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Freshwater shrimp genera Caridinaand Parisia (Decapoda: Caridea:Atyidae) of Madagascar, withdescriptions of four new speciesAnn R. Gurney aa Department of Zoology , British Museum (Natural History) ,Cromwell Road, London, SW7 5BD, EnglandPublished online: 17 Feb 2007.

To cite this article: Ann R. Gurney (1984) Freshwater shrimp genera Caridina and Parisia(Decapoda: Caridea: Atyidae) of Madagascar, with descriptions of four new species, Journal ofNatural History, 18:4, 567-590, DOI: 10.1080/00222938400770481

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JOURXAL OF NATURAL HISTORY, 1984, 18:567-590

F r e s h w a t e r sh r imp gene ra Caridina a n d Parisia

(Decapoda: Caridea: A ty idae ) o f M a d a g a s c a r ,

w i t h descr ip t ions o f four n e w species

ANN R. GURNEY

Department of Zoology, British Museum (Natural History), Cromwell Road, London SW7 5BD, England

Introduction Atyid shrimps were collected in the Grotte d'Andrafiab6 and Grotte

d'Antsatrobonko, Ankarana Massif, northern province of Diego Suarez, North Madagascar by Jane Wilson during the University of Southampton Expedition, 1981. The following 9 species were present in the collection including three new species of Caridina and one new species of Parisia.

Caridina Milne-Edwards, 1837 Registration Number:

C erurispinata sp. nov. C. isaloensis isaloensis Couti~re, 1899 C. nilotiea (P. Roux, 1833) C. norvestiea Holthuis, 1965 C. parvoeula sp. ~iov. C. unca sp. nov.

BM(NH) 1983.23-24 .40-41 .35-39 .33-34 .21-22 .25-29

Parisia Holthuis, 1956 P. dentata sp. nov. .30-32 P. maerophthalma Holthuis, 1956 .42-45 P. microphthalma (Fage, 1946) .46-48

A comprehensive review of the Atyidae of Madagascar by Holthuis (1965) lists 23 recorded species representing 5 genera. Holthuis drew attention to the genus Caridina, then represented in Madagascar by 16 species, as 'one of the notoriously difficult genera of the Decapoda Macrura'. Three new Caridina species are added in the present work and to facilitate species' determinations a re-evaluation is made of a valuable key-character previously given scant attention or completely ignored--an apical hook on the dactylus and propodus of pereiopods 1 and/or 2. Milne-Edwards (1837) made no mention of this feature in his original diagnosis of Caridina, but Bouvier (1925) redescribed the type species, C. typus, and clearly figured the hook on the dactylus and propodus of pereiopod 1.

Material from Madagascar representing all 21 species in the key has been examined with the exception ofC. troglophila Holthuis, 1965, which was unavailable from the Institut de Recherche Scientifique de Madagascar, Tananarive, at the time of the study. Since the presence or absence of an apical hook on the pereiopod could not be confirmed this species is keyed out before couplet 7 which refers to this character.

© 1984 The British Museum (Natural History)

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568 A . R . G u r n e y

Key to C a r i d i n a of Madagascar

1 Endopod of first pleopod of male with appendix interna 2 Endopod of first pleopod of male without appendix interna . . . . 6

2 Upper border of rostrum without teeth. Rostrum short, not reaching beyond second segment of antennular peduncle. Intermediate spine on posterior margin of telson longer than the lateral pair. Antennal spine placed on the lower orbital angle t y p u ,

- Upper border of rostrum with teeth. Intermediate spines on posterior margin of telson as long as or shorter than the lateral pair 3

3 Rostrum straight and short, not reaching beyond the second segment of antennular peduncle. Distal part of upper margin of rostrum unarmed, no subapieal teeth present. Antennal spine placed on lower orbital angle . j apon ica

- Rostrum with the distal part curved up, reaching distinctly beyond second segment of antennular peduncle. Distal part of upper margin of rostrum unarmed apart from one or more subapical teeth. Antennal spine placed below the very distinct lower orbital angle . . . 4

4 Posterior margin of telson ending in a median point. Preanal earina with a sharp posteriorly directed spine. Eggs numerous and small (0-2~)'4 x 0"3-0-6 mm) 5

- Posterior margin oftelson rounded. Preanal earina blunt without a spine. Eggs fewer and larger (0'5-0"6 x 0'7-1"0 mm) nilotica

5 Dorsal margin of rostrum with 13-23 spines in basal part, which are placed close together . longirostris

- Dorsal margin of rostrum with 5-13 spines in the basal part, which are placed widely apart . gracilirostris

6 Terminal margin of telson stepped, troglophila - Terminal margin of telson not stepped . 7 7 Daetylus and propodus on pereiopods 1 and 2 with robust apical hook. 8 - Dactylus and propodus on pereiopods 1 and 2 without apical hook 9 - Dactylus and propodus on pereiopod 1 with apical hook. Dactylus and propodus on

pereiopod 2 without apical hook. . . 15 8 Eyes well developed and fairly large. Rostrum with dorsal unarmed portion behind

subapical teeth . . . . . . norvestica - Eyes not well developed, although facets visible, no pigment in preserved specimens.

Rostrum without dorsal unarmed portion . unca sp. nov. 9 Rostrum most usually unarmed dorsally, rarely armed; ventrally 0-3 teeth . 10 - Rostrum armed dorsally and ventrally . . 11

l0 Rostrum short, reaching about to the end of the basal segment of the antennular peduncle without teeth ventrally or dorsally. Terminal margin of telson with medial spines shorter than laterals and even in length . lamiana

- Rostrum short, reaching about to the end of the basal segment of the antennular peduncle, rarely with dorsal teeth; 0 3 ventral teeth. Terminal margin of telson with medial spines longer than laterals, not even in length calmani

l l Eyes reduced, without facets; no pigment 12 - Eyes bulbous, with facets and pigment . . 13

12 Propodus and daetylus of pereiopods 3 and 4 with marginal row of strong spines crurispinata sp. nov.

- Propodus and dactylus of pereiopods 3 and 4 without marginal row of strong spines parvocula sp. nov.

