Contributions to Zoology, 72 (4) 227-252 (2003)

SPB Academic Publishing bv, The Hague

from Great Three new sympatric species of (Crustacea) Exuma

Island, Bahamas Islands

Stefan Koenemann Thomas+M. Iliffe² & Joris van der Ham³ ¹,

1 Institute of Biodiversity and Ecosystem Dynamics, University of Amsterdam, P.O. Box 94766, 1090 GT

2 Amsterdam, The Netherlands, corresponding author ([email protected]); Department of Marine

3 Biology, Texas A&M University at Galveston, Galveston, TX 77553-1675, USA; Department of Biology,

University ofLouisiana at Lafayette, 300 East St. Mary Blvd., Lafayette, LA 70504, USA

anchihaline Keywords:: Crustacea, Remipedia, Speleonectidae, Speleonectes, new species, Bahamas, caves,

sympatry, biogeography, evolutionary history

Abstract Introduction

Three new sympatric species of remipede , Speleo- To date, the class Remipedia Yager, 1981, nectes tanumekes,Speleonectesparabenjamini and Speleonectes is composed of two families, the Speleonectidae minnsi, are described from an anchihaline cave onGreat Exuma Yager, 1981, including nine species in three gen- Island in the central Bahamas. Speleonectes tanumekes is a and the & era, Gpdzilliidae Schram, Yager Emerson, comparatively long and slender species distinguished by the 1986, with three in three The first largest number of trunk segments found in remipedes to date. species genera.

Speleonectes parabenjamini is morphologically closely related remipede was discovered in an anchihaline cave to Speleonectes benjamini, but differs from the latter species on the Bahamas Islands (Yager, 1981). Between by several distinct autapomorphies. Speleonectes minnsi is cha- 1980 and 1999,11 additional species were described. racterized by comparatively robust maxillules. The occurrence All currently known Remipedia inhabit subtropi- ofthree sympatric species is a remarkable record for the Remi-

cal anchihaline caves. Their distribution is pedia adding to a total of 10 sympatric taxa. We discuss the disjunct, high diversity of remipedes in the larger West Indian region comprising recorded localities in the larger Carib- with to their evolutionary history and origin. Ca- regard bean region, western Australian, and on the

nary Islands.

We describe three new sympatric species of remi-

Contents pedes, Speleonectes tanumekes, Speleonectes para-

benjamini and Speleonectes minnsi, including de-

Introduction 227 tailed drawings and SEM images. The new species Abbreviations and definition of terms 227 were found during a recent diving expedition on Systematics 228 Great Exuma Island located in the central Baha- Remipedia 228 All collected from the Nectiopoda 228 mas. specimens were same

Speleonectidae 228 anchihaline cave between 33 and 43 m depth. The Speleonectes 229 implications of this new record are discussed with tanumekes 229 Speleonectes n. sp. regard to the biogeography and evolutionary his- 235 Speleonectes parabenjamini n. sp. tory of the Remipedia. Speleonectes minnsi sp. 241

Ecological profile of the type locality 247

Some remarks on the biogeography and evolutionary

history of Remipedia 247 Abbreviations and definitions of terms Acknowledgements 252

References 252

JvdH Joris van der Ham

ZMA Zoological Museum Amsterdam

Elbow Term referring to the main point of flexure in

maxillules, maxillae and maxillipeds.

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Tagmosis of maxillae and maxillipedsis a condition characterized fossils Tesnusocaris goldichi Brooks, 1955, and distinct size by and shape differences between the segment Cryptocaris hootchii Schram, 1974, as subordinate proximal to the elbow and those distal to the elbow. In tagmatized taxa. appendages, the proximal segment is much longer and wider

than the distal segments (see Fig. I2B, C). If tagmosis is com- Order 1986 pletely lacking, the segments proximal and distal to the elbow Nectiopoda Schram,

are subequal in width in length (Fig. 3D). Nectiopoda Schram, 1986: 36. Schram, Yager & Emerson,

Heteromorphic sternal bars are differently shaped transverse 1986: 6.

bars sternites. The male of herma- on some trunk gonopores the

phroditic remipedes are located on trunk 14. In some segment - Diagnosis. Cephalon with bifurcate, pre-anten- the sternal bars this taxa, on segment are modified as enlarged, nular ventral filaments. Antennular peduncle fused, flap-like structures (see Fig. 8B). However, heteromorphy can with aesthetascs. Antennae also serial sternal bars 1-14 long, filamentary small, occur as a modification, e.g., can bent have a concave distal margin, while those on the posterior trunk paddle-like; endopod 3-segmented, or as semi-

are convex. circular Mandibles arc; exopod 1-segmented paddle.

Isomorphic sternal bars are typically sublinear (with paral- with 6-7 lacking palps. Maxillules segments; ter- lel and thus, modified trunk margins) subequal, not on segment minal with segment developed as fang-like claw, a 14 other trunk or any segment (see Fig. 11D). single pore at the tip. Maxillae and maxillipeds

subequal; terminal segments developed as complex

cluster Systematics claws equipped with a or arc of denticles

thumb-like typically opposed by a pad bearing a

In the with following sections, we provide an outline of row of filamentary, flexible setae. Maxillae

the taxonomic structure of the class Remipedia. three lobate endites on basal segment. Maxillipeds

Although we consider the inclusion of higher taxa 9-segmented, with 5 segments distal to elbow; en-

for important revised diagnoses of lower catego- dites of basal segments reduced, pad-like. Trunk

ries this (in case Speleunectes ), diagnoses are omitted limbs with 3-segmented exopod; endopod 4-seg-

for if arc not relevant for our basal anterior taxa they compari- mented, segment reduced; and pos-

sons. See Remarks for additional information. terior trunk limbs more slender than those of mid-

Trunk counted the first segments are beginning at trunk, armature more reduced. Anal segment with post-cephalic segment (not including maxillipeds). simple caudal rami.

of are numbered from Segments appendages proxi- Remarks. - The Nectiopoda include two families, mal distal. to the Godzilliidae Schram, Yager & Emerson, 1986,

and the Speleonectidae Yager, 1981. The maxilli-

Class 1981 number of Remipedia Yager, peds of a nectiopods are described as Remipedia Yager, 1981: 328. Schram, Yager & Emerson, being composed of 8 segments. However, a rein- 1986: 6. vestigation of collection and type material did not

confirm this. Maxillipeds appeared to be 9-seg- Diagnosis. - Hermaphroditic crustaceans with 6- mented in all specimens evaluated, although some segmented cephalon (including maxillipeds), head taxa in both families, Speleonectidae and God- Trunk shield present. lacking tagmosis, composed zilliidae, show a tendency to reduce the size and of at least 15 segments; carapace absent; female articulation of the basal segments. on trunk male gonopores segment 7, gonopores on

trunk segment 14. Antennules and antennae bi- Family Speleonectidae Yager, 1981 Post-oral ramous. Labrum well developed. cepha- Speleonectidae Yager, 1981:328. Schram, Yager & Emerson, lic appendages modified as subequal, uniramous, 1986: 6.

