~oologicalJournal of the finnean Sock& (1996), 116: 149-1 66. With 2 figures New records of marine Tardigrada in the Mediterranean Sea SANTIAGO VILLORA-MORENO Laboratorio de Biologi Marina, Departamento de Biologii Animal, Uniuersitat de Vakncia, Auda. Dr, Moliner, 50, 461OO-Buls;assot, Valencia, Spain* AND SUSANNA DE ZIO GRIMALDI Istituto di <oologia ed Anatomia Comparatu., Uniuersitci degli Studi di Bad, Via E. Orabona 4, 70125 Bari, ItaQ Meiofauna from Chafarinas archipelago (western Mediterranean, N. Africa) were studied, and marine Tardigrada were identified to species level. A total of 15 species, belonging to families Stygarctidae, Halechiniscidae (SubfamiliesHalechiniscinae, Florarctinae, Styraconyxinaeand Tanarctinae), Batillipedidae, and Echiniscoididae, were found for the fwst time from the Alboran Sea and southwestern Mediterranean area. Four species and one subspecies are new records for the Mediterranean Sea: Floractus acer, ?I mpm, F. stellatus, Actinarctus dotyphonu and Echiniscoides sigirmundi hispanimris. The male of Floractus stellatus, previously unknown to science, is described. Many species have been recorded for the first time in different sedimentary environments, such as midlittoral pools, very shallow gravels, pools in a mixed rocky environment or between Posidonia oceanica meadows, ‘Amphioxus’ gravels, and coastal detritic. 01996 The Linnean Society of London ADDITIONAL KEY WORDS: -Stygarctidae - Halechiniscidae - Batillipedidae - Echiniscoididae geographical distribution - ecology - Alboran Sea. CONTENTS Introduction . . . . . . Material andmethods . . . Family Stygarctidae . . Subfamily Stygarcinae . Family Halechiniscidae . Subfamily Halechiniscinae Subfamily Florarctinae . Subfamily Styraconyxinae Subfamily Tanarctinae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 150 150 150 152 153 153 155 158 160 *Present address: Observatoire Ocetanologique de Banyuls, CNRS URA 117, F-66650 Banyuls Sur-Mer France 0024-4082/96/010149 + 18 $18.00/0 149 81996 The Linnean Society of London I30 S. VILLORX-SIORENO .LID S. DE ZIO GlUI\.IAI,DI Faniilj- Batillipedidae . . . . . Family Erhiniscoididae . . . . Concluduig commmts . . . . . . ‘Taxonomic arid systematic remarks Ecological records . . . . . BiogeoLgraph>- . . . . . . . Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kckrences . . . . . . . . . . . . . . . . . . . . . . . . . . . 161 162 162 162 163 163 164 164 IXTRODL‘CTION As extensive exploration of the meiobenthos continues, information on abundance and diversity of marine Tardigrada will increase giving rise to greater interest in the ,group. Marine Tardigrada are a common, occasionally abundant, component of the meiofauna found in all marine sediments. From the monograph on the phylum Tardigrada (Marcus, 1936),where only six marine species were recorded, to the review of Renaud-Mornant (1982a), with 22 genera and a total of 56 species, our knowledge of marine Tardigrada has drastically increased. Interest in marine Tardigrada worldwide during the last decade has nearly doubled the number of genera (-10) and more than doubted the number of known species (more than 125). Marine ‘I’ardi<gradarecords from the eastern and southwestern Mediterranean basin are scarce because this group has been studied mainly from the southern Italian coasts, where nearly 50 species are known (De Zio Grimaldi, unpublished data). Located in the southwestern Mediterranean basin, the Alboran Sea is of interest because of its geographical position between both Atlantic and Mediterranean waters. In order to expand our knowledge of the marine tardigrade fauna of the Mediterranean Sea, we investigated Tardigrada from the Chafarinas Islands in the southeastern Alboran Sea (north of Africa). A total of 15 species have been identified, five of which are new records for the Mediterranean Sea. hiATERL4L tLXD hlETHODS hfeiofauna from the Chafarinas archipelago (Alboran Sea, western Mediterranean) were collected during July-August 1991 during a benthos survey (Fig. 1). Samples for both qualitative and quantitative analyses were taken by means of a box corer (1 1) and a hand operated coring device (10 cm2 area). Samples