13 6-8 spines of the dorsal margin of rostrum placed behind the orbit. Telson, terminal margin sub-lateral spines shorter than lateral; medial spines setose and longer than laterals . . . serratirostris

- 0-6 spines of the dorsal margin of rostrum placed behind the orbit. Telson, terminal margin medial spines shorter than l a t e r a l s . . . 14

14 Rostrum in adults very long, curving upwards with dorsal margin unarmed distally, subapieal tooth absent . x iph ias

- Rostrum reaching to base or end of 2nd segment of antennular peduncle, straight with dorsally 14 20 large distinct teeth; ventrally 3-4 distinct teeth. . hova

15 Terminal margin of telson with medial spines as long or longer than laterals 16 Terminal margin of telson with medial spines shorter than laterals 19

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Madagasean a ty id shrimps 569

16 Rostrum with ventral teeth only . ~ 17 - Rostrum with teeth dorsally and ventrally 18

17 gostrum short, with only one subapical tooth ventrally. Telson terminal margin not produced to a point centrally . . . . brevirostris brevipes

- Rostrmn of medium length, not reaching beyond peduncle of antenna 1, with more than one ventral subapical tooth. Telson terminal margin produced to a point centrally . . . . . . . angulata

18 Rostrum of medium length, not reaching beyond peduncle of antenna 1. Telson terminal margin not produced to a point centrally. Anterior margin of carpus, pereiopod 1, not deeply excavated petiti

19 Daetylus of third pereiopod with 15-17 teeth on posterior margin edulis Dactylus of third pereiopod with 3 7 teeth on posterior margin 20

20 Rostrum usually reaching to middle or end of segment 2, antennular peduncle; dorsally 6-18 teeth, seldom less or more; ventrally 0-10 teeth isaloensis grandidieri

- l~ostrum usually reaching to base or middle of segment 2, antennular peduncle; dorsally typically unarmed; ventrally 0~2 teeth, seldom 4. isaloensis isaloensis

The genus Parisia contains six recognized species: P. macrophthalma Holthuis , 1956, P. microphthalma (Fage, 1946), P. edentata Holthuis , 1956 and P. dentata sp. nov. f rom Madagascar , P. gracilis Williams, 1964 and P. unffuis Williams, 1964 from Austral ia. Gordon and Monod (1968) described Parisia? fowleri f rom a sub te r ranean lake in Zanzibar not ing t h a t the branchiae consisted of only 4 p leurobranchs and 3 epipods; the fifth p leurobranch and four th and fifth epipods being absent. Hol thu is ' (1956) generic diagnosis of Parisia states 'each of the pereiopods with a single p leurobranch ' . I t seems likely t h a t Gordon and Monod 's species belongs in Halocaridina Holthuis , 1963 bu t this can only be confirmed by re -examina t ion of the type material .

K e y to Parisia 1 Eyes strongly reduced with, at most, only a pigment spot 2

Eyes with the cornea well developed and rounded, provided with dark pigment. Rostrum well developed, with teeth on both upper and lower margins . . . . . . . macrophthalma

2 Rostrum well developed, reaching about to the end of the antennular peduncle, with teeth on both upper and lower margins . microphthalma

Rostrum short, hardly reaching beyond the eyes 3 3 Pereiopods 1 and 2 with an apical hook on dactylus and propodus unguis

Pereiopods 1 and 2 without such a hook 4 4 Rostrum with 2-6 dorsal spines, unarmed ventrally dentata sp. nov.

Rostrum unarmed either dorsally or ventrally . 5 5 Carpus of pereiopod l short, less than twice as long as high. and deeply excavate.

Fifth abd<)minal pleurae posteroventral margin br<)adly rounded. Adult eye typically with pigment spot . . • . . . . edet~lata

Carpus <>f pereiopod 1, 2½ times as long as broad, not or scarcely excavate. Fifth abdominal i)leurae posteroventral angle acute. Adult without eye spot pigmentation gracilis

D e s c r i p t i o n s

Caridina parvoeula sp. nov. Etymology: F r o m the latin parvus meaning small; oculus meaning eye. Locality: Dark zone, la Grot te d 'An t sa t robonko , A n k a r a n a Massif, 40 km nor th

o f Ambilob6 and 60 km south of Diego Suarez, no r the rn province o f Diego Suarez. 18th September' , 1981.

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Type material: HOLOTYPE, ~ (figs. 1 3), 30mm., registration number BM(NH) 1983.21; rA~ATY~'E, ~ (figs. 1C & 3H), 24ram., registration number BM(NH) 1983.22.

Eyes: Reduced and without facets. Rostrum (fig. 1 B): Long, gradually curving upwards and reaching beyond the

antennal scale with 23 24 dorsal teeth and l l -14 ventral teeth.

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FIG. l. Caridina parvocula, 9, HOLOTYPE, BM(NH) 1983.21: A, antenna 2, B, rostrum, PARATYPE BM(NH) 1983.22, C, appendix masculina, 9, ttOLOTYPE, D, maxilla 2,

E. antenna 1. F, mandible. Scales= 1 mrrL

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Madagascan atyid shrimps

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FIG. 2. Caridina parvocula, ~, HOLOTYPE~ BM(NH) 1983.21: A, maxilla I, B, maxilliped 1, C. telson, l), maxilliped 3. E. maxilliped 2. F. ple<>t>(>d 1. ~cales= 1 ram.