Transverse ster- prehensile, raptorial mouthparts. MorlockiidaeGarcia-Valdecasas, 1984: 329.

nal bars present. Trunk limbs as biramous, paddle-

shaped swimming appendages. Type genus. - Speleonectes Yager, 1981.

Remarks. - The class includes two Remipedia or- Diagnosis. - Head shield subrectangular, often

ders, the Nectiopoda Schram, 1986, and the Enantio- slightly tapered anteriorly. Transverse sternal bars Birshtein, 1960. The latter order contains the poda of trunk isomorphic or heteromorphic. Ventral ra-

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mus of antennules well developed, typically with Speleonectes tanumekes new species 7-15 segments. Mandibles asymmetrical. Maxillules Figs. 1-3; PI. 1

7-segmented, with main point of flexure between

3 and 2 with lobate - segments 4; segment large endite, Type locality. Basil Minns Blue Hole, Great Exuma

with distoanterior and dis- equipped row of spines Island, Exuma Cays, The Bahamas (23° 28’ N, 75°

toposterior row of endite of 3 well 45’ setae; segment W); collected in 33-43 m depth of anchihaline but shorter than endites developed, 1 and 2; me- cave.

dial margin of segment 4 expanded. Maxillae 7-

segmented; distal elbow segments to to Material examined. - subequal Holotype (27 mm, 40 trunk that of maxillipeds, but shorter; main of flex- ZMA 1 point segments; Rem.204.577), paratype (20 mm, between 3 ure segments and 4. Maxillary and maxil- 38 trunk ZMA 10 segments; Rem.204.574), para-

lipedal distal to elbow 26 segments typically distinctly types (up to mm and 40 trunk segments; JvdH

than to elbow. longer segment proximal 12-01 A1-3); all specimens were collected from

Remarks. - The family Speleonectidae is composed the type locality by B. Kakuk and T. Iliffe, 12 Jan. of the three genera Cryptocorynetes Yager, 1987a, 2003. The and 1 holotype (whole specimen) paratype Lasionectes & Yager Schram, 1986, and Speleo- (dissected) are preserved in alcohol. The remain-

nectes 1981. Yager, ing paratypes are preserved in formalin and will be

retained in the research collection of the first au-

Genus 1981 Speleonectes Yager, thor Some are (JvdH). of the paratypes prepared Speleonectes Yager, 1981: 1994: 752. 328; Schram, Yager for SEM investigations. & Emerson, 1986: 6.

Morlockia Garcia-Valdecasas, 1984: 329. Etymology. - The epithet tanumekes (Greek for

refers to the and slender ‘long-stretched’) long ap- Type species. - Speleonectes lucayensis Yager, pearance of this species. 1981.

Diagnosis. - Maxillules typically more robust than

Diagnosis. - and slender to 27 and Long species, up maxillae maxillipeds; endite of segment 3 bear- mm, largest specimens composed of 38-40 trunk ing 2 prominent apical spines and several setae. segments (Fig. 1 A); pleural tergites weakly devel- Tagmosis of maxillae and maxillipeds weakly de- oped, with rounded lateral in setation broadly margins an- veloped; of segment proximal to elbow and terior of part trunk, becoming slightly pointed pos- those distal to elbow composed of long and short teriorly; sternal bars sublinear, isomorphic; frontal setae or spines, sparsely distributed, not covering filaments with short processes; dorsal flagella of more than about 75% of medial margins; terminal antennules short; segment 4 of maxillule subrec- claws as complex, horseshoe-shaped scrapers equip- tangular, with small enditic lobe 2 with of bearing long, ped an arc small, serrate denticles accom- rasp-like spines and several slender setae; maxilla 1 2 panied by or prominent, separate denticles. and maxilliped without distinct tagmosis, bearing Remarks. - The of discovery three new species of few clusters of described long, marginal setae; horseshoe-type remipedes herein reveals a number of claws of maxilla and maxilliped with 17-20 small inconsistencies concerning the diagnoses of the denticles; anal somite slightly wider than and the long; family Speleonectidae genera Speleonectes caudal rami slightly longer than anal somite. and Lasionectes. In particular, the tagmosis and

setation ofthe three prehensile cephalic limbs needs Description. - Body distinctly elongate and slen- to be redefined for both genera. A comprehensive der, with a maximum of 27 and revision of length mm up to Remipedia is in preparation by one of 40 trunk I segments (Fig. A). Pleural tergites nar- us (SK), which will address these inconsistencies with row, broadly rounded distolateral corners on in greater detail. trunk segments 1-16, becoming slightly acuminate

in posterior part of trunk. Sternal bars sublinear

and Female on trunk isomorphic. gonopores seg-

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I. 24 seventh 34th Fig. Speleonectes tanumekes ft. sp.; A, mm paratype; B-F, 20 mm paratype. B, first thoracopod;C, thoracopod;D,

thoracopod; E, feathered seta; F, serrate spine. Scale bar = 0.5 mm, for B-D.

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2. tanumekes 24 20 mm A, frontal filaments bar = 0.1 Fig. Speleonectes n. sp.; A, mm paratype; B-F, paratype. (scale mm); B,

= = antennule (scale bar = 0.5 mm); C, antenna (scale bar = 0.1 mm); D, labrum (scale bar 0.1 mm); E, right mandible (scale bar 0.1

lacinia mobilis and incisor of left mm); F, enlarged process tnandible.

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20 left Fig. 3. Speleonectes tanumekes n. sp.; mm paratype. A, right maxillule; B, maxillule; C, maxilla; D, maxilliped; E, rasp-like

spines ofsegment 2 (small spine), and segments 3-4 (large spine) of maxillule; F, claw ofmaxillae and maxillipeds; G, anal segment

= and caudal rami (scale bar 0.25 mm). Scale bar A-D = 0.5 mm.

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PI. I. lamimekes ventral view ofhead claw ofmaxilla detail of mandible Speleonectes n. sp., paratype. A, (60x); B, (850x); C, right

detail of left mandible female male (500x); D, (500x); E, gonopore (IGOOx); F, gonopore (1200x).

