Hydrobiologia 223: 293-299, 1991. L. Watling (ed.), Vllth International Colloquium on Amphipoda. 293 © 1991 Kluwer Academic Publishers. Printed in Belgium.

A new marine interstitial ingolfiellid (Crustacea, Amphipoda, Ingolfiellidea) from Tenerife and Hierro

Ronald Vonk1 & Elias Sanchez2 1 Institute of Taxonomic Zoology, University of Amsterdam, P.O. Box 4766, 1009 AT Amsterdam, Netherlands; 2Departamento de Zoologia, Universidad de La Laguna, Tenerife, Spain

Key words: Crustacea, Amphipoda, Ingolfiella canariensis n. sp., Tenerife, marine interstitial

Abstract

Ingolfiella canariensis n. sp., from coarse sand and gravel in the mediolittoral zone of Tenerife and Hierro, Canary Islands, is described. The new species shares supposedly apomorphous characters with species from comparable habitats from the Andaman Islands, Bermuda and Curacao (Netherlands Antilles). The female of Ingolfiella similis Ronde-Broekhuizen & Stock, 1987, from Fuerteventura is also described.

Introduction information about the male is often lacking in species from marine shallow waters. This may Along the rugged northwest coast of Tenerife account for the disjunct areas in which species of many good sampling sites for interstitial stygo- these subgenera occur. bionts are present. The narrow volcanic sand Since /. canariensis could not be placed in beaches sheltered by capes and ridges of often either the subgenera Tethydiella Ruffo & Vigna- recent lava outflows harbor Ingolfiella, Bogidiella, Taglianti or Gevgeliella S. Karaman (sensu Stock, Psammogammarus and other still unidentified 1976) to which it comes closest when character eyeless and microphthalmous amphipods. states are compared, we refrain from subgeneric The genus Ingolfiella now consists of 28 division. species, /. canariensis n.sp. included. Recently Ruffo & Vigna-Taglianti (1989) divided the genus Ingolfiella canariensis n. sp. into 7 subgenera. They drew the conclusion from their cladogram that Ingolfiella is divided into two Material distinct groups of subgenera characterized by the presence or absence of ocular lobes. The first TENERIFE (Canary Islands), 1 c? holotype. group occurs in the marine environment {Ingolf­ Station 88-594: Punta del Hidalgo, Playa de los iella s. str., Hansenliella, Tethydiella). The second Troches (UTM coordinates CS group is associated with fresh- and anchihaline 37170 x 316110), 0.3 m under sediment surface, waters. The two groups seem well defined from an low-tide mark, Bou-Rouch (BR) biophreatical ecological point of view. pump (see Bou, 1975); 12 Dec. 1988 (Zoologisch Biogeographically, they are less well defined for Museum Amsterdam, ZMA, Amph. 108.650). 294

