing one long as first segment of endopod, with dis­ distolateral movable spine, distal margin tal tuft of long setae; epipod digitiform, less broadly rounded, with long plumose setae than half the length of exopod; arthro- on distal and internal margins. Internal ra­ branches present (Fig. 3c). mus bearing marginal long plumose setae First pereiopod with ischium and merus except on proximal third, distal margin sub­ of about same length and width, carpus acute (Fig. 4g). wider proximally, propodus articulating Etymology.—The specific name is de­ with carpus at one third of its length, palm rived from "Acklins", the name of the Ba­ as long as fingers, cutting edges of both fin­ hamian island where the new species was gers with minute sharp teeth, dactylus with captured. long setae arising from proximal half teeth (Fig. 3d); exopod as long as ischium and Key to the species of Agostocaris merus combined, with apical tuft of long setae; arthrobranch and pleurobranch pre­ \. First maxilliped with palp 2-segmented, sent (Fig. 3d). Second pereiopod longer ischium of fifth pereiopod devoid of than first one, with merus slightly shorter spines, outer ramus of uropods devoid of than ischium, carpus becoming wider dis­ distolateral spines, . . Agostocaris williamsi tally and as long as merus, propodus with - First maxilliped with palp unsegmented, palm shorter than fixed finger, dactylus ischium of fifth pereiopod with spines, outer ramus of uropods with spines, out­ heavier and longer than fixed finger; exo­ er ramus of uropods with distolateral pod shorter than ischium and merus com­ spines 2 bined, bearing apical tuft of long setae; ar­ 2. Ischium of fifth pereiopod with two throbranch and pleurobranch present (Fig. spines, outer ramus of uropod with two 3f). Third pereiopod with ischium with two distolateral spines, telson with five pairs spines, merus the longest segment, carpus of dorsal spines and propodus of about the same length. Agostocaris bozanici VOLUME 117, NUMBER 3 375

- Ischium of fifth pereiopod with one spine, outer ramus of uropod with one distolateral spine, telson with four pairs of dorsal spines . . Agostocaris acklinsensis .s ts

Remarks.—Agostocaris acklinsensis can (U D (U •" be easily distinguished from the other two o " o o D. "*- _ known species in the genus by the presence O O £ ^ (U (U « C M of: exopods on the third and fourth pereio- '• C pods, a fifth pereiopod with one spine on ^ 1 « the ischium and one distolateral movable c/i .-- •-« ° ^ i .^>^ >^ §^ ^ ^^ spine on the outer ramus of the uropods. « a & -r: "^ ^^ " I 5 •£ 2i CK Other taxonomically important characters Cu CO PQ •^ < ^ o vary among the three species. A second maxilla with a palp devoid of setae and an unsegmented palp of the first maxilliped distinguish A. acklinsensis from A. william- si, whereas the number of dorsal spines on the telson, unpigmented eyes and two dis­ tolateral spines on the outer ramus of the T3 o o O 'a _2J uropods seprate A. bozanici (Table 1). C/5 O a -o O o OH S u X X O o ^ u c This plate is projected outside the orbits 3 n c *V3 5 13 t^ c« U 3 creating the eye-like structures, which in m a PH PQ CO < ^ o the three species are pointed distally. Since all the species of Agostocaris are cave 60 dwellers it is reasonable to suppose that the cornea was lost and later the peduncle was c« reduced, in such a way that the "eyes" we u — c u see now are part of the basal plate. This O singular morphology merits further studies c/1 •- ^d t*oe on its ontogeny and functionality. 5 O X a 4-( •" 4- O ) O O cS The placement of the genus Agostocaris U -^ U (U o .5 is a matter of controversy. Holthuis (1993), S o > .3 bo x; U T3 by synonymizing Agostocarididae with the 4^ .5a 5^ X O ^ S '•n ^ Bresiliidae, gave more weight to characters -I (N ^ that are shared by many taxa in the Caridea •3 C^ § ^ OH d, PQ ^ o (mandible with palp, carpus of second legs undivided, first two pairs of legs chelate, first pair of legs more robust than second one, Williams, 1984) with little resolution S3 among families, than to exceptional auta- ft S o pomorphic characters such as the fused U --^ (U O O_H 5 O ::= a g- o eyes and the particular morphology of the X ;3 O o "? first two pereiopods of Agostocaris. X c.