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ment 7 with tiny, triangular projections (PL IE); setae on distal margins. Claw well developed.

rounded lobes Maxillae Endites first male gonopores with ovate, (PL IF). (Fig. 3C): of segment equip-

with few Cephalon subrectangular, tapered anteriorly, as ped 1 apical spine accompanied by a long long as trunk segments 1-4 (Fig. 1A). Frontal fila- and short setae, respectively. Endite of segment 2

with short mid-medial with short and ments processes (Fig. 2A; broadly rounded, a single, spine a

PL 1A). few setae. Third segment with weakly expanded,

2 about 8 Antennules (Figs. 1A, 2B): Peduncle composed straight inner margin bearing rows of of at least 2 fused segments; proximal component long and 2 short setae. Segment 4 shorter than third

distal with cluster of subrectangular (barely dilated ventrally) bearing segment; margin expanded, a dense rows of long aesthetascs. Dorsal flagellum ca. 4 long setae and I short seta on inner margin, relatively short, about twice as long as head shield and 2 short setae on dorsal margin. Fifth segment and 7% of of with 13 shorter than but length body; up to segments; segment 4, not expanded distally, proximal-most segment apparently partially fused setation subequal to that of segment four. Segment to peduncle. Ventral flagellum with 7-8 segments, 6 slightly shorter than fifth segment, equipped with

less than half as long as dorsal flagellum, slightly separate clusters of setae distally, and a few longer

midmedial shorter than head shield. setae on margin. Arc of horseshoe-type

Antennae (Fig. 2C): Protopod 2-segmented, distal claw finely serrate, composed of 17-20 fine den- segment with row of7-8 setae. Exopod longer and ticles flanked by 2 stouter, separate denticles (PL wider than adjacent distal segment of protopod, IB; Fig. 3F).

in bearing 26-28 long setae. Endopod bent a semi- Maxillipeds long, slender, 9-segmented (Fig. 3D); circular first with 8 elbow between 4 and 5. Proximal way; two proximal segments segments seg- and 7 setae, respectively; distal segment with 15- ments 1-3 with oblique, interconnected articulation;

17 setae arranged in two rows (11-12 + 4-5 setae). segments 1 and 2 bearing a few short medial setae;

All setae faintly feathered (see Fig. IE). segment 3 broadly rounded medially, with about

Labrum fleshy, with posterolateral lobes bear- 6-7 long and 2 short setae. Fourth segment with ing several clusters of fine setules (Fig. 2D). weakly expanded, straight ventral margin equipped

incisor Mandibles (Fig. 2E, F): Right process with with some 8-7 long and 2 short setae proximally. three lacinia mobilis 5-7 in large denticles; right equipped Segments gradually decreasing length; seg-

5 with three larger and several smaller denticles (Fig. ment slightly shorter than segment 4, distal mar-

3E; PL 1C). Left incisor process with four large gin weakly expanded, with a medial cluster of about

denticles; left lacinia mobilis crescent-shaped, bear- 6 long and short setae. Segment 6 bearing a disto- ing several large and small denticles (Fig. 3F; PL medial cluster of 8 short and long setae. Segment

1D). Molar processes prominent; distal surface long, 7 equipped with 9-12 short and long setae along ovoid. mid-to distomedial margin. Segment 8 longer than

Maxillules (Fig. 3A, B): First segment equipped segment 7, with several separate clusters of setae 6-7 and with long, narrow endite bearing naked, slen- on distal distomedial margins. Claw subequal der Endite of 2 to that of maxillae spines distally. segment broad, spa- (PL IB; Fig. 3F). tulate, with 5 setae along distoanterior margin, and Trunk appendages (Fig. 1B-D; description based

10 short spines on distoposterior margin; all short on larger thoracopods): Segment 1 ofexopod with spines naked except 1 rasp-like spine (Fig. 3E), long setae on lateral margin, and about 3 serrate

inserted anteriorly adjacent to row of setae. Third spines on distolateral corner (Fig. 1C); segment 2 segment short, endite broadly rounded, bearing 2 bearing setae on both lateral and medial margins, long, slender, rasp-like spines (Fig. 3E), and a few and serrate spines on both distolateral and -medial

enditic lobe setae. Segment 4 subrectangular, small, corners; segment 3 ovate, with long, marginal se- with 2 long, slender, rasp-like spines, and a few tae. Endopod slightly shorter thanexopod, but simi-

5 shorter and in and setae. Segment slightly narrower than lar shape setation. All setae faintly feath- fourth segment, with a row of distomedial setae. ered (Fig. IE).

Sixth of Anal segment very short, bearing separate rows segment slightly wider than long, lateral

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few short margins bearing a setae and spines (Fig Etymology. - The specific epithet parabenjamini, caudal rami 1.1-1.2 times than anal 3G); longer meaning ‘nearby benjamini’, alludes to the mor-

somite, with a few setae on curling medial and apical phological resemblance of this new species to Spe- margins, respectively. leonectes benjamini.

Remarks. - Speleonectes tanumekes has the larg- Diagnosis. - A relatively small species (Fig. 4A);

est numberoftrunk for segments recorded remipedes pleural tergites well developed, broadly rounded,

to date. The number of in large segments, combi- becoming increasingly pointed in posterior part of nation with weakly developed pleurotergites, gives trunk; trunk sternites with pointed posterolateral

this a rather and slender sternal bars species elongate appear- corners; heteromorphic; frontal filaments

ance. Both S. Yager & with epilimnius Carpenter, 1999, short, stout processes; antennules with long and S. gironensis Yager, 1994, also have narrow dorsal flagella; segment 4 of maxillule subrectan-

but are characterized a smaller 1 pleurotergites, by gular, bearing very long, serrate spine and sev-

trunk number of segments to 21 and 25 eral slender claw (up seg- setae; maxillulary very long; max- In S. ments, respectively). addition, tanumekes can illa and maxilliped slender, without distinct tagmosis,

be from S. dif- of clearly distinguished gironensis by setation sparse; arc horseshoe-type claw com-

fering shapes and setations of maxillae of 7-8 small maxillules, posed denticles; anal somite longer and maxillipeds. Speleonectes tanumekes seems than wide; caudal rami shorter than anal somite.

morphologically similar to S. epilimnius, but dif-

fers from the latter - species by having a 13-segmented Description. Body short, up to 13 mm, composed

dorsal antennular flagellum (10-11 in S. of 24 segments postcephalic segments (Fig. 4A). Pleural terg- epilimnius)', a relatively large antennal exopod; ites well developed, with broadly rounded disto- and slender longer more on corners spines maxillulary seg- lateral on trunk segments 1-7, becoming ments 3 and 4; less expanded, straight margins of increasingly acuminate posteriorly. Posterolateral 3 and 4 (proximal to elbow) of maxillae corners segments of trunk sternites with distinctly pointed and maxillipcds, (more expanded and respectively corners (Fig. 8B; PI. 2C). Sternal bars heteromor-

rounded in S. and an anal wider epilimnius)-, segment phic, slightly concave on segments 1-13, posterior

than long, with caudal rami 1.2 times than to 14 longer segment as small, convex flaps; segment 14 the anal somite than (anal somite wide, with with convex longer large flap (PI. 2C). Male gonopores

caudal rami 2-2.5 times in trunk longer S. epilimnius). on segment 14 with rounded lobes.