were treated with 7.5% MgCl, as a narcotic agent and fixed with 5% buffered formaldehyde. ‘I’ardigrada were removed from the extracted meiofauna and identified to species level. All specimens were mounted in glycerine, using a coverslip with supports to prevent body deformations. General environmental and biological (meiofauna) characteristics of the area were described by Villora-Moreno (1993). Details on the collection localities and other data are provided for each species. Family Stygarctidae Schulz, 1951 (emend. McKirdy, Schmidt & McGinty-Bayly, 1976) Arthrotardigrada with five pairs of cephalic appendages; anterior clavae either Figure 1. Map of the Chafarinas Archipelago showing collection sites. See test (zZfatprialexamined subsections) for further explanation 152 S \~IIdI,ORA-h1ORENO AWD S. DE ZIO GRIMALDI elongate or modified as hemispherical structures. Cuticle forming unpaired dorsal thickenings or plates. Feet non-digitate; in adults, either four claws per foot with medial pair bearing a long filament or accessory spine, three claws on each leg each with an accessory spine, or four claws on legs 1-3, and two larger claws on leg 4, all with a short dorsal accessory spine. Claws often attached to foot by a basal membrane. Subfamily Stygarcinae Schulz, 1951 (emend. De Zio Grimaldi et al., 1992) Stygarctidae with a complete set of cephalic appendages. Primary clavae either elongated or hemispherical. Dorsal cuticle forming five dorsal thickenings: a cephalic plate, three body plates, and a caudal plate; ventral plates sometimes present, Bulb shaped papilla on the fourth leg. Genus SpgarctuJ Schulz, 1951 (emend. Pollock, 1989) Stygarctidae with dorsal cuticle thickened to form three unpaired body plates between an undkided cephalic plate and a caudal plate; legs with four claws each, the central pair Lvith long filamentous appendages: anterior clavae elongate. Spgarrtus bra&us Schulz, 1951 Sbgarctus b r ~ d y p uSchulz, ~ 1951; Renaud-Mornant & Anselme-Moizan, 1969; Grimaldi De ZLo et ai., l980b, 1983; Pollock, 1989. hfatprial examined. Rey Francisco Island: sublittoral sand and gravel, -1 m deep, from station RE03c 1 (1 female). Dzaynoszs. S!jgarctus with a single pair of acute processes at posterior rim of cephalic plate; with large bitid spikes from midposterior margin of body plate 11; large recurved anterior clavae, long caudal spikes, body plates 1-111 with flared lateral edges. Ecology. Intertidal interstitial, from 10 to 50cm deep in medium to coarse sand (Renaud-Debyser, 1956; hlcGinty & Higgins, 1968; Grimaldi De Zio et al., 1980b; Pollock. 1989). In the Charfainas archipelago, Spgarctus bradypus was identified from shallow sublittoral coarse sand and gravel. According to the literature, this species is usually interstitial and intertidal, but may be occasionally found in the sublittoral. Chgraphical dzrtribution. Based on reports by Lindgren (197 1) and Pollock (1989), Sogurctus brat$pus is a boreal and circumglobal species because it has been reported from the Mediterranean Sea and the Pacific, Atlantic and Indian Oceans (all records in the northern hemisphere). However, the data are insufficient to conclude that this is the actual geographical distribution not only of this species, but also of the majority of marine Tardigrada. MEDITERRANEAN TARDIGRADA 153 Family Haleschiniscidae Thulin, 1928 (emend. Grimaldi De Zio et al., 1990) No strongly sclerotized dorsal segmental plates. Secondary clavae sometimes clubshaped or dome-shaped or indistinguishable. Adult legs with four digits bearing claws (simple, with accessory hooks; or with distal section of uncus shaped like a bird’s beak, ‘aviculaire’ s m u Renaud-Mornant, 1989).Peduncles sometimes present in the digits. Cirrus E often with a basal accordion-shaped articulation. Subfamily Halechiniscinae (Thulin, 1928) Usually with complete set of cephalic appendages. Cylindrical tibia followed by a small tarsus. Claws either simple or with a dorsal spur. Peduncles absent. Sense organ on the fourth pair of legs short and non-branched. Genus Haleschinkcus Richters, 1908 (emend. Grimaldi De Zio et al., 1990). Flat head extending into lateral