silioidea, which includes five famiUes, and liam Keegan (Florida Museum of Natural concur with the opinion that this taxon still History) for providing invaluable logistical represents an artificial grouping. While it is information on Crooked and Acklins Is­ beyond the scope of this paper to discuss lands. The drawings were prepared by Ro­ the relationships among bresilioids, it is lando Mendoza. clear that Agostocarididae represents a dis­ tinct family that can be easily separated Literature Cited from the other four bresilioid families. The Caiman, W. T. 1896. On deep sea Crustacea from the Christoffersen, 1986, and south west of Ireland.—Transactions of the Mirocarididae Vereshchaka, 1997, lack ex- Royal Irish Academy 31:1-22. opods on all pereiopods, whereas the Agos­ Christoffersen, M. L. 1986. Phylogenetic relationships tocarididae can have exopods on all five pe­ between , Atyidae, , reiopods. The Diascididae Rathbun, 1902, Alvinocarididae fam. n., Bresiliidae, Psalido- popidae and (Crustacea Caridea have well developed eyes with peduncle Atyoidea).—Boletim Zoologico, Universidade and cornea, a dorsoventrally flattened ros­ do Sao Paulo 10:273-281. trum and a disc-like dactylus of the first pe- De Haan, W. 1849 (1833-1850). Crustacea. In P. F. von reiopod, contrasting with the fused eyes, Siebold, ed.. Fauna Japonica sive descriptio an- acuminate rostrum and typically shaped imalium, quae in itinere per Japonium, Jussu et dactylus of pereiopod 1 of the Agostocari­ auspices superiorum, qui summum in India Ba- tava imperium tenent, suscepto, annis 1823- didae. Finally the Bresiliidae, and the rest 1830 collegit, notis, observationibus et adum- of the bresilioid families, can be separated brationibus illustravit. Lugduni-Batavorum, 243 from the Agostocarididae based on the car- pp. pus-propodus articulation of the first pereio­ Hart, C. W., Jr., & R. W. Manning. 1986. Two new pod which is normal in the former, being shrimps (Procarididae and Agostocarididae, new family) from marine caves of the western the distal end of the carpus articulated to north Atlantic.—Journal of Biology the proximal end of the propodus; while in 6:408-416. the latter the carpus is articulated to an area Holthuis, L. B. 1993. The Recent Genera of the Car- close to the middle portion of the propodus. idean and Stenopodidean Shrimps (Crustacea, In addition, the chela of the second pereio­ ). Nationaal Natuurhistorisch Muse­ pod in the Agostocarididae is unique in that um, Leiden, 328 pp. Kensley, B. 1988. New species and records of cave the digitiform dactylus is longer than the shrimps from the Yucatan Peninsula (Decapoda: fixed finger and lacks teeth or spines. Agostocarididae and Hippolytidae).—Journal of Crustacean Biology 8:688-699. Martin, J. W., & G. E. Davis. 2001. An updated clas­ Acknowledgments sification of the recent Crustacea. Natural His­ tory Museum of Los Angeles County, Science Collection of shrimp described herein Series 39, 124 pp. was part of the January 1999 Anchialine Rathbun, M. J. 1902. Papers from the Hopkins Stan­ Caves Expedition to the southern Bahamas ford Galapagos Expedition 1898-1899. VIII. led by Thomas Iliffe. Other members of the Brachyura and Macrura.—Proceedings of the Washington Academy of Sciences 4:275-292. expedition included Texas A&M University Vereshchaka, A. L. 1997. A new family for a deep-sea graduate students Brett Dodson and Shelley caridean shrimp from North Atlantic hydrother- Fetterolf. This expedition was funded by mal vents.—Journal of the Marine Biological National Science Foundation, Biotic Sur­ Association of the United Kingdom 77:425- veys and Inventories Program award num­ 438. Williams, A. B. 1984. Shrimps, lobsters, and crabs of ber 9870219. We thank Neil Sealey (Media the Atlantic coast of the eastern United States, Publishing Ltd, Nassau, Bahamas), Dr. Maine to Florida. Smithsonian Institution Press, Nancy Elliott (Sienna College) and Dr. Wil­ 550 pp.