Cephalon subrectangular, with concave antero-

Speleonectes new lateral trunk 1-4 parabenjamini species excavations, as long as segments

Figs. 4-8; PI. 2 4A). Frontal filaments (Fig. bearing short, corpu-

mid-medial lent, processes (Fig. 5A). Type locality: Basil Minns Blue Great Exuma Antennules with Flole, long dorsal flagella, reaching Exuma The Bahamas Island, Cays, (23° 28’ N, 75° up to 60% of length of body (Figs. 4A, 5B). Pe-

45’ collected in 33-43 m of anchihaline duncle ofat least 2 fused W); depth composed segments; ven-

cave. tral margin expanded, bearing densely inserted, long

with aesthetascs. Dorsal flagellum 14 segments.

Material examined: Flolotype (9 22 trunk Ventral with 15 mm, seg- flagellum up to segments, less than

ZMA Rem.204.579) and 1 as ments; paratype (13 mm, half long as dorsal flagellum, but slightly longer 24 trunk segments; JvdH 12-01 Al); collected by than head shield.

B. Kakuk and T. Iliffe, 12 Jan. 2003. The holotype Antennae(Fig. 5C): Protopod 2-segmented, proxi-

is dissected and in alcohol. The with preserved paratype mal segment equipped 7 marginal setae; dis-

is prepared for SEM investigations and will be tal with 11 segment setae. Exopod longer and wider retained in the research collection of the first au- than distal adjacent segment of protopod, bearing thor (JvdH). about 37 long setae. Endopod forming a semicir-

cular first two arc; proximal segments with 6-8 setae,

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13 9 first Fig. 4. Speleonectes parabenjamini n. sp.; A, ram paratype; B-E, mm holotype. B, thoracopod;C, seventh thoracopod; D,

of and Scale bar = for thoracopod posterior trunk; E, rasp-like spine (left) serrate spine (right). 0.5 mm, B-D.

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5. 13 9 filament bar = 0.1 Fig. Speleonectes parabenjamini n. sp,; A, mm paratype; B-C, mm holotype. A, frontal (scale mm); B,

antennule (scale bar = 0.5 mm); C, antenna (scale bar = 0.5 mm).

respectively; distal segment with up to 17 setae of segment 2 ovate, spatulate, with 4-5 long setae

arranged in two rows. All setae faintly feathered on distoanterior margin, and 8 short spines and 1

(cf. Fig. IE). short seta on distoposterior margin (all short spines

Labrum with clusters of fine setules naked 1 at end of fleshy, on except rasp-like spine spine row;

posterior margin (Fig. 6A). Fig. 7D). Third segment short, with conical endite,

2 Mandibles (Fig. 6B, C; PI. 2A): Right incisor bearing long, slender, rasp-like spines, and a few

process and lacinia mobilis with three large den- setae. Segment 4 subrectangular, medial margin

Left incisor with four 1 ticles; respectively. process even, bearing very long, slender, rasp-like spine

large denticles; left lacinia mobilis crescent-shaped, on proximal corner (Fig. 7E), and 7-9 long and short

Molar 5 but apical margin serrate. processes prominent; setae. Segment as long as segment 4, slightly distal surface long, ovoid. narrower, with a cluster ofdistomedial setae. Sixth

Maxillules little maxillae and a more robust than segment short, equipped with separate rows of long

First and shorter on distal Claw maxillipeds (Fig. 7A, D-E). segment equipped setae margins. very long.

with and endite 1 Maxillae long narrow bearing large, promi- (Fig. 7B): First endite of first segment and 1 nent spine 5-6 small spines (all naked). Endite bearing apical spine and a few setae; endites 2

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Pi 2. detail of left mandible claw of trunk Speleonectes parabenjamini n. sp., paratype. A, (500x); B, maxilliped (IOOOx); C, sternites

and sternal bars (sb) of trunk segments 2-4 (pointed posterolateral corners indicated by arrows) (90x).

and 3 with 1 prominent, apical spine, 3 smaller by 2 stronger, separate denticles (Fig. 7F).

and several Endite subapical spines, setae. of seg- Maxillipeds long, slender, 9-segmented (Fig. 7C);

short and elbow ment2 equipped with a single, spine about between segments 4 and 5. Segment 1 bear-

9 with 2 short and long setae. Third segment long, ing a few short medial setae; segment equipped

barely expanded, straight medial margin bearing with 2 slender medial spines; segment 3 rather long,

10-12 long and short setae. Segment 4 shorter than with a cluster of distomedial long and short setae.

distomedial with of 10- 4 and of segment 3; margin a row Segment very long narrow, bearing a row

12 de- setae. Fifth segment half as long as segment 4, 7-8 midmedial setae. Segments 5-7 gradually

bearing about 6 setae on distomedial margin. Seg- creasing in length; segment 5 a little shorter than

ment 6 distinctly longer than fifth segment, with 4 segment 4, distal margin expanded, with a cluster

sparsely inserted setae along medial margin, and of distomedial setae. Segments 6-8 bearing rows

clusters of Arc of horseshoe- of mid- and distomedial separate setae distally. setae on margins, respec-

type claw composed of7-8 small denticles flanked tively. Segment 8 distinctly longer than segment

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labmm scale bar = 0.5 left Fig. 6. Speleonecles parabenjamini n. sp.; 9 mm holotype. A, (partly damaged; mm); B, mandible, with

bar = detail of mandible. Please that the of slightly enlarged incisor process (scale 0.1 mm); C, enlarged right note angled appearance

the molar processes is an artefact caused by preservation.

7, with several separate clusters of setae on distal spines (Fig. 4E); distolateral corner of segment 2

margins. Claws subequal to those of maxillae (Fig. bearing 4 rasp-like spines and 1 strongly serrate

7F; PI. 2B). spine, distomedial corner with 2 serrate spines;

Trunk appendages (Fig. 4B-D): Segment 1 of segment 3 with 2 serrate spines on distolateral comer

exopod equipped with long setae on lateral mar- and 1 serrate spine on distomedial corner. Endopods

of gin, and about 3 serrate spines on distolateral cor- anterior and posterior trunk limbs comparatively

2 ner (Fig. 4C, E); segment with setae on both lat- strongly reduced. All setae faintly feathered (cf.

eral and medial margins, and 2-4 serrate spines on Fig. IE).

distolateral corner; segment 3 ovate, bearing long Anal segment longer than wide (Fig 8A); length

marginal setae. Endopod slightly shorter and nar- of caudal rami about 66% of length anal somite;

rower than exopod; distribution of feathered setae caudal rami bearing 1-2 curling setae on midmedial

similar that of with the 6 to exopod, following ex- margins up to curling apical setae, respectively.

ceptions: basal segment with 3-4 rasp-like, lateral

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7. 9 with of Fig. Speleonectesparabenjaminin. sp.; mm holotype. A, maxillule; B, maxilla, enlarged spine segment2; C, maxilliped:

D, enlarged maxillulary endite of segment 2; E, long rasp-like spine of maxillulary segment 4; F, claw of maxillae and maxillipeds.