Carapace: Lower orbital angle rounded, antennal spine sharply pointed. Ptery- gostomial angle broadly rounded.

Antenna I (fig. 1 E): Peduncle 3-segmented. Stylocerite ¼ length of basal segment, pointed, divided from segment almost to base. Segment 1 with submarginal setose setae, antero-lateral angle distinct and pointed. Segment 2 with submarginal setose setae. Segment 3 with expanded lobe fringed laterally and apically with setose setae. Flagella long. simple.

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13 /

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l Fl~. 3. Caridina parvocula, ~, HOLOTYPE, BM(NH) 1983.21: A, pereiopod l, B, pereiopod 2,

C, pereiopod 3, D, pereiopod 4, E, pereiopod 5, F, pleopod 2, G, diaeresis of uropod, H, ~' PARATYPE BM(NH) 1983.22 pleopod 1. Scales = l ram.

Antenna 2 (fig. 1 A): Scaphocerite longer than the antenna 1 peduncle in situ. Setose setae on inner and anterior margins. Outer margin without setae, ending anteriorly in a strongly pointed subapical tooth. Peduncle about ½ length of scaphocerite. Flagellum long and simple.

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Madagascan atyid shrimps 573

Maadibles (fig. 2 F): Laterally 7 or 8 incisor teeth; medially two groups of simpte setae, first containing l0 or 12, second approximately 30, finer, regularly placed setae; ridged molar process.

Maxilla 1 (fig. 2A): Lower lacina (not figured) with broadly rounded margin bearing rows of setae, centrally very long. A submarginal band of setae and a few proximally. Upper lacinia broadly elongate, inner margin straight bearing rows of strong spines and short setae. Setae on outer and lower inner margins. Palp truncate with long setose seta. Robust spine apically plus vertical row of long, unarmed setae.

Maxilla 2 (fig. 1 D): Upper endite cleft with marginal and submarginal simple and dentieulate setae on the distal portion; proximal portion with many transverse rows ofsetae. I,owcr endite with regular row of long marginal setae and a few irregularly placed submarginal setae. Palp narrow and rounded apically, not quite as long as cleft of upper endite, with a few setae. Seaphognathite not reaching as far as endite with marginal setose setae continuing down to proximal triangle process which has many long unarmed setae on proximal margin.

Maxilliped 2 (fig. 2 E): Endopodite ultimate and penultimate segments indis tinetly divided, reflected against basal segments. Inner margin of ultimate segment submarginal long rows of dentieulate and simple setae plus transverse rows of simple setae on lower portion. Palp triangular with broad base and one sub-apical seta. Exopod flagellum long and narrow with marginal setose setae. Caridean lobe not broad with marginal setose setae.

Maxilliped 2 (fig. 2 E): Endopodite ultimate and penultimate segments indist- inctly divided, reflected against basal segments. Inner margin of ultimate segment with long simple setae. Inner margin of penultimate segment with long setose setae and shorter, simple setae. Inner margins of basal segments wit h long simple setae and some setose setae proximally. Exopod long with marginal setose setae distally and proximally. Podobranch broad and fiat, with reduced branchial lamella.

Maxilliped 3 (fig. 2 D): Endopod 3-segmented. Segment 1 about 5 times as long as broad with a few simple marginal setae. Segment 2 about 6 times as long as broad but shorter than segment 1 with groups of transverse rows of simple setae. Segment 3 shorter than 2 with large robust apical spine surrounded by simple setae and approximately 6 spines on distal inner margin; proximally approximately 12 transverse rows of'simple setae. Exopod narrow and reaching midway along 2nd endopod segment and armed with distal marginal setose setae. Epipod hooked at distal end and with proximal extension. 2 arthrobranchs (not figured), the second, distal much smaller than the first, proximal.

Branchi<d .fo~ula: Maxillipeds Pereiopods 1 2 3 1 2 3 4 5

pleurobranehs 1 1 1 1 1 arthrobranchs - 2 1 podobranchs - 1 - epipods - 1 1 1 1 1 1 - exopods 1 1 1

Pereiopod 1 (fig. 3 A): Chela long and slender, dactylus slim, propodus similar, both with tips evenly rounded without hooks but with numerous long, simple and denticulate setae. Carpus as long as rectus but narrower and distally excavate. Merus slightly broader than carpus. Ischium about twice length of coxa. Epipod hooked distally. Two setobranchs.

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Pereiopod 2 (fig. 3B): Chela long and slender, two thirds as long as carpus, dactylus much longer than palm of propodus, tips evenly rounded without hooks but with numerous long, simple and denticulate setae. Carpus narrow and slightly longer than merus, slight excavation distally. Merus broader than carpus. Ischium over twice length of coxa. Epipod hooked distally. Two setobranchs.

Pereiopod 3 (fig. 3 C): Dactylus about 1 length of propodus with prominent apical spine and 5 marginal spines reducing in size proximally. Propodus about twice length of carpus. Carpus with strong spine near margin of propodus plus smaller medial spines. Merus slightly longer than propodus with 3 strong marginal spines. Ischium about twice length of coxa with one strong spine distally. Epipod hooked distally. Two setobranchs.

Pereiopod 4 (fig. 3 D): Dactylus about ¼ length of propodus with strong apical spine and 6 marginal spines reducing in size proximally. Propodus about twice length of carpus with several small spines medially. Carpus with a strong spine distally and approximately 4 smaller spines running proximally. Merus about 3 times length of ischium with 4 strong spines on lateral margin. Ischium about twice length of coxa with one strong spine distally. Epipod hooked distally. Two setobranchs.

