Bijdragen tot de Dierkunde, 52 (1): 3-12 1982

Amsterdam Expeditions to the West Indian Islands, Report 16.

A representative of the mainly abyssal family Pardaliscidae

(Crustacea, ) in cave waters of the Caicos Islands

by

Jan H. Stock & Jan J. Vermeulen

Institute of Taxonomic Zoology, University of Amsterdam,

P.O. Box 20125, 1000 HC Amsterdam, The Netherlands

Abstract the km from sea (the nearest sea coast is some II/2

A new and species of Amphipoda is described from the cave entrance). The accessible part of the water

cave waters on Providenciales (Caicos Islands) as Spelaeoni- hole is in semidarkness.

cippeprovo. Another species belonging to the same new genus this have been ob- The Amphipoda from cave is known from a cave on Lanzarote (Canary Islands). The served in genus belongs to the Pardaliscidae, a predominantly bathyal/ free-swimming ("pelagic") the water. abyssal family. A possible evolutionary scenario for both with and They are unpigmented, a pink intestine, species is discussed. fauna devoid of eye pigment. The accompanying

is of several of relatively poor, consisting species Résumé Gastropoda. Dr. H. E. Coomans (Institute of décrit des On un genre et espèce nouveaux d’Amphipodes Taxonomic of eaux d’une grotte de Providenciales (Iles Caicos), sous le Zoology, University Amsterdam) nom de Spelaeonicippe provo. Une autre espèce appartenant has been kind enough to examine these mollusks est d’une de Lanzarote au même genre connue grotte (Iles and he informs us (in litt.) that seven species are Le famille Canaries). genre appartient aux Pardaliscidae, con- Dead des represented. shells of Echininus nodulosus nue surtout étages bathyal et abyssal. On présente un scénario évolutif deux plausible pour ces espèces. (Pfeiffer, 1839), Truncatella caribaeensis (Reeve,

and 1842), Melampus (Microtralia) sp. (possibly

M. minusculus) were present, together with live INTRODUCTION and dead of Of E. specimens Lyrodes sp. these,

nodulosus and T. caribaeensis are living in the of the fauna of the During a survey groundwater marine fringe of the supralittoral of the coast; Bahamas and the islands in the West adjoining lives in and Lyrodes pools springs, near the sea; of taken in Indies, a sample Amphipoda was a is inhabitant Melampus an of brackish waters. All so-called "water hole" on an Providenciales, and all certain are superficial species, indicate a island in the western of the Caicos to part group, relation to the marine environment. In addition, the north of Haiti. The "water hole" in question the terrestrial cave yielded a number of snails, all is in an oblique fissure in the calcareous rocks, no doubt washed in from the surface, viz., Bo- direct with the table. This contact underlying water and thriopupa sp., Gastrocopta sp., members of "water hole" be classified particular may as a the No other family Chrondropomidae. macro- "marginal cave" Riedl & Ozretic, 1969) (sensu or microfaunal elements were observed in this or anchihaline habitat (sensu Holthuis, 1973), particular water hole, but a nearby water hole, since the high chlorinity of its waters (ll.l°/oo) presumably part of the same subterranean water points to extensive subterranean connections with harboured Ma- system, Typhlatya sp. (Decapoda

and Sesarma crura) sp. (Decapoda Brachyura).

Typhlatya is a wide-spread stygobiont element in has been in *) Report no. 15 published Bijdr. Dierk., 51 (2): the West Indies, of marine 375-387 (1981). probably origin (cf. 4 J. H. STOCK & J. J. VERMEULEN - CAVE PARDALISCIDAE

the Stock, 1981), and Sesarma is a marine and mixo- related to Nicippe, is still more close to cave

haline In faunal from and is therefore genus. summary, most elements Providenciales,

in the holes the of Providen- reckoned the Certain dif- water near airport to genus Spelaeonicippe. ciales have distinct marine affinities. ferences in its appendages, in addition to the

The called ocean-wide between the caves of Amphipoda belong to a new genus, geographical gap in related to the Ni- Providenciales and Lanzarote, justify our Spelaeonicippe, closely genus

member of that the Canarian and West Indian cippe Bruzelius, 1859, a the family opinion popu-

lations treated Pardaliscidae. This family is well-characterized by are to be as separate species.