Scale bar A-C = 0.5 mm.

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8. 9 anal and caudal rami bar = 0.1 sternites and Fig. Speleonectes parabenjamini n. sp.; mm holotype. A, segment (scale mm); B, sternal bars oftrunk segments 2, 3, 13 and 14 (scale bar = 0.5 mm).

is Remarks. - Material examined: 30 trunk Speleonectes parabenjamini morpho- Holotype (18 mm,

S. 1987a. ZMA B. Ka- logically closely allied to benjamini Yager, segments; Rem.204.580) collected by

Both species share several distinct apomorphies, kuk and T. Iliffe, 12 Jan. 2003. The holotype is

for example, antennules with long dorsal flagella; dissected and preserved in alcohol.

maxillules with rather long claws; trunk sternites with and hetero- Etymology. - Speleonectes minnsi is named after pointed posterolateral corners; Basil who first information the some Minns, provided on morphic sternal bars. However, of these syn-

cave assigned as locality for this species. apomorphies are also characterized by interspecific type

differences. Unlike S. benjamini, S. parabenjamini

Diagnosis. - A small to medium-sized, slender has less sharply pointed sternites, and the shape of spe- cies (Fig. 9A); pleural tergites developed, broadly the anterior sternal bars is only slightly concave. rounded, becoming angular in of trunk; dis- posterior part Furthermore, S. parabenjamini can be easily sternal bars isomorphic; frontal filaments with long tinguished from S. benjamini by possessing ven- processes; antennules with short dorsal flagella; tral antennular flagella that lack a fusion of proxi- 4 segment of maxillule expanded; maxilla and maxil- mal segments; maxillules and maxillae that are more liped exhibiting tagmosis (see definitionsof terms); slender (less expanded), with much longer and slen- arc of horseshoe-like claw composed of 7 small der segments proximal to the elbows; an anal seg- denticles; anal somite longer than wide; caudal rami mentthat is longer than wide; and caudal rami shorter shorter than anal somite. than the anal segment.

Description. - Body slender, length 18 mm, 30 trunk minnsi Speleonectes new species segments (Fig. 9A). First trunk segment much smal- Figs. 9-12 ler Pleural than adjacent posterior segments. terg- ites developed, with broadly rounded distolateral Type locality: Basil Minns Blue Hole, Great Exuma trunk corners on segments 1-10, becoming angular Island, Exuma Cays, The Bahamas (23° 28’ N, 75° to pointed posteriorly. Sternal bars sublinear, iso- 45’ W); collected in 33-43 m depth of anchihaline morphic (Fig. 11D). Male gonopores on trunk seg- cave. ment 14 with small triangular lobes.

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18 dorsal of first seventh 27th Fig. 9. Speleonectes minnsi n. sp.; mm holotype. A, view body; B, thoracopod; C, thoracopod; D,

setulose Scale bar = 0.5 for B-D. thoracopod; E, serrate spine; F, spine. mm,

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10. minnsi 18 filaments bar = 0.1 antennule = 0.5 Fig. Speleonectes n. sp.; mmholotype. A, frqjital (scale mm); B, (scale bar mm); C,

bar = 0.1 mandible bar = 0.1 lacinia mobilis and incisor antenna (scale mm); D, right (scale mm); E, enlarged (left) process (right) of

and incisor right mandible; F, enlarged lacinia mobilis (left) process (right) of left mandible.

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II. 18 = 0.1 of of Fig. Speleonectes minnsi n. sp.; mm hoiotype. A, labrum (scale bar mm); B, detail apical margin labrum; C; anal

segment and caudal rami (scale bar = 0.5 mm); D, sternal bars 4, 13 and 18 (from top).

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12. minnsi 18 endite of Fig. Speleonectes n. sp.; mm holotype. A, maxillule; B, maxilla; C, maxilliped; D, enlarged maxillulary

setulose and 4 claw of segment 1; E, spines ofmaxillulary segments 3 (left) (right); F, maxillae and maxillipeds. Scale bar A-C = 0.5 mm.

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with distomedial Sixth Cephalic shield slightly trapezoidal, sinu- setae. segment short, bearing

soid lateral margins, as long as trunk segments 1- separate rows of setae on distal margins. Claw well

4 (Fig. 9A). Frontal filaments equipped with long developed.

mid-medial Maxillae First endite of basal processes (Fig. 10A), reaching up to (Fig. 12B): seg-

distal tips of filaments, apparently articulated (see ment bearing 1 apical spine and 2 setae; endites 2

Remarks). and 3 with 1 prominent apical spine, 2-3 smaller

Antennules with short dorsal about 20% and several Endite of flagella, subapical spines, setae. seg-

of length of body (Figs. 9A, 10B). Peduncle com- ment 2 equipped with a single apical spine and about posed of at least 2 fused segments; ventral margin 6 long subapical setae. Third segment long, with

expanded, broadly rounded, with densely inserted, pear-shaped medial margin, bearing rows of densely

long aesthctascs. Dorsal flagellum 12-segmented. inserted long and short setae. Segment 4-6 increas-

Ventral flagellum less than half as long as dorsal ingly shorter, bearing rows of setae on midmedial

shorter than head of 8 and distomedial medial of flagellum, shield, composed margins; margin seg-

segments, of which the proximal-most 2 segments ment 5 almost entirely covered with setae. Arc of

7 are fused. horseshoe-like claw composed of small denticles

Antennae flanked 2 more robust denticles (Fig. 10C): Protopod 2-segmented, pro- by (Fig. 12F).

ximal segment with about 5 marginal setae; distal Maxillipeds long, slender, 9-segmented (Fig.

segment with 7-8 setae. Exopod longer and wider 12C); elbow between segments 4 and 5. Segment

of with cluster of than adjacent distal segment protopod, equipped 1 a 5 medial setae; segment 2 short,

2 with 27 long setae. Endopod forming a semicircu- with medial setae; segment 3 bearing 2 clusters

lar arc; first two segments with 7 and 8 setae, re- of short and long, distomedial setae. Segment 4 long,

spectively; distal segment with about 20 setae ar- medial margin pear-shaped, bearing rows of densely

in ranged two rows (composed of ca. 14 and 5-6 inserted setae. Segments 5-7 gradually decreasing

setae, respectively). All setae faintly feathered (cf. in length; segments 7 and 8 subequal in length; distal

of 5 setations of Fig. IE). margin segment expanded; seg-

Labrum fleshy, with row of fine setules on pos- ments distal to elbow similar to those of maxillae. terior, apical margin (Fig. II A). Claws subequal to those of maxillae (Fig. 12F).