Pereiopod 5 (fig. 3 E): Dactylus about i length of propodus with a strong apical spine and a regular row of medial spines. Propodus narrow, just over twice as long as carpus with two small spines on the distal margin and several medial spines. Carpus with one strong spine distally. Merus not as long as propodus, but broader with 3 strong marginal spines. Iscbium about twice length of coxa. No epipod. Two setobranchs.

Pleopod 1: HOLOTYPE female, BM(NH) 1983.21 (fig. 2 F), endopod nearly ½ length ofexopod gradually tapering to apex with setose setae on inner margin. Exopod with marginal setose setae plus submarginal siml)le setae. PARATYPE male. BM(NH) 1983.22 (fig. 3 H). endopod ¼ length of exot)od with marginal setose setae. No appendix interna.

Pleopod 2: HOLOT~ePE female, BM(NH) 1983.21 (fig. 3 F), endopod only slightly shorter than exopod both with marginal setose setae. Appendix interna of endopod with many retinacula at tip. PAI%ATYPE male, BM(NH) 1983.22 (fig. 1 C), similar to holotype with appendix masculina at base of endopod which has 12 long robust setae in figured specimen.

Pleopods 3-5: Similar to pleopod 2 of tbmale. Telson (fig. 2 C): Very nearly as long as 6th abdominal segment, tapering distally.

Dorsal surface with 4 and 5 spines, usually in pairs and submarginal. Terminally 1 pair of very short spines outermost, a lateral pair longer than the 5 intermediate setose setae. In the paratype specimens there are 4 pairs of dorsal, submarginal spines and a total of 10 terminal spines including the shorter outermost and longer lateral pairs.

Uropods: Exopod slightly larger than endopod both with marginal setose setae and small unarmed setae submarginally. Diaeresis (fig. 3 G) on exopod produced laterally to pronounced point with 11 and 12 robust spines. Paratype specimen with 13 spines on both diaereses.

Caridina erurispinata sp. nov. Etymology: From the latin cru8 meaning leg; spina meaning spine. Locality: Dark zone, la Grotte d'Antsatrobonko, Ankarana Massif, 40 km north

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o[" A m bilob6, 60 km south of Diego Suarez, northern province of Diego Suarez. 18th September, 1981.

7'!11~, m¢deri<d: HOI:OTYPE, ~ (figs. 4 7). 30mm., registration number BM(NH) 19S3.23: PAmX'rYPE. 3-27 mm.. registration number BM(NH) 1983.24.

Eyes: l~edueed and without/i~cets. Rostrum (fig. 4 B): Long, gradually curving upwards and reaching beyond the

antennal scale, with 21-26 dorsal teeth, 6-11 ventral teeth. Carapace: Similar to that of C. parvocula. Antenna 1 (fig. 4 A): Similar to that of C. parvocula. Stylocerite ¼ length of basal

segment, pointed and divided from segment by just under half its length. Antenna 2 {fig. 4C): Similar to that of C. parvocula. Mandibles (fig. 4E): Laterally 7 8 incisor teeth, medially two groups of fine

simple setae. The second group with about twice as many very regular setae. A ridged molar process laterally.

Maxilla 1 (fig. 4 D): Lower laeinia not seen clearly in the dissection. Upper lacinia similar to that of C. parvocula. Palp truncate with robust spine and two long setose setae distally plus median rows of long simple setae.

Maxilla 2 (fig. 5 B): Similar to that of C. parvocula. Maxilliped I (fig. 5 A): Similar to that of C. parvocula. Endite of basis long with

small subapical protrusion on inner border. Palp with broad base, extended apically and with row of marginal setose setae distally.

Maxilliped 2 (fig. 5 C): Similar to that of C. parvocula. Maxilliped 3 (fig. 5 D): Endopod 3-segmented. Segment 1 about 6 times as long as

broad with a few marginal setae proximally. Segment 2 about 7 times as long as broad but shorter than segment l, with groups of transverse rows of setae. Segment 3 shorter than segment 2 with large robust apical spine surrounded by short setae; 8 spines on distal margin, proximally approximately 12 transverse rows of setae decreasing in length proximally. Exopod, epipod and arthrobranchs similar to that of C. parvocula.

Branchial formula: Similar to that of C. parvocuta. Pereiopod 1 (fig. 6 B): Chela long and slender but shorter than chela of pereiopod

2; dactylus slim, propodus similar, tips evenly rounded without hooks but with numerous long, simple and denticulate setae. Carpus slightly longer than merus, narrower and excavate distally. Ischium about 11 times length of coxa. Epipod hooked distally. Two setobranchs.

Pereiopod 2 (fig. 6 D): Chela long and slender, dactylus slim, propodus similar, tips evenly rounded without hooks but with numerous long, simple and dentieulate setae. Carpus narrower but just longer than merus and excavate distally. Ischium about twice length of eoxa. Epipod hooked distally. Two setobranehs.

Pereiopod 3 (fig. 6 C): Dactylus about ¼ length of propodus with robust apical spine and approximately 9 marginal spines, reducing in size proximally. Propodus about twice length of carpus with a row of strong spines running along the proximal +. Carpus also with row of strong spines running along its length. Merus just over three times the length of the ischium and with three large, robust marginal spines. Coxa approximately ½ length of ischium. Epipod hooked distally. Two setobranchs.

Pereiopod 4 (fig. 6 A): Daetylus about ¼ length of propodus with prominent apical spine and 8 marginal spines decreasing in size proximally. Propodus just under twice the length of carpus with a row of strong spines running proximally along its length. Carpus also with a row of strong spines running along its length. Merus about 3 times

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Flu. 4. Ca~'idip,I Cr~vis'yit,lt~l. ~. ,IOL(YrV,'E. BM(NH) 1983.23: A. antenna 1, B, rostrum, C, antenna 2, 1). maxilla I. E. mandible. Scales=.l ram,

length ofischium with 3 large robust marginal spines. Ischium about twice the length of coxa with one large, marginal spine. Epipod hooked distally. Two setobranehs.