the distinct molar mandible, lacking a process;

small inner and by the very outer plates of the THE HABITAT OF SPELAEONICIPPE maxillipeds; by the short coxal plates; and by the

elongate, lanceolate rami of the third uropods Both localities from which Spelaeonicippe is

Lanzarote and in at (Barnard, 1969). known, Providenciales, agree

least three respects:

(1) The caves are of relatively young geological

BATHYMETRICAL DISTRIBUTION OF THE del Lanzarote is age. The Jameos Agua on part PARDALISCIDAE lava of a tube formed 3000-5000 years ago (Wil-

The Pardaliscidae are predominantly bathyal/ kins & Parzefall, 1974). The extensive karst ero-

which abyssal/hadal, pelagic Amphipoda (Bousfield, sion, during caves were formed by ground-

and Birstein Providenciales 1978), & Vinogradov (1962: 249) water solution, on developed during

them "one the families the consider as of basic abyssal low sea-level stages (at -100 to -120 m) of

of The which the Pleistocene thus sometime be- gammarids". genus Nicippe to (Mather, 1975),

and present cave amphipods are most closely related, tween 10,000 1,000,000 years ago.

is in and Providen- not restricted to the deep sea. Most of its records (2) The cave waters Lanzarote

the lower ciales The Lanzarote are from part of the continental shelf are salty. lava tube runs from

and the of the continental the volcano la Corona to the coast where it is upper part slope (110-

600 cf. flooded The blue holes and other m; Metzger, 1875; Sars, 1893; Chevreux, by seawater.

karst extend 1900; Enequist, 1949; Gurjanova, 1951; Ledoyer, features on Providenciales often

in- with down below sea level and are 1973, 1977), exceptional deeper ones, present nowadays

to 1393 m (cf. Stephensen, 1931). According to undated by seawater, due to sea-level rises at the

certain authors Karaman end of the ice and — to lesser (Barnard, 1959; & ages, perhaps a

this extent — due to the subsidence Schiecke, 1973) genus penetrates also in general regional

waters of the of the of the Bahamas upper part shelf, though platform.

In in fluctuates always in the sublittoral (from 35 m down). Lanzarote, the salinity the cave

Bousfield states that the around 37°/ with a Although (1978) 00 (corresponding chlorinity

Ene- which is members of the Pardaliscidae are carnivorous, of some 20.4°/oo)> slightly (0-l°/oo)

that of the quist (1949) supposes that at least Nicippe might inferior to adjoining ocean (Wilkins

table The be a detritus feeder, and Wilkins & Parzefall & Parzefall, 1974, I). water hole on

diatoms (1974) consider the primary food source Providenciales has a chlorinity that is distinctly

of Spelaeonicippe buchi (as Nicippe). lower than that of the surrounding ocean, viz.

the which is still for Only one member of family Pardaliscidae about ll.l%o, but very salty

has been found outside the in waters: inland fresh water mixes open sea cave groundwater. Apparently,

described with of marine not Andres (1975) a new species, under salty groundwater origin, a

the name of buchi from a lava tunnel uncommon situation in the Bahamian Nicippe , archipelago.

well-known has in (the Jameos del Agua) on the island (3) Spelaeonicippe been found both the of Lanzarote in the Canarian archipelago. As will West Indian and the Canarian locality in con- be shown in the taxonomic part of the present ditions where light could penetrate and not in

the As Par- paper, Canarian cave species, though closely total darkness. to Lanzarote, Wilkins & BIJDRAGEN TOT DE DIERKUNDE, 52 (1) - 1982 5

zefall that this is leaves the from time (1974) believe not accidental, gravel on the cave bottom to

since the feed swim amphipods are supposed to on time, to around freely, and to disappear

in the in the after while. The microscopic algae developing cave waters. again substrate a new

For West Indian less certain of found in the West Indies the locality, we are species Spelaeonicippe

shows similar but Gamma- about this point, because the totally dark parts a behaviour, hadziid

the "water hole" ridae have been of were not explored (so no never observed swimming around

information is available about absence in the water. The recent presence or freely relatively origin

of in and marine Spelaeonicippe those parts), no diatom (from ancestors) of the pardaliscid cave

with growth was observed in the entrance area of Amphipoda seems in good agreement (1)

the cave. the existence of close relatives in continental shelf

and and their in slope waters, (2) occurrence

formed in geologically young caves, pleistocene/

ORIGIN OF THE FAUNA IN THE MARGINAL CAVES holocene times.