Mandibles incisor Trunk of (Fig. I0D-F). Right process appendages (Fig. 9B-D): Segment 1

lateral and lacinia mobilis with three large denticles; re- exopod equipped with long setae on mar-

spectively. Left incisor process with four large denti- gin, and about 2 serrate spines on distolateral cor- cles; left lacinia mobilis crescent-shaped, apical ner (Fig. 9E); segment 2 with setae on both lateral

Molar and medial and margin serrate. processes prominent. margins, up to 4 serrate spines on

Maxillules much robust than maxillae and distolateral 3 bear- more corner; segment distally tapered,

maxillipeds (Fig, 12A). First segment equipped with ing long marginal setae (segment 3 of posterior trunk

long and narrow endite bearing 5 spines, one of limbs round, paddle-shaped). Setation of endopod which Endite of similar that of with weakly serrate (Fig. 12D). seg- to exopod, the following ex- ment 2 ovate, spatulate, with about 3 long setae ceptions: Limbs from mid-trunk posteriad lacking

and 5-6 short 1 along distoanterior margin, spines feathered setae on proximal segments; segment on distoposterior margin (all short spines naked and 2 with setulose spines on distolateral and -medial

setulose end of with except 1 spine at spine row; Fig. comers (Fig. 9F); segment 3 2 serrate spines

Third with distolateral and -medial 12E). segment short, cone-shaped endite, on corners, respectively. bearing 2 long, slender, setulose spines (Fig. 12E) Larger (mid-trunk) endopods and exopods subequal and several long and small setae. Segment 4 with in length, but endopod slightly shorter than exopod obliquely expanded medial margin even bearing two on anterior and posterior trunk. All setae faintly rows of 8-9 short and tong setae; proximal corner feathered (cf. Fig. IE).

3 (endite) equipped with setulose spines (2 of which Anal segment 1.3 times longer than wide (Fig

and 1 5 caudal 69% long and slender rather short). Segment as 11C); length of rami about of length long as segment 4, expanded, with a cluster of anal somite, bearing a single, fine seta on midmedial

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and 5-6 fine, and Dissolved concentration margins apical subapical setae, oxygen (DO) dropped ra-

respectively. 1.06 pidly from mg/l at the surface to 0.1 mg/l at 2

and 0.5 m depth stayed below mg/l to 20 m where

Remarks. - The midmedial of the fron- it increased processes steadily to 3.5 mg/l at 46 m. The pH at

tal filaments show several fine the sutures, indicating surface was 7.0; it spiked from 6.9 at 2.6 m to

articles or since the 7.4 at 3.2 and separate segments. However, m was followed by a general increase

of minnsi is based on a to 7.55 46 DO description Speleonectes at m depth. The low layer corre-

single we cannot exclude the specimen possibility sponded to a murky, hydrogen sulfide zone. Most of this an caused being artefact, e.g., by preserva- , including remipedes, were observed and tion. collected from the clear, marine saline water be-

minnsi is similar low the sulfide 25 Speleonectes morphologically hydrogen zone at to 40 m depth. S. and S. S. to epilimnius gironensis. However, Remipedes were observed swimming in the water minnsi can be from these column and clearly distinguished spe- were collected either by hand in indi-

cies different and with by shapes setation patterns of vidual vials or a suction bottle.

the maxillae Other maxillules, and maxillipeds; a larger animals present in the terminal breakdown

numberof trunk both in S. chamber where segments (than epilimnius the remipedes were collected in- and S. frontal filaments with mid- cluded copepods (three of undescribed gironensis)-, long genera epac-

medial in S. processes (short epilimnius): dorsal teriscids and many other unknown species; A. Foss-

antennular with flagella 12 segments (10-11 in s. hagen, pers. commun.); the halocyprid ostracodes and 10 in S. epilimnius, gironensis); caudal rami Danielopolina exuma and Deeveya medix; the lepto- being shorter than anal segment (distinctly longer stracan Speonebalia sp.; the peracarid Thetispe- in both S. epilimnius and S. lecaris remix; the amphipods Bahadzia gironensis). sp. and

Socarnopsis catacumba; a thermosbaenacean, prob- Tulumella ably sp.; a mysid, probably Stygiomysis of and the Ecological profile the type locality sp.; polynoid polychaete Pelagomacelli-

All of these cephala iliffei. organisms are cave-

The Basil Minns Blue Hole is located southeast of adapted, stygobiontic taxa characteristic of hydro-

Georgetown on Great Exuma Island in the central logically isolated anchihaline caves. Copepod tax-

Bahamas (see Fig. 14). A karst window consisting onomist Audun Fosshagen of the University of of two sinkhole the Basil Minns entrances at bottom of a salt- Bergen considered Blue Hole one of water lake provides access to the submerged cave. the most interesting copepod caves in the Baha-

A 10-12 m wide passage extends to the northeast mas. from the lake for several hundred meters at 50 m depth, before opening into a 50 m wide collapse- floored This circular room. room contains a large Some remarks on the biogeography and breakdown mound in the center with an air dome evolutionary history of Remipedia above, but no apparent opening to the outside. Sev- eral large root masses, penetrating through cracks The three species described herein present another in the down A ceiling, hang into the pool. Hydrolab remarkable example of sympatry for remipedes,

DataSonde 3 water quality analyzer carried by a especially since these crustaceans are exclusively diver used the column in was to profile water the known from marine subterranean habitats. Ground- terminal breakdown chamber 13). (Fig. Salinity water environments are typically characterized by increased from 9.8 at the surface, to 32.7 at a limitation of in ppt ppt nutrients and, part as a conse-

14 m and then to 34.8 at depth more gradually ppt quence, low abundances of stygobiont organisms.

46 m increased depth. Temperature from 26.7 de- The fact that are remipedes hermaphrodites may

C at the surface, to a maximum of 29.1 de- also grees point towards an adaptation to small popula- C between 14 and 19 m and then tion grees dropped m sizes. Interestingly, the sympatry of the three several 23.8 C 46 steps to degrees at m depth. new species described herein is not exceptional for

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13. Water column of and in terminal of Fig. profile salinity, temperature, dissolved oxygen pH versus depth the breakdown room

Basil Minns Blue Hole, Great Exuma Island, Bahamas. Data were collected with a Hydrolab DataSonde 3 water quality analyzer carried by a diver.