Pereiopod 5 (fig. 7 A): Dactylus approximately ¼ length of propodus with marginal row of spines plus one strong apical spine. Propodus about twice length of carpus and with a large and small spine on antero-lateral margin and 6 marginal spines decreasing in size proximally. Merus just over twice the length of ischium with 2 large, robust marginal spines. Coxa about ~ length of ischium. No epipod. Two setobranchs.

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FIG. 5. Caridina crurispinata, ~, HOLOTYPE, BM(NH) 1983.23: A, maxilliped 1, B, maxilla 2, (?. maxillil)ed "2. !). maxillit)e(l 3. Scales= 1 ram.

Pleopod 1 (fig. 7B): Exopod approximately 4 times length of endopod with marginal setose setae. Endopod broad with setae on ~ of inner margin, and along the whole length of outer margin. No appendix interna.

Pleopod 2 (fig. 7 C): Exopod just longer than endopod, both with marginal setose setae. Appendix masculina ½ length of endopod with about 14 long marginal setae. Appendix interna similar in length with many retinacula at tip.

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FH,. 6. ('<tridimt crurispinatel. ~, HOLOTY|'E. [~M(NH) 1983.23: A, pereiopod 4, B, pereiopod I. C. l)ereiot)od 3, 1), i>ereiol)od 2. Scales = 1 mm.

Pleopods 3 5: Similar to pleopod 2 but without appendix masculina. Telson (fig. 7 D): Tapers distally. Dorsal surface with 4 submarginal pairs of

spines. Terminally 10 spines; including the very short outer pair: the longest lateral pair, with 6 intermediate pairs.

Uropods: Exopod slightly longci than the endopod both with marginal setose setae. Diaeresis (fig. 7 E) on exopod produced laterally to a pronounced point with 13 robust spines on holotype, 13-15 on paratype.

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Madagascan atyid shrimps 579

B

f C

)t

D

E

\ \

FIG. 7. Caridina cruri,~'pinata, ~, HOLOTYPE, BM(NH) 1983.23: A, pereiopod 5, B, pleopod l, C, appendix interna and appendix masculina on pleopod 2, D, telson, E, diaeresis of uropod. Scales= 1 mm.

Caridina unca sp. nov. Etymology: From the latin uncus meaning hook, Locality: Dark zone, Grotte d'Antsatrabonko, Andarana Massif, northern

province of Diego Suarez, North Madagascar. 18th September, 1981. Type material: HOLOTYPE, ~ (figs. 8--11), 32 mm., registration number BM(NH)

1983.25; PARATYPES, C~, 21'5 ram., 9, 29 ram, c~, 28 mm., and 9, 33 ram., registration numbers BM(NH) 1983.26 29.

Eyes: Not large or bulbous, but facets visible although there is no pigment in the preserved specimens.

Rostrum (fig. 8C): Long, gradually curving upwards and reaching beyond antennal peduncle. 27-28 dorsal spines, 12-13 ventral spines.

Carapace: Lower orbital angle distinct, antennal spine sharply pointed. Ptery- gostomial angle broadly rounded and slightly produced.

Antenna I (fig. 8A): Similar to that of C. parvocula. Antenna 2 (fig. 8 I3): Similar to that of C. parvocula.

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580

A

A. E. Gurney

B

L__ I

1

FIG. 8. tWridim~ Hl~r( I . ~', H()IJOTYPE BM(NH) 1983.25: A, antenna 1, B, antenna 2, U, rostrum, D, maxilla 2, E, maxilla 1. Scales= 1 ram.

Mandibles (fig. 9 C): Laterally W9 incisor teeth, medially one group of 8-9 simple setae, second group of approximately 30 regularly placed setae; lateral ridged molar process. Proximal portion bearing several short setose setae.

Maxilla 1 (fig. 8E): Upper lacinia broadly elongate with outer marginal and submarginal setose setae. Inner margin straight with 2 rows of strong spines and submarginal row of simple setae. Palp reduced with long simple setae and robust spine apically. Lower lacinia submarginal rows of short simple setae. Marginal rows of long setose setae and shorter setose setae.

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Madagascan atyid shrimps

R

581

i

FIG. 9.

J

E D L _ _ r

Caridina unca, ~, ROLOTYPE BM(NH) 1983.25: A, maxilliped 3, B, maxilliped 1, C, mandible, D, maxilliped 2. E, telson. Scales= 1 mm.

Maxilla 2 (fig. 8 D): Similar to tha t of C. parvocula. Scaphognathite just longer than endite.

Maxilliped 1 (fig. 9 B): Similar to tha t of C. parvocula. Maxill@ed 2 (fig.. 9 D): Similar to tha t of C. parvocula. Maxilliped 3 (fig. 9 A): End0pod 3-segmented, segment 1 about 4 times as long as

broad covered in par t with simple setae. Segment 2 about five times as long as broad but shorter than segment 1 with transverse groups of simple setae. Segment 3 shorter

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582 A. t~. Gurney

B

I

i! !% D

C

f -

\

/

E

Fro. 10. ('<~ridina unca, ~2, HOLOTVP~ BM(NH) 1983.25: A, pereiopod 1, B, pereiopod 3, C. I~ereiopod 2, l), pereiopod 4. E. diaeresis of uropod. Scales = 1 mm.

than second with a large robust apical spine surrounded by simple setae and distal robust spines on inner margin. Proximal ly approximate ly 12 transverse rows of simple setae. Exopod, epipod and ar throbranehs similar to tha t of C. parvocula.