OF LANZAROTE AND PROVIDENCIALES At if blind hadziid any rate, Spelaeonicippe or

It accidental del to discovered can hardly be that in the Jameos Gammaridae are be in the open sea,

will be meio- Agua on Lanzarote, in addition to Spelaeonicippe we suppose they shallow-water,

faunal buchi (a species which is undoubtedly related to elements in coarse substrates. It is from

the tumida such and not from the sublittoral/bathyal species Nicippe elements, abyssal ones, that

Bruzelius, 1859), two other organisms with deep- cave forms presumably evolved. In agreement with

water affinities occur: Munnidopsis polymorpha this view is the fact that the temperatures in the

Koelbel, 1892 (Anomura, Galatheidae) and Mu- Lanzarote cave (18.0-18.5° C) and in the Provi-

similar cellicephala jameensis Hartmann-Schroder, 1974 denciales cave (23.8° C) are closely to

(Polychaeta, Polynoidae). shallow-water temperatures in the neighbouring

We do not share Wilkins & Parzefall's opinion ocean, and not at all to bathyal/abyssal tem-

(1974: 431) that the animals in the Jameos del peratures of 10-4° C in the neighbouring ocean.

Agua might be wide-spread in the deeper waters

of the neighbouring Atlantic. Of course, the ab-

of and of of the TAXONOMIC PART sence eyes body pigmentation

cavernicolous forms is analogous to similar reduc- Spelaeonicippe n. gen. found in animals. On the tions many deep-sea

other hand, the pardaliscid cave Amphipoda of

Lanzarote differ in number Pardaliscidae a of morphological Family (Amphipoda). Resembling

the in the of the characters from the nearest marine relative. Fur- genus Nicippe general shape

of known and the thermore, a second species amphipod telson, of the mouthparts, of large, oval

from the hadziid in the 1 and Differs from same cave, a (see Andres, 1978) propodus gnathopods 2.

has relatives all in It in the characters enumerated in table I. no at deeper waters. must Nicippe

S. be stressed that both species of cave amphipods Type-species: provo n. sp.

from Lanzarote have their closest relatives in fresh Other species: S. buchi (Andres, 1975) (as Ni- and brackish groundwaters of the West Indies. cippe).

This makes it more likely that they descend from Distribution: Cave waters on Providenciales (Cai-

marine ancestors that during some phase of the cos Islands) and Lanzarote (Canary Islands).

evolutionary scenario became adapted to a life in Derivatio nominis: From CT7TT]XOUOV (Greek, cave)

and in the interstitia of coarse substrates (e.g. gravel). Nicippe (a generic name pardaliscid Am-

The this lie based and adaptation to kind of substrate might phipoda, on Greek mythology named

farther in the past for the exclusively benthic had- after the daughter of Pelops). Gender feminine.

ziid Gammaridae, than for the predominantly Remarks: We consider in particular the non-trans-

pelagic Pardaliscidae. Wilkins & Parzefall (1974: formed first male antenna as a diagnostic character

have that buchi the 430) observed Spelaeonicippe of new genus. - 6 J. H. STOCK & J. J. VERMEULEN CAVE PARDALISCIDAE

TABLE I

Salient between the and n. differences genera Nicippe Bruzelius, 1859, Spelaeonicippe gen.

Nicippe Spelaeonicippe

1. Lateral head lobes large, acutely produced Inconspicuous, rounded

2. Al > A2 **) Al « A2

of of Al transformed Normal 3. Segments accessory flagellum $

Al with without 4. Flagellum segment 1 of $ elongated, extra Short, special setation

setation

of Md dense 5. Second and third segment palp with With sparse setation

setation

of distal dilated; and medial ns 6. Distal palp segment Mxl clavate (with strong Not armature on terminal marg :

dilatation); armature restricted to terminal margin

7. Outer lobe of Mxl less wide than 2nd palp segment Wider than 2nd palp segment

8. Outer lobe of Mxl reaches to 75% of the length of To 33%

the palp

claw of slender 9. Segments 2, 3 and Mxp palp Plump 10. Outer lobe of Mxp setiferous Spiniferous