7 2 2 paratypes. Stn. 88-592: Puntadel Hidalgo, EL HIERRO (Canary Islands) beach in harbor (UTM coord. CS 1 S, 5 2 2 paratypes. Stn. 87-47: Tamaduste, 36985 x 316015), 0.5 m under sediment surface, harbor (UTM coord. BR 21515 x 308120), low-tide mark, BR pump; 10 Dec. 1988 (ZMA Karaman-Chappuis method (digging a hole) in Amph. 108.651). muddy sand in mediolittoral zone; 29 Apr. 1987 2 2 2 paratypes. Stn. 88-593: Punta del Hidalgo, (ZMA Amph. 108.660). beach left of harbor (UTM coord. CS 4 2 2 paratypes. Stn. 88-560: La Restinga, 36975 x 316015), 0.4 m under sediment surface, Jameos del Puerto (UTM coord. BR low-tide mark, BR pump; 11 Dec. 1988 (ZMA 20590 x 306075), in volcanic debris of anchiha- Amph. 108.652). line cave, washings of gravel, conductivity 4 2 2 paratypes. Stn. 88-596 B: boulevard of 50.5mS/cm; 13 Nov. 1988 (ZMA Amph. Punta del Hidalgo (UTM coord. CS 108.661). 36987 x 316120), in gravel of rockpool in medio- littoral zone, 0.5 m under sediment surface, BR pump; 12 Dec. 1988 (ZMA Amph. 108.653). Description 3 2 2 paratypes. Stn. 88-616 A: Punta de Teno, Body length up to 1.8 mm. Body (Fig. 1) elon­ small bay (UTM coord. CS 31175 x 313630), gate; body somites about as long as high, bearing 0.4 m under sediment surface, low-tide mark, BR few setules. 'Ocular lobe' present, triangular. pump; 29 Dec. 1988 (ZMA Amph. 108.654). First antenna (Fig. 3d) with 3 stout peduncle 1 S, 1 2 paratypes. Stn. 88-616 B: same locality segments and 4-segmented flagellum: aesthetascs as previous station, a few metres higher on the on 3 last flagellar segments and also a ribbed beach in coarse volcanic debris, 0.5 m under sedi­ small aesthetasc-like structure on distal margin of ment surface, BR pump; 29 Dec. 1988 (ZMA terminal flagellum segment. Amph. 108.655). Second antenna (Fig. 3i) with 5-segmented peduncle (Fig. 1) and 5-segmented flagellum; ter­ 1 2 paratype. Stn. 88-617; Punta de Teno, north minal segment with aesthetasc-like structure on beach (UTM coord. CS 31175 x 313655), strong distal margin. swell, 0.2 m under sediment surface in coarse Right mandible (Fig. 4i) with 7 teeth on pars gravel between boulders, BR pump; 29 Dec. 1988 incisiva; lacinia mobilis with several small teeth; (ZMA Amph. 108.656). pars molaris sharply pointed. Left mandible 1

Fig. 1. Ingolfiella canariensis n. sp., male holotype, 1.8 mm. Playa de los Troches, Tenerife.

than ischium and merus combined; merus with 1 seta anterodistally; carpus relatively elongate, palm bearing 4 spines: 1 long, 3 short and 3 Y-shaped spinules with a sensory hair coming out of the core; dactylus with 4 sharply pointed teeth. Second gnathopod (Fig. 2a, b) carpus more oval than of first gnathopod; palm carrying 3 spines and 3 Y-shaped spinules; margin with 8 serrations (inset a' of Fig. 2a drawn after a SEM photo). In the male a heavy forked spine near the palmar margin is present, as well as a foliaceous structure hanging from the posterior margin of the merus; dactylus with four sharp teeth. Third pereopod (Fig. 2c) with trifid unguis (Fig. 4g). Fourth pereopod similar to third. Oostegites small and suboval (Fig. 4e), some­ times with 1 seta. Fifth pereopod (Fig. 3a) robust, short, with broad basis; heavy spines on merus, carpus and propodus; dactylus separated from unguis by a very faint demarcation line; unguis bifid. Oostegite small, suboval. Coxal gill present. Fig. 2. Ingolfiella canariensis n. sp., paratypes. a, second gnathopod ?, a' inset drawn after a SEM photo; b, second Sixth pereopod (Fig. 3b) more slender and gnathopod cf; c, third pereopod $ ; d, first gnathopod cf; longer than fifth; dactylus faintly separated from e, first gnathopod cf; f, telson, cf; g, first uropod cf • unguis; unguis bifid. 296

/ V

Fig. 5. Ingolfiella similis Ronde-Broekhuizen & Stock, 1987, female, a, second gnathopod; b, first gnathopod; c, tip of second antenna; d, tip of first antenna.

Seventh pereopod (Fig. 3c) more slender and longer than sixth; merus with 2 long spines on distoposterior corner; carpus with 6 distal spines and 1 spoon-shaped denticulate element; dac­ tylus faintly separated from unguis; unguis bifid. Fig. 3. Ingolfiella canariensis n. sp., paratypes. a, fifth pereo­ Pleopods (Fig. 3f, g, g) triangular. First pod d; b, sixth pereopod cf; c, seventh pereopod d; d, first antenna d', e, second uropod £ ; f-h, pleopods 1-3 !j>; pleopod in male with two distal spinules. i, second antenna d- First uropod (Fig. 2g, 4c) biramous; peduncle with 2 setae; exopodite pointed, bearing 1 seta; endopodite longer and wider than exopodite; medial surface with 7 long setae; apex with 4 to 5 spiniform processes. Second uropod (Fig. 3e, 4b) peduncle carrying 3 rows of spinules, bifid at tips; rami subequal, curved, pointed; exopodite with 3 setae, endo­ podite with 2 setae. Third uropod (Fig. 2f) small, 2-segmented; first segment with 2 setae; second segment square at apex with 1 long distal seta. Telson (Fig. 2f) with 2 setae.