Branchial formula: Similar to tha t of C. parvocula. Pereiopod 1 (fig. 10A): Chela broad, daetylus stout, not long, propodus broad,

both with a strong hook at apex surrounded by long simple and dentate setae. Carpus shorter' than chela and broad, distal margin excavate. Merus about same length as

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Madagasean atyid shrimps 583

A

B

f

Fro. 11. Cardina unca, 2, HOLOTYPE BM(NH) 1983.25: A, pereiopod 5, B, pleopod 1. ScMes = I ram.

carpus and stout. Isehium approximately 1 length of merus similar to length of eoxa. Coxa with pronounced lobe and ridge with a vertical row of short simple setae, plus a group of 4 long simple setae on inner proximal corner. Epipod hooked distally. Usually 3 setobranchs, occasionally 2.

Pereiopod 2 (fig. 10C): Chela, although broad not as broad as in pereiopod 1. Dactylus stout, fairly long, propodus broad, both with a strong apical hook although these are not as robust as those on 1st pereiopod. Each hook surrounded by simple and dentate setae. Carpus long, not broad, but widening distally. Merus about the same in length as carpus, not broad. Ischium shorter than merus. Coxa shorter than ischium with pronounced lobe and ridge which has vertical row of short, simple setae, plus a group of 4 long simple setae on inner distal corner, several simple setae covering remainder. Epipod hooked distally. Usually 3 setobranchs, or 2.

Pereiopod 3 (fig. 10 B): Dactylus approximately ½ length ofpropodus with strong apical spine and 5 marginal spines reducing in size proximally. Propodus with marginal and submarginal spines and many simple setae. Carpus just over ½ the length of merus with a strong distal spine, 3 small marginal spines and many small simple setae. Merus with 3 strong marginal spines and many simple setae. Isehium less than ½ length of merus with a strong marginal spine distally. Coxa shorter than isehium and produced distally and with a trace of a vertical row of small simple setae; approximately 5 simple setae on inner distal anterior corner, several simple setae covering remainder. Epipod hooked distally. Usually 3 setobranehs, or 2.

Pereiopod 4 (fig. 10 D): l)actylus approximately ½ length of propodus with strong apical spine and 5 marginal spines reducing in size proximally. Propodus with marginal and submarginM spines and many simple setae. Carpus just over ½ the length of merus with a strong distal spine; approximately 4 small marginal spines distally and several simple setae. Merus with 3 strong marginal spines and many simple setae. Ischium approximately ½ length of merus covered with many simple

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584 A . g . Gurney

setae. Coxa shorter than merus produced distally and with a vertical row of small simple setae and approximately 5 simple setae on inner distal anterior corner, several simple setad covering remainder. Epipod hooked distally. Usually 3 setobranchs, or 2.

Pereiopod 5 (fig. II A): Dactylus approximately ¼ length of propodus with a strong apical spine and a regular row of smaller marginal spines and simple setae submarginally. Propodus with marginal and submarginal spines and many simple setae. Carpus just over ½ length of merus with approximately 2 strong marginal spines and many simple setae. Ischium approximately ½ length of merus with many simple seta. Coxa approximately ½ length of merus produced distally, with approximately 4 simple setae on distal anterior corner, several simple setae on remainder. No epipod. Usually 2 setobranchs.

Pleopod I (fig. 11 B): ~ endopod } length of exopod and gradually tapering to apex with setose setae on outer margin. No appendix interna. Exopod with regular rows of long setose setae on both margins and several small simple setae submarginally. Peduncle, outer margin, with marginal row of small spinules distally and curving submarginally proximally. Small spinules also present on inner margin submedially. 6 strong spinules grouped on inner proximal corner of peduncle.

Pleopod 2: ~ endopod 3 length of exopod, both with marginal setose setae. Appendix interna arising near to base ofendopod and with rectinacula at tip. Outer margin of peduncle with a few sparse setae running proximally. Inner margin with a few spinules; a group of 6 strong spinules in inner proximal corner.

Pleopod 3: Similar to pleopod 2. Pleopod 4: Exopod and endopod similar to pleopod 2. Peduncle outer margin with

a few simple setae. Inner margin with a few spinules including a group of 4 on inner proximal corner.

Pleopod 5: Exopod and endopod similar to pleopod 2. Peduncle outer margin with long setae some setose. Inner margin with less, shorter setose setae and a group of inconspicuous spinules.

Telson (fig. 9E): Almost as long as 6th abdominal somite. 4 pairs of spines on dorsal surface. Between 9 and 19 terminal setae. The outer spines are veryshort, the laterals long, the medials vary in length but are shorter than the laterals.

Uropods: Exopod larger than endopod both with marginal setose setae. Diaeresis (fig. 10E) produced laterally to pronounced point and with l 1-13 small, strong spines.

Parisla dentata sp. nov. Etymology: From the latin dentatus meaning toothed. Locality: Dark zone, Grotte d'Andrafiabe, Ankarana Massif, 40km north of

Ambilob6 and 60 km south of Dieg0 Suarez, northern province of Diego Suarez. 17th September, 1981.

Type material: HOLOTYPE, ~ (figs. 12-14), 14 mm., registration number BM(NH) 1983:30; PARATYPES, ~, l l ' 5 m m , 2, 9-5mm., registration numbers BM(NH) 1983:31-32, respectively.

Eyes: Reduced without facets and very little pigment, if any. Rostrum (fig. 12C): Very short, not reaching further than the eyes, unarmed

ventrally. Antenna 1 (fig. 12 A): Typical for genus. Styloeerite reaching ~- length of first

segment of the peduncle, but not divided from it by more than a ½ length of stylocerite.

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A B

Madagascan atyid shrimps

C

(

585

/

s

/+

?