lobes. Claws either simple or with distal dorsal spur, calcar always absent. Primary clava may be longer than the lateral cirrus. Halechinkcus greueni Renaud-Mornant & Deroux, 1976 HalechiniscusgreueniRenaud-Mornant & Deroux, 1976; Kristensen, 1981;D’Addabbo Gallo et al., 1987; Grimaldi De Zio & D’Addabbo Gallo, 1987; D’Addabbo Gallo et al., 1989; Grimaldi De Zio et al., 1990. Material examined. Congreso Island: sublittoral mixed environments from stations CGOl (2 females), CG03 (10 females, 6 males and 2 preadults), CG04c1 (1 male), CG04c2 (2 females, 1 male and 1 preadult); ‘Amphioxus’gravel from stations F06 (24 males and females), F07 (5 males and females). Diagnostic characteristics are according to the species revision of Grimaldi De Zio et al. (1990), including all additional characters in the emended diagnosis of the species. Ecology. The original description was made with material from a sink in the Biological Station of Roscoff (Renaud-Mornant & Deroux, 1976). Kristensen (1981) recorded the species in the natural environment, from a midlittoral rock pool at St. Pol-deLeon (Atlantic Ocean, France). Atlantic records are intertidal however, all new records of the species in the Mediterranean sea are sublittoral, mainly from biodetritic sediment (Italian coasts). In the Chafarinas archipelago, the species were identified in sublittoral coarse mineral sediment, mainly basalt, and in biodetritic sediment. HalechinzScus greueni was not observed either in midlittoral rock pools or in the midlittoral sediment patches. The species is identified in high numbers from both organic and mineral sublittoral sediments with well-developed interstitial spaces. Geographical distribution. Mediterranean and Atlantic-Oriental distribution: English Channel, Roscoff, (Renaud-Mornant & Deroux, 1976; Kristensen, 1981); Mediterranean Sea, Ionian Sea (D’Addabbo Gallo et al., 1987; Grimaldi De Zio & D’Addabbo Gallo, 1987), Tyrrhenian Sea (D’Addabbo Gallo et al., 1989), Gulf of Valencia, Spain, (Villora-Moreno, unpublished materials). This work is the first record of the species in the Alboran Sea and southwestern Mediterranean area. S \’ILLORA-,ZlOKENO /UYD S. DE ZIO GRIMALDI 1% Halechzniwus perfpctus Schulz, 1955 Hakchiniscu.~pefer&iisSchulz, 1955; Swedmark, 1956; Fize, 1963; Renaud-Mornant, 1967, 197 1 a, b, 197613; Salvat & Renaud-Mornant, 1969; D’Addabbo Gallo et al., 1978; Grimaldi De Zio et al., 1980a, b, 1983; Morgan & O’Reilly, 1988. Halechinixus guiteli Grell, 1936 (in Ramazzotti & Maucci, 1983) i24atrrial mamined. Congreso Island: sublittoral mixed environment from station CG04c2 (1 preadult). Remarks. The identified animal corresponds to the previous descriptions in all essential characters. ‘I’hisspecies, like Halerhiniscus greveni and H. guiteli, has no lateral expansions. General morphology of the head and size of primary clava are the more obvious characters that separate this group of sister species. Ecology. Halechinisrus pefectus is known from both the midlittoral and sublittoral sediments. In the midlittoral it has been recorded in medium to coarse sediment (Fize, 1963; D’Addabbo Gallo et al., 1978; Grimaldi De Zio et al., 1980a, b, 1983) and coralline sand (Rcnaud-Mornant, 1967). Sublittoral records correspond to the original description (Schulz, 1955) in ‘Amphioxus’ gravel (also found in this environment by Sw:edmark, 1956).Other sublittoral records are in coarse biodctritic sediment, to 173m deep (Renauld-Mornant, 1971a, b). In the Charafinas archipelago this species was found in medium biodetritic sediment from a sublittoral mixed en\ironnient with rock. This species is frequent in medium-coarse sediment, both biogenous and terrigenous. There are no records from deep sea sites in the literature. Halechzniscuspefertzrs was described by Schulz ( 1955) from two distinct populations with one (Banyuls-sur-mer)and three (Naples)specimens, respectively. The species is often present, with very few specimens, from both midlittoral and sublittoral sediments. This very low abundance and the absence of large samples have prevented a proper evaluation of intrapopulation variability, making redescription of the species difficult. ~~ Geographical di.Ptribution. Atlantic north-oriental, Mediterranean and South Pacific: Scottish coasts (Morgan & O’Reilly, 1988),Brest, France, (Renaud-Mornant, 197 la, b); Gulf of Aigues Mortes (Fize, 1963), Ionian Sea (D’Addabbo Gallo et al., 1978; Grimaldi De Zio et al., 1980a, b, 1983), Coral Sea, New Caledonia (RenaudMornant, 1967),and French Polynesia (Rcnaud-hfornant, 197613; Salvat & RenaudMornant, 1969). l’his work is the first record of the spccics in the Alboran Sea and southwestern Mediterranean Area. Ha.lechiniscus remand Schulz, 1955 HalechinZrcus reinanei Schulz, 1955; Rcnaud-Debyser, 1959, 1963; Fize, 1963; De Zio, 1964; Swedmark & Teissier, 1967; De Zio & Grimaldi, 1966; McGinty & Higgins, 1968; Rao & Ganapati, 1968; McGinty, 1969; Renaud-Mornant, 197 1b; D’Adcfabbo Gallo et al., 1978; Grimaldi De Zio et al., 1980a, b, 1983; Renaud-Mornant 8r Gourbault, 1984; Renaud-Mornant, 1981, 1984. :tfatm’al examined. Isabel 11 Island: shallow sublittoral sediment between rocky blocks at station IS06c2 (1 male). Dzty.noszs. Secondary clavae vestigial. Sensorial setae on legs 1 to 4. Papilla on fourth MEDITERRANEAN TARDIGRADA 155 leg on a strong base. Cuticular lateral expansions between each leg and after the head. Two subspecies are known (Renaud-Mornant, 1984). Remarks. The one specimen identified corresponds with Halechiniscus remanei remanei; however, the cuticular lateral expansions and tail are longer than in Schulz’s description. Ecology. Records of the species are mainly from midlittoral medium-fine sand (Renaud-Debyser, 1959; McGinty & Higgins, 1968; D’Addabbo Gallo et al., 1978; Grimaldi De Zio et al., 1980a, b) or shallow sublittoral (Fize, 1963; this work). The original description was made from coarse sublittoral sand (Schulz, 1955) and new records have been established in the same environment (Swedmark & Teissier, 1967). Only one record is known from -1 30 m deep (Renaud-Mornant, 197 lb). The subspecies antillensis is only known from midlittoral coralline sand in Antilles and the coast of Florida (Renaud-Mornant, 1984). The species is midlittoral and sublittoral, inhabiting the interstitial system in fine, medium and coarse sands, both siliceous and biogenous, but it is not an eurytopic species. Geographical distribution. Recorded from the Atlantic, Indian, and Pacific Oceans and the Mediterranean Sea, the species shows a circumglobal distribution. It is not possible to designate a cosmopolitan distribution (Renaud-Mornant, 197lb, 1984) because no records are available from the Arctic and Antarctic Oceans. This work is the first record of the species in the Alboran Sea and southwestern Mediterranean area. Halechiniscus sp. nov. Material examined. Isabel I1 Island: sublittoral mixed environment near the seagrass community of Posidonia oceanica meadows, from station IS 1 1c 1 (1 female). Diagnosis. Halechiniscus with cephalic region divided into two lobes: a ventral one bearing the external ventral cirri and the internal dorsal ones, and a dorsal lobe with its central part protruding dorsally to support the strong median cirrus and with the lateral sides expanding with large cirrophori bearing lateral cirri and primary clavae. Ventral mouth opening in the middle of an elevated square-like area. Kidney-shaped sensory areas surrounding the mouth. Three large conical lateral expansions above the first and second legs and between the third and fourth ones. Smaller lateral expansions alternating with the larger ones. Legs and claws as usual in Halechiniscus. Definitive description of this new species is postponed because only a single specimen (a mature female) has been found. Subfamily Florarctinae Renaud-Mornant, 1982 emend. Kristensen, 1984). Halechiniscidae with cuticular aliform expansions. Complete cephalic sense organs with secondary clavae transformed to dome-shaped papillae or H-shaped flat sacs. Four toes with claws present in adults; external digit with a hook-shaped pedunculus; uncus with a calcar externum. Females with two seminal receptacles, each consisting of a spheroid vesicle and S-shaped genital duct. 156 S. \’IIZORA-MORENO AND S. DE ZIO GRIMALDI Genus Florarctus Delamare-Deboutteville & Renaud-Mornant, 1965 Three to five large lateral and caudal lobate aliform expansions, more or less completely covering the legs. Caudal ala may be present with spines. Procuticular expansion (caestus)frequently present in the ala. External digits short with distal part of claws shaped like a bird’s beak (also called ‘aviculaird’).Internal digits with simple claws and accessory spines. Florarctus actr Renaud-Mornant, 1989 Material examined. Congreso Island: sublittoral mixed environment between rocks, from station CG04c2 (1 preadult). Remarks. General features as in original description of Renaud-Mornant (1989). Caestus evidently pointed in the distal part. Ecology. Identified by Renaud-Mornant (1989) from the sublittoral, -25 m deep, in coarse sand (Dentalium sand), and from the intertidal. In the Chafarinas archipelago, this species was observed in a medium sand pool, -3 m deep, located between rocky blocks, near the community of photophilic algae (PCrb, 1967).As in other species of the genus, it is found only as interstitial meiofauna. Geographical distribution. Atlantic north-oriental: Bay of Morlaix and Roscoff in France (Renaud-Mornant, 1989). No data are available after the original description. This is the first record for the Mediterranean Sea. The species shows a Mediterranean and Atlantic north-oriental distribution. Florarctus mpr Renaud-Mornant, 1989 hlateriul examined. Rev Francisco Island: circalittoral station RE 12 (2 specimens). Remarks. According to the original description of Renaud-Mornant (1989). As in other species of the genus Florarctus, the caestus is the more useful character for species identification. Ecolo~.Lagoon coralline sand -3 m deep (Renaud-Mornant, 1989). Two specimens were identified in the Chafarinas archipelago at -27 m deep in the community of the Coastal Detritic (Perks, 1967), in the interstitial system of coarse sand and gravels. Sediment was 90% biogenous, similar to the original description. Geographical distribution. New Caledonia in the Coral Sea and French Polynesia in the Pacific Ocean (Renaud-h.lornant, 1989). No data after the original description. This is the first record for the Mediterranean Sea. The species shows a PacificMediterranean distribution. Florarctus cinctus Renaud-Mornant, 1976 Florarctus cinctus Renaud-Mornant, 1976a, 1979; D’Addabbo GaUo et al., 1987; Grimaldi De Zio & D’Addabbo Gallo, 1987; Grimaldi De Zio et al., 1980a. Material examined. Isabel I1 Island: sublittoral mixed environment near meadows of the seagrass Poszdonia oceanica, at -4- m deep, from station IS 1 1c2 (3 specimens). Rey Francisco Island: sublittoral sand and gravel, -1 m deep, from station RE03c1 (1 specimen). Remarks. All external characters as in the original description (Renaud-Mornant, 1989).Florarctus cinctus is closely related to F. mper and F. acer by the presence of dorsal MEDITERRANEAN TARDIGRADA 157 cuticular structure and caestus. The former species differs from the others in the morphology of the cuticula, sculptured with much more and smaller dots. Ecology. Sublittoral coarse sand, -1 1.5 m deep, and coralline sand (Renaud-Mornant, 1976a). Also in biodetritic coralligenous sediment, between -20 and -200 m deep (Grimaldi De Zio & D’Addabbo Gallo, 1987). In the Chafarinas archipelago Florarctus cinctus has been identified in the interstitial system: in mineral (basalt) coarse sand from a pool, -4.m deep, near the Posidonia oceanica meadows, and from -1 m deep in rocky environment with similar granulometrical characteristics. This sublittoral species is only found when a true interstitial system is present. Geogaphical distribution. Mediterranean: Gulf of Naples and Ionian Sea (RenaudMornant, 1976a; Grimaldi De Zio & D’Addabbo Gallo, 1987).Atlantic NE (Roscoff) and Madagascar in the SW Indian Ocean (Renaud-Mornant, 1976a, 1979, 1986). This work is the first record of the species in the Alboran Sea and southwestern Mediterranean area. Florarctus hulingsi Renaud-Mornant, 1976 Florarctus hulingsi Renaud-Mornant, 1976a, 1986, 1987, 1989; D’Addabbo Gallo el al., 1978, 1987, 1989; Renaud-Mornant & Gourbault, 1984; Grimaldi De Zio & D’Addabbo Gallo, 1987; Grimaldi