Etymology Named after the Canary Islands.

Fig. 4. Ingolfiella canariensis n. sp., paratypes. a, first maxilla Remarks ? ; b, second uropod $ ; c, first uropod $ ; d, maxilliped $ ; e, coxa of third pereopod with gill and oostegite; f, second maxilla d', g, dactylus of fourth pereopod d; h, left The male differs from the female in the following mandibular body d\ i, right mandibular body d', j, first respects: maxilla d- - On the carpus of the second gnathopod a 297

broad, stubby, forked spine is present near lated element on the carpus of the seventh pereo­ the palmar margin. pod (vs. absent); 7 serrations on the palmar mar­ - On the merus of the second gnathopod a gin of second gnathopod carpus (vs. 8); first transparent foliaceous process is present in gnathopod without sensorial setae near edge of the same position of a normal seta in the palmar margin; oval third pleopod (vs. triangu­ female (this process has a different structure lar); bifid dactylus of third and fourth pereopods than the 'sackartigen Gebilde' on the carpus (vs. trifid). of/, petkovskii S. Karaman, 1957 where it is /. quadridentata differs from /. canariensis in clearly made up of two segments - rather like having: 1 palmar spine on carpus of first gnatho­ an aesthetasc - or than the spiniform and pod (vs. 4); unguis of third and fourth pereopods ciliated process as in, for instance, /. putealis simple with inner row of spinules (vs. trifid Stock, 1977). without spinules); carpus with 2 modified spines - There are 2 spinules at the distal and of the posterodistally (vs. absent). Of /. quadridentata first pleopod (naked in cO. the mouthparts are not described. - Peduncle of the second uropod with a sub- basal, hook-shaped spine. /. xarifae Ruffo, 1966, female, from shallow reefs in the Maldives, has a striking resemblance to Only those species possessing a 4-toothed /. canariensis especially with respect to gnathopod dactylus on the first and second gnathopod, and setation and spination, as well as other features of having dissimilar claws on the third to seventh pereopodal dactyli. However, the dactylus of the pereopods, are considered. These conditions first gnathopod has 3 teeth on the inner margin, seem to be derived because they stand out against while the second gnathopod has 4. This character the 'normal' gammaridean condition (Dahl, needs to be checked in the two existing specimens, 1977). Most other species in Ingolfiella have 0-3 and the mouthparts are to be described. teeth on the dactyli of the gnathopods and claws of similar form in the third to seventh pereopods. /. kapuri Coineau & Chandrasekhara, 1972 Ingolfiella similis (Andaman and Nicobar Islands, mediolittoral), /. longipes Stock, Sket & Iliffe, 1987 (Bermuda, Ref.: Ronde-Broekhuizen & Stock, 1987, anchihaline), and /. quadridentata Stock, 1979 p. 441-450 (Curacao, infralittoral) share these character states with /. canadensis. The species are known Material through females only, however. FUERTEVENTURA (Canary Islands). /. kapuri differs from /. canariensis in having: 3 1 . Station 87-119: end of Barranco de los palmar spines on the carpus of first gnathopod Molinos (UTM coord. ES 59175 x 315760) (versus 4), 7 serrations on palmar margin of car­ method Karaman-Chappuis, conductivity pus of second gnathopod (vs. 8); heavier unguicu- 12.6 mS/cm; 6 May 1987 (ZMA Amph. 108.690). lar spines pereopod dactyli; 1 spine on exopodite of first uropod (vs. 0 in $, 1 in d); seventh 1 d- Stn. 87-77: Las Playitas (UTM coord. pereopod with 1 long spine on posterodistal cor­ ES 59990 x 312315) well about 500 m from the ner of merus (vs. 2). The mouthparts of/, kapuri sea, cond. 10.7mS/cm; 6 May 1987 (ZMA are not described. Amph. 108.689). /. longipes differs from /. canariensis in having: 3 spines and 2 setae on outer lobe of first maxilla Description of female (vs. 6); 1 distal spine on terminal segment of Body similar to that of male; all appendages