11. Outer lobe of Mxp reaches to the end of the inner Does not reach the end

margin of palp segment 1

12. Merus of P5 setiferous Spiniferous

13. Urosomite 1 with dorsal teeth Smooth

illustrates male tumida Bruzelius in which Al is shorter than but Karaman **) Sars, 1893, pi. 145, a of Nicippe slightly A2,

& Schiecke (1973: 166) consider this an exceptional case.

armed with of Spelaeonicippe provo n. sp. ventrally a row spinules, dorsally

No with groups of short setae. calceoli. Flagel-

Material. — One 9 (holotype), one $ (allotype), eight lum 16- to 17-segmented. Amsterdam Indian paratypes. Expeditions to the West Islands, Labrum sta. 79-151. Providenciales (Caicos Islands): nameless "water (fig. Id) bell-shaped. hole" N.W. of the airfield (estimated position 21°46'50"N Masticatory part of mandible asymmetrical semi-dark; 1 72°16'30"W); stones, silt; water depth over m; (figs, le, If): left mandible with, right mandible water temperature 23.8° C; chlorinity 11,108 mg/1; 16 No- vember without lacinia mobilis. Pars molaris reduced; 1979. Zoologisch Museum Amsterdam coll. no. ZMA 107.495. Amph. cutting edge long, almost toothless; a row of 8-11

spines is present between the cutting edge and the of Description. — Body length adult speci- implantation of the palp. Palp slender; segment 1

ventral mens 3-4 mm; body unpigmented (intestine pink); unarmed; segment 2 with 7 or 8 setae;

and 3 with 1 2 with no eye pigment. Urosomites 2 or third segment narrower than the second, 2 minute dorsal setules; dorsum otherwise smooth. terminal and 2 subterminal setae only.

Coxal plates low. Lateral head lobes inconspicuous, Labium (fig. lg) deeply cleft, without inner

round- rounded; no rostrum. Epimeral plates with lobes. ed posterior corners (fig. la). First maxilla: left (fig. 2b) and right (fig. 2a)

First antenna (fig. lb) reaching to the end of appendages almost symmetrical. Inner lobe trian- the pedunculus of the second antenna. Pedunculus gular, with 1 apical seta; outer lobe squarish,

and segments heavy short. Accessory flagellum armed with 7 strong distal spines. Palp 2-seg-

3-segmented, almost as long as second pedunculus mented; distal segment ovate, not dilated distally;

aesthe- segment. Flagellum 17- to 18-segmented; medial and distal margins with some 13 short tasks 4 on segment 1 (6 long aesthetasks), 3 and spines.

and Second maxilla of (each 2 long aesthetasks), 6, 7, 8, 9 10 (each (fig. 2c) consisting two very

1 short aesthetask). slender, digitiform, lobes. Outer lobe with 3 distal

medial Second antenna (fig. lc): gland cone short, setae; inner lobe with 8 or 9 setae. conical. Pedunculus segments 4 and 5 elongate; Maxilliped (fig. lh): inner lobe small, ovate, BIJDRAGEN TOT DE DIERKUNDE, 52 (l) - 1982 7

1. Fig. Spelaeonicippe provo n. gen., n. sp., ￿ paratype: a, entire from the right (real size abt. 4 mm); b, first

second labrum mandible left mandible antenna (scale A); c, antenna (A); d, (B); e, right (B); f, (B); g, labium (B); h, maxilliped (C). Scales see fig. 4. 8 H. STOCK - J. & J. J. VERMEULEN CAVE PARDALISCIDAE

2. Fig. Spelaeonicippe n. provo n. a, right first maxilla (scale B); b, left first gen., sp., ￿ paratype: maxilla (B); c, second maxilla (B); d, first pleopod (A); second (A); f, third e, pleopod pleopod (A). Scales see fig. 4. BIJDRAGEN TOT DE DIERKUNDE, 52 (l) - 1982 9

with distal Outer the much wider than that of 2 setae. lobe very short, not long as sixth; basis

the distal end of the first the reaching segment of sixth, but likewise with a strongly overhanging

the inner margin of the palp, armed with 3 distal lobe. Both sixth and seventh pereiopod armed

and several short medial spines setae. Palp seg- with spines only.