De Zio et aL, 1980a, 1983. Material examined. Isabel I1 Island: midlittoral pool from stations ISlOcl and c2 (2 females, 1 male and 2 preadults). Remarks. Florarctus hulingsi and F. stellatus are the only two species of the genus Florarctus, without caestus in the alae. F. hulingsi may be separated from F. sbllatus by its smooth cuticle. Observations on populations of both species in the Chafarinas Islands add other distinctive characters such as claw morphology. F. stellatus claws are sharp and sickle-shaped, differing from the stronger and square-shaped ones of F. hulingsi. Ecology. Medium and coarse midlittoral sand, to -20cm deep in the sediment (Renaud-Mornant, 1976a; D’Addabbo Gallo et al., 1978; Grimaldi De Zio et al., 1980a). Sublittoral biodetritic coralligenous sediment between -20 and -80 m deep (D’Addabbo Gallo et al., 1987, 1989; Grimaldi De Zio & D’Addabbo Gallo, 1987). In the Chafarinas Islands Florarctus hulingsi was found in mineral and biogenous sediment from a midlittoral pool. Geogfaphical distribution. Mediterranean and Indian Ocean (Madagascar) (RenaudMornant, 1986). This work is the first record of the species in the Alboran Sea. Florarctus stellatus Renaud-Mornant, 1989 Material examined. Isabel I1 Island: sublittoral mixed environment near Posidonia oceanica meadows, -4. m deep, from station IS 1 1c 1 and c2 (2 1 specimens). Congreso Island ‘Amphioxus’ sand from station F06 (8 males, 7 females). Rey Francisco Island: sublittoral channel between Posidonia oceanica meadows, -5 m deep, from station RE05 (1 specimen). 7jpe material. Allotype male from Isabel I1 Island, station IS11 (slide CHAF-782). Paratype female from the same station (slide CHAF-790). Dzagnosis and remarks. Mediterranean specimens, both males and females, are smaller 158 S VILLORA-MORENO M I D S. DE 210 GRIMA1,DI than specimens (only females) of the original description (Polynesia, RenaudMornant, 1989). Males were previously unknown so a description is reported here. Allotype male, 130 pm long (Fig. 2D). The primary clavae are a little longer in the male than in the female (50--70pm). However, there is not a strong sexual dimorphism as in other species of Florarctus. The secondary clavae have H-shape and surround the mouth cone (Fig. 2A). Males and females have cephalic vesicles with symbiotic bacteria (Fig. 2B). A large vesicle U-shaped is located ventrally, around the mouth opening and a smaller one is frontal, placed between external cirri. The dorsal cuticle is sculptured as in the female description (Renaud-Mornant, 1989)(Fig. 2C). The digiti externi are shorter than the interni. An accessory spine is present in the claws of the digiti interni (Fig. 2E). Testis is dorsal and medium-sized. Large lateral seminal vesicles are located at both sides of gonopore. The germ cells in the caudal region of testis and in the seminal vesicles are shaped like wheat grains. The male gonopore is slightly papillar, with ovoid opening and a refringent, triangularshaped, subcuticular structure between gonopore and anus (Fig. 2F,G). Ecology. Sublittoral coralline sand, -32 m deep (Renaud-Mornant, 1989). In the Chafarinas archipelago it has bccn observed in mineral (basaltic) and biogenous sediment (80% of carbonates), both with similar granulometric composition (medium to coarse sand). Geyaphical distribution. Pre\.iously known only from the original description: French Polynesia in the Pacific Ocean. This is the first record for the Mediterranean Sea. The species shows a South Pacific and Mediterranean distribution. Subfamily Styraconyxinae Kristensen & Renaud-Mornant, 1983 (emend. D’Addabbo Gallo et ul., 1992) Haleehiniscidae with peduncles present either on all digits or only on the first and fourth ones, or absent. Heart-shaped proximal pads may be present at the base of second and third digits. Crescent-shaped claws with one, two or no accessory points, depending on the genus. Complete set of cephalic cirri always present. Secondary clavae sometimes absent. Primary clavae and fourth papillae similar. Cuticular plates absent. Genus S@xonyx Thulin, 1942 External digits always supported by pedunclcs; internal digits with proximal