Remipedia. There are several other instances of two occurrences of sympatry could be result of dispersal

three invasions of available or sympatric species (Tab. 1). However, sym- and, subsequently, multiple patric remipedes are known only from the Baha- anchihaline caves. mas and nearby West Indian islands (Fig. 14). The Sympatric remipedes are likely to be subjected high abundance of remipedes and their taxonomic to strong competition, which could lead either to diversity at and below the family level in this re- niche differentiation or competitive exclusion. For gion forms a sharp contrast to the disjunct occur- example, the new species S. tanumekes (12 speci- rences of a few remote taxa. The Remipedia in the mens collected) seems to be more abundant than S. northwestern region of the West Indies andlthe parabenjamini (2 specimens) and S. minnsi (1 speci-

12 2 this Bahamas comprise species (6 genera and fami- men). However, whether is a sampling bias, a lies). Of the remaining three taxa, S. tulumensis seasonal fluctuation or indeed related to popula-

Yager, 1987b, from the Yucatan Peninsula could tion dynamics of the three remipedes in Basil Minns

of the cluster. Blue be be regarded as a peripheral isolate main Hole needs to confirmed by additional

By contrast, S. ondinae (Garcia-Valdecasas, 1984) collections. from the Canary Islands and L. exleyi Yager & Since their discovery, a plethora of opinions has

Humphreys, 1996, from western Australia are ex- been offered regarding the evolutionary history and

of Inver- treme disjunct occurrences (Fig. 15). phylogenetic status Remipedia (see any

The high availability of suitable habitats (anchi- tebrate Zoology textbook). Yet, we can be fairly haline caves) within a relatively small region (Ba- certain that Remipedia are an ancient group of crus-

share well hamas, West Indies) may have had, and still have, taceans. Remipedes several defined fea- an important impact on the unique geographic dis- tures with the Carboniferous fossil Tesnusocaris tribution of Remipedia. In this light, the frequent goldichi Brooks, 1955, from the Tesnus Forma-

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Fig. 14. Geographic distribution of Remipedia on the Bahamas and in the northwestern West Indies, including 12 of a total of 15

species known to science. Rectangular labels indicate occurrences of two or three sympatric species; circles show records of non-

= sympatric occurrences. Abbreviated taxa: Ch Cryptocorynetes haptodiscus; Gf = Godzilliognomusfrondosus; Gr = Godzillius

robustus ; Le = Lasionectes entrichoma; Pa = Pleomothra apletocheles; Sb = Speleonecles benjamini; Se = S. epilimnius; Sg = S.

= SI = S. S. Sm = S. minnsi St = S. tanumekes gironensis; lucayensis; Sp parabenjamini n. sp.; n. sp.; n. sp.

tion (Upper Mississippian/Lower Pennsylvanian) 1985). These theories are particularly interesting

since in in Texas (see Emerson & Schram, 1991, for a de- all remipedes occur regions that are com-

tailed redescription). The most distinctive shared paratively young in geological time. For example,

characters comprise a homonomously segmented the Canary Islands were probably formed during

trunk equipped with paddle-shaped, biramous swim- the Late Tertiary, and anchihaline cave systems

three ming appendages, and the modification of post- resulting from volcanic activity, such as lava tubes,

have oral cephalic appendages as subequal prehensile must been colonized subsequently. Yet, we

limbs. However, does the global, circum-tropical have to be careful not to jump to conclusions. Too

distribution of remipedes support an ancient origin little is still known about the biology ofRemipedia,

for this group? Such disjunct occurrences of taxa and at present, each additional discovery seems to

are often interpreted as relicts of an ancient marine add a piece to a more complete mosaic. The cur-

distribution consists of of distribution, as postulated for various crustacean rent a prominent cluster

stygobionts, e.g., hadziid and bogidiellid amphi- taxa in the northern Caribbeanregion including the

pods (Stock, 1981; Holsinger, 1986; Koenemann Bahamas. Whether this cluster is an ancient center

& Holsinger, 1999). Alternative theories favor a of origin and the disjunct taxa are isolated relicts

At this it is deep-sea origin for some stygobiontic crustaceans remains to be seen. point, equally con-

(Wilson, 1999; Manning et al., 1986; Hart et al., ceivable that we are observing a distribution pat-

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Bahama (AL); Bahama Peninsula Island Province (AL); Island Cave Island Lanzarote Grand Range Bahama Cenote Airport Matanzas (XL), (AL), Bahama Salvador (TL), Cape Grand (TL),(XL), (TL),(XL), Najaron locality. System Grand (TL),(XL), San Atlantida Cave (XL),(TL), (TL),(XL), species Cave (TL),(XL), C-28 (TL),(XL), la additional Hill CavernCavern Carboneros de Cave Islands Cave Cave Cenote Blue TunelXunel = single Freetown los AL Asgard Old Caicos Major’s LucayanLucayan de with Old Australia: Carwash RooRoo Cueva Islands: locality, Indies:Indies; and Localities Bahamas: Bahamas:Bahamas: West Western Bahamas: Bahamas: Canary Mexico: Quintana type Island none none West XurksTurks nonenone none none none Cuba:

=

TL Island Island Island 14). Islands Islands Island Island Fig. Exuma Exuma Exuma Caicos Caicos also Bahama (see Island Island and and Great Great Great of Island Island; Island; Grand (TL),(XL), (TL),(XL), (TL),(XL), Bahama Grand TurksXurks TurksXurks species Abaco Abaco Bahama Abaco Bahama species (TL),(XL), (AL), (AL), Hole Hole Hole Grand (AL), Grand Blue Blue Blue sympatric (TL),(XL), (TL),(XL), (AL), Cave Pond Pond sympatric Cave Cave (AL), Cave (AL), Minns Minns Minns of Cottage Cottage with Dan’s Dan’sDan’s Cave Dan’s Cave Sagittarius Basil Basil Basil occurrences Indies: Indies: Localities Bahamas: Bahamas: Sagittarius Bahamas: Sagittarius Bahamas: West West Bahamas: Bahamas: Bahamas: none nonenone nonenone nonenone none none several 1999 with -— 1986 1984) 1996 1987a 1989 Schram, 1986 Carpenter, Remipedia, 19891989 1987a 1981 19941994 1987b & al., sp. & Yager, & al., Humphreys, of Yager,Yager, n. Yager,Yager, et sp. Yager, Yager & Yager Yager, Yager,Yager, (Garcia-Valdecasas, Yager, Schram n. sp. localities haptodiscus frondosus n. Yager apletocheles benjamini entrichoma tanumekes parabenjamini minnsi epilimnius lucayensis gironensis ondinae tulumensis Recorded robustus exleyi 1. Godzilliognomus Table Species Cryptocorynetes PleomothraPleomolhra SpeleonectesSpeleonecles Lasionectes Godzillius SpeleonectesSpeleonecles SpeleonectesSpeleonecles SpeleonectesSpeleonecles Lasionectes SpeleonectesSpeleonecles SpeleonectesSpeleonecles SpeleonectesSpeleonecles SpeleonectesSpeleonecles SpeleonectesSpeleonecles

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equator.

the

of south

and north

30° latitudes

and equator,

the indicate

lines Horizontal

species. individual represent

circle Filled Remipedia.

of distribution

Global

15.