ments short and 1 to similar in robust. Pleopods 3 (figs. 2d, e, f)

First gnathopod (fig. 3a): Coxal plate with morphology; pleopod 3 slightly shorter than the

a rounded anterior projection. Carpus triangular. others. All pleopods are biramous and the rami

Propodus large, ovate. No palmar angle marked; are untransformed in both sexes. Two retinacula

of each retinacula the entire posterior edge the propodus forms on pleopod; the are exceptionally

the palmar margin. This margin (fig. 3b) is long; the extreme distal part bears rake-like teeth

densely packed with some 40 shorter setae (the (fig. 2e, detail).

of which bears proximal margin many cilia, the Uropods 1 (fig. 4d) and 2 (fig. 4e) not much

distal margin fewer cilia) and 5 longer setae (the different in length; both with short exopodite

of which bears proximal margin few cilia, the (0.6-0.7 times as long as the endopodite).

distal Third both rami margin many). Claw long, curved; one uropod (fig. 4f) equiramous,

very short setule on its outer margin. 1-segmented, rather wide, lanceolate to foliaceous.

Second gnathopod (fig. 3c) slightly larger than Outer margin of exopodite armed with 3 or 4

the but otherwise similar. Outer of with limited first, very spines. margin endopodite a

and of Oostegites on gnathopod 2 pereiopods 3 to number of plumose setae. Inner margin exo-

5; linear, armed with long setae. Coxal gills with and endopodite with numerous long, plumose

an indistinct basal stalk, ovate to sausage-shaped, setae.

decreasing in size from anterior to posterior, Telson (fig. 4g) deeply cleft; lateral margin

half with present on gnathopod 2 and pereiopods 3 to 7. of each slender 2 spines; distal part with

Third with and pereiopod (fig. 3d) modified car- 2 plumose (sensory) setae 2 spines.

this and No pus: segment is slightly swollen armed secondary sexual differences have been

with numerous long, plumose setae on its poste- observed in the appendages of males and females. rior margin. The propodus on the contrary is

for Distinction. —, is similar practically unarmed, except some short plumose S. provo n. sp. very elements the distal end its S. buchi from Lanzarote. The on of posterior margin. to (Andres, 1975)

The claw is extremely elongate. following differences exist between the two

Fourth pereiopod (fig. 4a) with trapezoidal species:

coxal plate, without posterior excavation. Carpus (1) Body length 3-4 mm in S. provo (versus

armed with 7-10 in S. buchi 1 of antenna not swollen and a limited number of mm ?'); (2) segment short with 1 than wide in S. plumose setae only. Propodus numer- only slightly longer provo

short in S. buchi 2 of Al ous plumose setae on the entire posterior (elongate ); (3) segment

Claw in the 4 the of in S. margin. as third pereiopod. about /s of length segment 1 provo

in Fifth pereiopod (fig. 3e) slightly longer than (about 1/2 the length S. buchi); (4) molar the fourth. in of mandible 8 11 in Basis almost linear shape; postero- process replaced by to spines distal into S. to 3 in S. buchi 2nd mandible comer produced a strongly overhanging provo (2 f); (5)

with ventral in S. lobe. Merus armed with a few short spines only. palp segment 7 to 8 setae provo

swollen but in S. buchi 2nd of maxilla Carpus not armed with a row of long (1 I; (6) palp segment

least in in setae and some spines along its anterior margin. 1 with at 10 spines S. provo (5 S.

i Anterior of with similar buchi) ost:er or of basis of arma- — (7) margin gnatho- margin propodus it P ture. pods 1 and 2 with several long setae in S. provo

of Sixth pereiopod (fig. 4b) much longer than the (unarmed in S. buchi); (8) propodus pereio- fifth. Basis less linear than in the 4 with in S. fifth pereiopod, pod setae on the posterior margin with and in S. a very strongly produced overhanging provo (spinules only buchi); (9) over-