as in maxilliped palp (vs. 2); a spoon-shaped denticu­ male (see Ronde-Broekhuizen & Stock, 1987) 298 except for those described below: second gnatho- lations (Hart, Manning & Iliffe, 1985), can be pod (Fig. 5a) without reversed element on lower termed a 'continuous crevicular corridor hypothe­ margin of carpus; first pleopod without spines; sis'. He considers this hypothesis 'has considera­ second uropod without subbasal hook-shaped ble merit when explaining distributions between spine; oostegites oval. different islands within a particular archipelago, such as the Canary islands, but that it is implausi­ ble merit when explaining distributions between Zoogeographic remarks remote island systems surrounded by ocean floor covered by a significant depth of pelagic sedi­ Geological data from the Canary Islands suggest ments'. As to the affinities with such distantly that at least the western islands are separate vol­ located species as /. kapuri (Andaman and Nico- canic edifices with an overall irregular and com­ bar Island), /. longipes (Bermuda) and /. quadri- plex decrease in age from east to west. They dentata (Curacao); they all live in coastal habitats evolved independently for at least the last 20 in the former realm of the circumtropical Tethys million years (Ronde-Broekhuizen, unpublished). Sea (early Cretaceous, Smith etal, 1981) and Hierro, the youngest and westernmost island has their phenetic similarities may stem from the time a subaerial existence of 0.75 My, while Tenerife that gene flow was continuous. surfaced 15.7 My ago (Pitman & Talwani, 1972). They are separated by 3000 m deep water. /. canadensis is found in marine groundwater of Acknowledgements both islands.This fact suggests three possible explanations: In the course of the research project on the origin - /. canadensis spread, actively or passively of insular groundwater biotas in the Atlantic (ships, not vicariance), from Tenerife to Hierro; financial support has come from - it lives not only in the mediolittoral but also on NATO Collaborative Research Grants Pro­ the ocean floor and, when seabottom eruptions gramme, contract SA. 5-2-05 (RG.0011/88); occur, follows the rise of a volcanic slope until ERASMUS programme, contract ICP 88-0079 it reaches shallow water. NL; NWO, Den Haag. The hospitality of the staff - or /. canadensis invaded both islands only very of the Departamento de Zoologia, Universidad de recently, having evolved into a mediolittoral La Laguna (Director: Prof. Dr. M. Ibaflez Genis) stygobiont from a common marine benthic is gratefully acknowledged. ancestor. Also we wish to thank Drs. H.P. Wagner for staining part of the material, D. Platvoet for tak­ The second explanation requires fewer specula­ ing SEM photos and Prof. Dr. J.H. Stock for tive steps and is favored. In this case it should be critically reading the manuscript and for provid­ possible to find /. canadensis also on the other ing material from Fuerteventura. western Canary Islands, i.e. La Palma, Gomera and Gran Canaria and in the seabed between those islands. For that matter; ingolfiellids have earlier been found in oceanfloor debris at great References depths (/. abyssi Hansen, 1903; /. atlantisi Mills, Bou, CL, 1975. Les methodes de recolte dans les eaux 1967) and in dredge samples from bottoms of souterraines interstitielles. Annls. Speleol. 29: 611-619. shallower water (/. britannica Spooner, 1960; Boxshall, G. A., 1989. Colonization of inland marine caves I.fuscina Dojiri & Sieg, 1987). by mysophrioid copepods. J. Zool., Lond. 219: 521-526. Coineau, N. & Rao G. Chandrasekhara, 1972. Isopodes et Boxshall (1989) remarks that the idea of a mar­ Amphipodes des sables intertideaux des iles Andaman et ine crevicular fauna that is interconnected from Nicobar (Golfe du Bengale). Vie Milieu (A) 22 (1A): one island to the other through deep water popu­ 65-100. 299

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