Fig.

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tern subjected to biased sampling in less accessible Bogidiella arista, new species, from Turkey. J. Crust. Biol.

the 18 (2): 383-404. diving regions. We cannot even exclude possi- Koenomann S, Holsingcr JR. 1999. Phylogenetic analysis of that the localities the bility two most disjunct on the amphipod crustacean family Bogidiellidae, s. lat., and Canary Islands and in western Australia are the result revision of taxa above the species level. Crustaceana 72 (8): of kind of Koenemann some passive dispersal (see 781-816. et al. 1998). Manning RB, Mart JW jr, Iliffe TM. 1986. Mesozoic relicts

in marinecaves of Bermuda. Stygologia 2 (1/2): 156-166,

Schram FR. 1974. Late Paleozoic Peracarida ofNorth America.

Fieldiana: Geology 33: 95-124. Acknowledgments Schram FR. 1986. Crustacea. New York: Oxford University

Press.

We are grateful to Jill Yager for providing us with specimens Schram FR, Yager J, Emerson MJ. 1986. Remipedia. Part I. (in connectionwith another project) that facilitated the identifi- Systematics. Mem. San Diego Soc. Nat. Hist. 15: 1-60. cation of these new species. We also wish to thank Jan van Stock J.H. 1981. The and zoogeography ofthe fam- Arkel for digital processing of all line drawings, Bruce Felgen- ily of Bogidiellidac (Crustacea, Amphipoda), with emphasis hauer for assistance with the processing ofspecimens for SEM, on the West Indian taxa. Bijdr. Dierk. 51: 345-374. Texas A&M University graduatestudents Rebecca Belcher, Darcy Vonk R, Schram FR. 2003. I ngolfiel 1 idea (Crustacea, Gibbons, Lara llinderstein and Colin Kliewer for sorting and Malacostraca, Amphipoda): a phylogenetic and biogeo- photographing collected material, and cave diver Brian Kakuk graphic analysis. Contrib. Zool. 72 (I): 39-72. (Caribbean Marine Research Center) for assistance with the Wilson GDF. 1999. Some of the deep-sea fauna is ancient. collection This research ofspecimens. was supported by a grant Crustaceana 72 (8): 1019-1030. from the Schure-Beijerinck-Popping fund of the Royal Dutch of from Yager J. 1981. A new class Crustacea a marine cave in Academy of Sciences (KNAW, SBP/JK/2003-25) to J. van der the Bahamas. J. Crustacean Biol. I: 328-333. Ham, and the National Oceanic and Administration Atmospheric J. 1987a. Yager Cryptocorynetes haptodiscus, new genus, new (NOAA) Caribbean Marine Research Center at Lee Stocking species, and Speleonectes benjamini, new species, of

Island, Bahamas, to T. Iliffe. described here were Specimens crustaceans remipede from anchialine caves in the Bahamas, collected under marine resource collecting permits issued by with remarks on distribution and ecology. Proc. Biol. Soc. the Bahamas ofFisheries to T. Iliffe and B. Kakuk. Department Wash. 100: 302-320.

This DIVERSITAS-IBOY paper is a contribution to project J. 1987b. Yager Speleonectes tulumensis, n. sp. (Crustacea, “Exploration and Conservation ofAnchialine Faunas”. Remipedia) from two anchialine cenotes of the Yucatan Pe-

ninsula, Mexico. Stygologia 3: 160-166.

Yager J. 1989. Pleomothra apletocheles and Godzilliognomus

two new and of crusta- References frondosus, genera species remipede

ceans (Godzilliidae) from anchialine caves ofthe Bahamas.

Bull. Mar. Sci. 44 (3): 1195-1206. Birshtein YA. I960. Podklass Cephalocarida: Chlenistongie, Yager J. 1994. Speleonectes_gironensis, new species (Remi- Trilobitoobraznie, Rakoobraznie. In: Orlov YA (ed.). Osnovy pedia: Speleonectidae) from anchialine caves in Cuba, with Paleonlologii. Akadetnii Nauk, Moskva: 421-422. remarks on biogeography and ecology. J. Crust. Biol. 14: Brooks 1IK. 1955. A crustacean from the Tesnus formation of 752-762. Texas. J. Paleont. 29: 852-856. Yager J, Carpenter JH. 1999. Speleonectes epilimnius new Emerson MJ, Schram FR. 1991. Remipedia, Parf 2, species (Remipedia, Speleonectidae) from surface waters of Paleontology. Proc. San Diego Soc. Nat. Hist. 7: 1-52. an anchialine cave on San Salvador Island, Bahamas. Garcia-Valdecasas A. 1984. Morlockiidae new family of Crustaceana 72: 965-977. Rcmipcdia (Crustacea). Eos 60: 329-333. WF. 1996. Lasionectes Yager J, Humphreys exleyi, sp. nov., Hart CW jr, Manning RB, Iliffe, TM. 1985. The fauna ofAt- the first remipede crustacean recorded from Australia and lantic marine caves: Evidence ofdispersal by sea floor spread- to the Indian Ocean, with a key the world species. Invertebr. ing while maintaining ties to deep waters. Proc. Biol. Soc. Taxon. 10: 171-187. Wash. 98 (1): 288-292. Schram FR. 1986. Lasionectes Yager J, entrichoma, n. gen., ofNorth Ameri- Ilolsingcr JR. 1986. Zoogeographic patterns from anchialine in the n. sp. (Crustacea, Remipedia) caves crustaceans. can subterranean amphipod In: Gore RH, Heck Turks and Caicos, B.W.I. Proc. Biol. Soc. Wash. 99 (1): 65- KL, eds. Crustacean Biogeography (in Crustacean Issues). 70. Rotterdam: Balkema, 85-106.

Koenemann S, Vonk R, Schram FR. 1998. Cladistic analysis

of 37 MediterraneanBogidiellidae (Amphipoda), including Received: 8 May 2003

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