Seventh lobe the basis of posterodistal lobe. pereiopod (fig. 4c) as hanging posterodistal on pereio- 10 J. H. STOCK & J. J. VERMEULEN - CAVE PARDALISCIDAE

of of Fig. 3. Spelaeonicippe provo n. gen., n. sp., ￿ paratype: a, first gnathopod (scale A); b, palmar margin propodus

first second third fifth Scales 4. gnathopod (D); c, gnathopod (A); d, pereiopod (A); e, pereiopod (A). see fig. BIJDRAGIiN TOT DE DIERKUNDE, 52 (1) - 1982 11

fourth b, sixth pereio- Fig. 4. Spelaeonicippe provo n. gen., n. sp., a-f, ￿ paratype; g, ￿ allotype: a, pereiopod (scale A);

seventh first second (A); f, third (A); telson (C). pod (A); c, pereiopod (A); d, uropod (A); e, uropod uropod g, 12 - J. H. STOCK & J. J. VERMEULEN CAVE PARDALISCIDAE

sto 7 in S. revised classification and pods very strongly developed provo BOUSFIELD, E. L., 1978. A phylo- of Trans, Soc. geny amphipod . roy. Canada, (moderately developed in S. buchi); (10) outer (4) 16: 343-389. of of 3 with margin endopodite uropod long des CHEVREUX, E., 1900. Amphipodes provenant campagnes plumose setae in S. provo (with spines only in de l'«Hirondelle». Result. Camp, scient. Prince Albert I, 16: i-iv, 1-196, pis. I-XVIII. S. buchi ); (11) telson with 2 lateral spines in Studies soft-bottom 1;NEQUIST, P., 1949. on the amphipods S. lateral in S. buchi) sen- provo (1 spine ); (12) of the Skagerak. Zool. Bidr. Uppsala, 28: 297-492, 1 the of the sory setae implanted near tip telson chart.

SSSR i lobes in S. the middle in GURJANOVA, E., 1951. Bokoplavy morej sopredel'nykh provo (slightly over vod (Amphipoda-). Opred. Faune SSSR S. buchi). (Akad. Nauk SSSR), 41: 1-1029.

HOLTHUIS, L. B., 1973. Caridean shrimps found in land-

locked saltwater pools at four Indo-West Pacific local- Derivatio nominis. — The specific name, ities (Sinai Peninsula, Funafuti Atoll, Maui and Hawaii provo, is the popular name for the type-locality, Islands), with the description of one new genus and

128: the island of Providenciales. four new species. Zool. Yerhand. Leiden, 1-48.

KARAMAN, G. S. & U. SCHIECKE, 1973. Some interesting

Amphipoda of the Pardaliscidae family (Amphipoda,

Gammaridea) of the Adriatic and Mediterranean Seas. ACKNOWLEDGEMENTS Memorie Mus. civ. Stor. nat. Verona, 20: 149-168.

The fieldwork which the is based has been on present paper LEDOYER, M., 1973. Amphipodes gammariens nouveaux ou

grants from the Netherlands Foundation for de la de Marseille. supported by peu connus region Tethys, 4 (4): the Advancement of Research The Tropical (WOTRO), 881-898.

The Treub The de de la faune Hague; Maatschappij, Utrecht; Beijerinck- 1977. Contribution a 1'etude l'ecologie

Popping Fonds, Amsterdam; and the Fonds Landbouwhoge- vagile profonde de la Mediterranee nord occidentale, I. school, Wageningen. Les Gammariens (Crustacea, Amphipoda). Boll. Mus.

Dr. Steven Weinberg is thanked for assistance during the civ. Stor. nat. Verona, 4: 321-421.

in the Bahamas. 1979 sampling program MATHER, J. D., 1975. Turks and Caicos Islands. In: R. W.

H.M. Embassy of The Netherlands in Kingston (Jamaica) FAIRBRIDGE ed., The encyclopedia of world regional logistic which provided support, is gratefully acknowledged. geology, X. Western hemisphere: 500-501 (Dowden, Hut- We are also indebted to Dr. H. E. Coomans (Institute of chinson & Ross, Stroudsburg, Pa., U.S.A.). Taxonomic of for iden- Zoology, University Amsterdam) METZGER, A., 1875. Zoologische Ergebnisse der Nordsee- the found with the tifying Gastropoda along Amphipoda at fahrt, 10. Crustaceen aus den Ordnungen Edriophthal-

the type-locality. mata und Podophthalmata. Jber. Comm. wiss. Unter-

such. dt. Meere, 2/3: 277-310, pi. VI.

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Received: 6 November 1981