? / / / /

N

I +//

[ _ _ j

FIG. 12. Parisia dentata, ~, HOLOTYPE BM(NH) 1983.30: A, antenna l, B, antenna 2, C, rostrum, D, maxilla 2, E, maxilla 1. Scales= 1 ram.

Antenna 2 (fig. 12 B): Scaphocerite about 2¼ times as long as broad tapering distally and with a regular row of setose setae on inner and anterior margins. Outer margin with only a few simple setae, ending with a subapical projection. Peduncle about ½ length of scaphocerite, flagellum long and simple.

Mandibles (fig. 13 B): Lateral incisor process with 5-7 teeth. Medially 4-6 setae followed by regular row of approximately 20 setae. Lateral ridged molar process. Proximal portion with a few short, simple setac.

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586 A.R. Gurney

h B , C

\ / /

[g \

Fro. 13. Parisia dentata, ~, 1KOLOTYPE BM(NH) 1983.30: A, telson, B, mandible, C, maxilliped 1, D, maxilliped 2, E, maxilliped 3, F, segment 3 of maxilliped 3.

Scales= l ram.

Maxi l la 1 (fig. 12 E): Lower lacinia (not figured) with long marginal setae and submarginal row of much smaller setae. Upper lacinia broadly elongate with outer marginal and submarginal long setae. Inner margin straight with marginal rows of strong spines and short setae. Palp reduced with long apical and submarginal setae plus one robust spine on subapical lobe.

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Madagascan atyid shrimps 587

h 13

D

E

G

F

FIG. 14: Parisia dentata, ~, I{OLOTYrE BM(NH) 1983.30: A, pereiopod 3, B, pereiopod 2, C, pereiopod 4, D, pleopod l, E, pereiopod 5, F, diaeresis of uropod, G, pereiopod I. Scales = 1 ram.

Maxilla 2 (fig. 12D): Typical for genus. Maxilliped 1 (fig. 13 C): Typical for genus. Maxilliped 2 (fig. 13D): Typical for genus. MaxiUiped 3 (figs. 13E, F): Endopod 3-segmented. Segment 1, 5 times longer

than broad with simple marginal setae. Segment it about 4½ times as long as broad

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588 A.R. Gurney

with a few medial groups of simple setae. Segment 3 shorter than 2 with robust apical spine and 5 medial spines distally; proximally approximately 12 transverse groups of setae. Epipod hooked distally, extended proximally.

Branchial formula:

pleurobranehs arthrobranehs - - 1

podobranehs - 1 -

epipods 1 1 1 exopods 1 1 1

Maxill@eds Pereiopods 1 2 3 1 2 3 4 5

1 1 1 1 1

1 1 1 1 -

Pereiopod 1 (fig. 14 G): Chela broadish, dactylus and propodus with tips evenly rounded without hooks but with tufts of long, denticulate and simple setae. Carpus shorter than merus broadening distally and excavate. Merus slightly longer than carpus. Ischium about l½ times longer than coxa. Epipod hooked distally. 1 setobranch.

Pereiopod 2 (fig. 14 B): Chela broadish, dactylus and propodus with tips evenly rounded without hooks and with tufts of long, denticulate and simple setae. Carpus about as long as merus. Ischium about 3 times length of coxa. Epipod hooked distally. 1 setobranch.

Pereiopod 3 (fig. 14 A): Dactylus less than length of propodus with strong apical spine and 4 marginal spines reducing in size proximally. Propodus about l½ length of carpus with several small spines and setae on surface. Carpus also with a small spine and setae. Merus broader but about same length as propodus with 3 strong medial spines. Ischium shorter than carpus with a medial strong spine distally. Coxa damaged, not figured. Epipod hooked distally. 1 setobranch.

Pereiopod 4 (fig. 14 C): Daetylus with strong apical spine and five marginal spines. Propodus over 4 times as long as dactylus with marginal and submarginal spines and setae. Carpus about 2 length ofpropodus with a small spine and setae. Merus broader but about same length as propodus with 3 strong medial spines. Ischium shorter than carpus with a strong spine distally. Coxa damaged, not figured. Epipod hooked distally. 1 setobraneh.

Pereiopod 5 (fig. 14E): Dactylus about ¼ length of propodus with strong apical spines and a regular row of smaller marginal spines. Propodus with marginal and submarginal spines. Carpus about ½ length of merus. Merus broader than propodus but approximately same length with 2 strong marginal spines. Isehium almost twice length of coxa. No epipod. 1 setobranch.

Pleopod 1 (fig. 14 D): Female endopod } length of exopod with setose setae on inner margin. Exopod with marginal setose setae distally.

Pleopod 1: Male endopod about ¼ length of exopod with setose setae on inner margin, 4 setae on outer margin. No appendix interna. Exopod with setose setae on both margins.

Pleopod 2: Male exopod with marginal setose setae. Endopod similar to exopod but shorter. Appendix interna with retinacula at tip, appendix masculina with approximately nine strong, unarmed setae.

Telson (fig. 13 A): About same length as 6th somite. Dorsal surface with 3 pairs of submarginal spines. Terminally, 1 pair of small spines outermost; the lateral pair

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Madagascan atyid shrimps 589

longest; the sublateral pair shorter than the next intermedial pair; the inner, medial, pair, short.

l'r,p<Ms: Exopo<ts slightly larger than en<tol~o<ls both with marginal setose selae. I)iaeresis (fig. 13 F) pro<tueed laterally to pronounced point with 7-9 robust spines.

Discussion The apical hook character occurs independently in a number of the genera in the

family Atyidae. Hiekson (1888) clearly figured hooks on the dactylus and propodus ofpereiopods 1 and 2 in his description ofAtya wyclcii from Celebes, since relegated to a variety ofCaridina nilotica by de Man (1908). In 1928, J. goux reported 'un court et mince stylet ' between the hairs on the chela of pereiopod 1 on the type specimens of Pseudoatya beauforti from B atjan Indonesia. Recently, Williams (1964) described Parisia unguis from Australia with 'tips sharply clawed' on pereiopods 1 and 2, although his figures show a robust hook. In 1906 Caiman figured Caridella minuta from Tanganyika indicating the presence of an apical point on the daetylus and propodus of pereiopods 1 and 2 but made no mention in his description to substantiate this observation. Re-examination of the types revealed the presence of robust hooks on the dactylus and propodus of the first and second pereiopods. Apical hooks are not present on the types of Caridella cunningtoni Calman, 1906. In his description of C. paslci Calman (1928) notes that it is 'distinguished from its two congeners by a conspicuous too th on the moveable finger'. Bouvier (1925), in his 'Monographie des Atyides', noted this character for the first time in several species from various genera, notably the genus Limnocaridina. Bouvier figures apical hooks in all the known species whereas Caiman (1899) in his description of the genotype L. tanganyilcae, misleadingly described the 'fingers' of the first pereiopods as 'opposed margins [that] bear series of small stout spinules increasing in size towards the tip, but there is no strong terminal hook as in C. wyckii'. Subsequently Bouvier (1925) figured 3 strong robust apical hooks and draws an analogy with Xiphocaris elongata (Gubrin-Men6ville, 1857) regarded by Fryer (1977) as the most primitive extant atyid.

Further observations on the presence or absence, size and number of apical hooks on the chela of the 1st and/or 2rid pereiopod should prove a most useful key- character for species determination in the Atyidae.

Summary Three new species of Caridina and one new species of Parisia from caves in

northern Madagascar are described and figured. A useful key-character, the presence or absence of an apical hook on the daetylus and propodus ofpereiopods 1 and/or 2, is re-evMuated. Revised keys are provided for both genera.

Acknowledgments I would like to thank Drs. A. A. Fincham and R. J. Lincoln for reading and

commenting on the manuscript. I am also grateful to Drs. L. B. Holthuis and J. Forest for arranging the loans of Madagasean specimens.

References BOUVlEI~, E.-L., 1925, t~echerches sur la Morphologie les Variations, la Distribution

syst6matique des erevetes d'eau douce de la famille Atyid~s, Encyclopedic Entomo- logique, IV, 1-370.

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590 M a d a g a s c a n a t y i d sh r imps

CALMAN, W. T., 1899, On two species of Macrurous Crustaceans from Lake Tanganyika, Proceedings of the Zoological Society of London, 1899, 704-712. 1906, Zoological results of the third Tanganyika Expedition. Report on the Macrurous Crustacea, Proceedings of the Zoological Society of London, 1906, 187-206. 1928, On Prawns of the Family Atyidae from Tanganyika, Proceedings of the Zoological Society of London, 1928, No. XLIX, 737-741.

COUT~RE, H., 1899, Sur quelques Macrures des eaux douces de Madagascar, Bulletin du Musdum d'Histoire Naturelle, Paris, 1899, 382-383.

FAOE, L., 1946, Sur une Caridine nouvelle cavernicole de Madagascar, Caridina microphthalma nov. sp., Bulletin du Museum d'Histoire Naturelle, Paris, 18, 324-327.

FRYER, G., 1977, Studies on the functional morphology and ecology of the atyid prawns of Dominica, Philosophical Transactions of the Royal Society of London, 277 (B), 57-128.

GORDOn, I., and MONOD, TH., 1968, Sur quelqucs Crustac~s des eaux douces de Zanzibar, Bulletin de l'Institut Fondamental d'Afrique Noire, XXX, set A, No. 2, 497-517.

GU~aI~-Mnc*~VILLE, F.-E., 1857, Crustac~s (Histoire . . . de File de Cuba par Ramon de la Sagra), trad. VII.

HIcssoc*, S. J., 1888, On a new species of the genus Atya (A. Wyckii) from Celebes, Annals and Magazine of Natural History, London, 6 (2), 357-367.

HOLTItUIS. L. B., 1963, On red coloured shrimps (Decapoda. Caridea) from trot)ieal land- locked saltwater pools, Zooloffi,sche Mededeelingen, Leiden, XXXVIII. (16), 261 279.

- - - 1956 a, An crmmeration of the ( 'rustacea l)eeapoda Natantia inhabiting subteH'a,man wate,'s. |'is el 31ilie~. 7, 43 76. 1956b, The trogh)bic Atyi(lae of Madagascar, Mdmoires de l'lnstitut Seienlifique de .lladaga,~car. set A, 11, 97 110. 1965, The Atyidae of Madagascar, M dmoires du Mu~vdum National d'Histoire Naturelle, 33 (1), 1-48.

MAC*, DE J. G., 1908, On Caridina nilotica (t~oux) and its varieties, Records of the Indian Museum, 2, 225-283.

MILC*n-EDwARDS, H., 1837, Histoire naturelle des Crustac6s, 2, 362-363. Roux, J., 1928, Notes carcinologiques de l 'archipcl Indo-Australian. I. D6capodes macroures

d 'eau douce, Treubia, 10, 197-224. Roux, P., 1833, Lettre relative ~ diverses eoquilles, Crustac6s, Insectes, Reptiles et Oiseaux

observ6s en Egypte, Annales des Sciences Naturelles, Paris, 28, 72-77. WILLIAMS, W. D., 1964, Subterranean freshwater prawns (Crustacea: Decapoda: Atyidae) in

Australia, Australian Journal of Marine and Freshwater Research, 15, 93-106.

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