pads, peduncles present or absent. Claws, with three exposed points (hooks), may be retracted within claw sheath. Configuration of primary clavae and lateral cirri variable; secondary clavae dome-shaped papillae or indistinct flat sacs. Spines present on legs 1-3, clavoid papillae on leg 4. SQraconyx cruticulus (Pollock, 1983) Buthvechiniscus cruticulus Pollock, 1983 S$vrucon_yxcraticulus: Kristensen & Higgins, 1984; D’Addabbo Gallo et ul., 1987, 1989; Grimaldi De Zio & D’Addabbo Gallo, 1987 iLfaterial examined. Isabel I1 Island: midlittoral pool from stations ISlOcl and c2 (2 MEDITERRANEAN TARDIGRADA Figure 2. Florurctus ste1htu.r. AUotype male. A, secondary clavae (arrows) and external cirri; B, cephalic vesicles with symbiontic bacteria; C, cirrus E and sculpture of the dorsal cuticle; D, dorsal view; E, claws of leg 3; F & G, ventral view of male gonopore, showing papilla (arrows) in two optical sections. Scale bar A,C,F,G = 15 pm; B,E = 20 pm; D = 50 pm. 159 160 S . VILLORA-MORENO AND S. DE ZIO GRMALDI females). General features according to the emended diagnosis of Kristensen & Higgins (1984). Ecology. Midlittoral, from barnacles with coralline algae in intertidal coralline sand (Kmtensen & Higgins, 1984). In agreement with previous identifications, in the Chafarinas archipelago S&racoryx craticulus was found inhabiting coarse sand and gravel in a midlittoral pool. Geogaphical distribution. Caribbean Sea (Pollock, 1983), Red Sea and Coral Sea (Kristensen & Higgins, 1984), Mediterranean Sea (D’Addabbo Gallo et al., 1987, 1989; Grimaldi De Zio & D’Addabbo Gallo, 1987). With these records, the species shows a circumglobal distribution (Indian, Pacific, Atlantic and Mediterranean). This work is the first record for the Alboran Sea and the southwestern Mediterranean area. Subfamily Tanarctinae Renaud-Mornant, 1980 Halechiniscidae with lanceolate tibia in the legs, followed by a conical tarsus with opposite vertex forming a thin wrist. Simple claws with external calcar, medial claws sometime with a dorsal accessory spur. Papilla of leg 4 modified as a long caudal appendage, simple or branched. Internal epicuticle supported by pillars. Genus Actinarctus Schulz, 1935 Tanarctinae with raised cuticle all around the body, and supported by extremely long piUars. Claws without external calcar, with or without accessory spur. Actinarch dolyphorus Schulz, 1935 Actinarch d o l y p h Schulz, 1935; Grell, 1936; Swedmark, 1956; Swedmark & Teissier, 1967; Renaud-Mornant, 1971b. Material examined. Congreso Island: ‘Amphioxus’gravels in station F06 (1 specimen), very coarse sand and gravels with biodetritic sediment, -25m deep, in the station E03 (1 specimen). Remarks. General features as in the original description and according to the redescription by Renaud-Mornant (197 1b). The subspecies Actinarctus dolyphorus ocelhtm erected by Renaud-Mornant (197 lb) is based on the presence of refringent, maybe lipid, droplets. We suppose, after numerous observations in other species of the genus Actinarctus and other genera of the family Halechiniscidae, that these lipid dropkts may be present in some species, depending on the physiological state of the animal, so we can not use this element as a diagnostic character. Therefore we consider the specimens with the lipid droplets found in Chafarinas as belonging to the species Actinarctus dolyphom Schulz, 1935, synonymizing with it the subspecies erected by Renaud-Mornant (197 1b). Ecology. Described originally as commensal of Echznocyamus pllfillus. (Echinodermata), this species was found free in the sediment and redescribed by Grell (1936) from ‘Amphioxus’ gravel. In the same community it was identified by Swedmark (1956). It was also found in detritic sediment, -120 to -130m deep (Renaud-Mornant, 197lb). According to previous data, Actinarctus doybhom has been recorded in the MEDITERRANEAN TARDIGRADA 161 Chafarinas Islands from both ‘Amphioxus’ gravel and from very coarse sand and gravel with biodetritic sediment, -25 m deep. Actinarctus do2yphorus is sublittoral, strictly interstitial, inhabiting medium to coarse sand and gravel, both biogenous and mineral. Geographical distribution. Atlantic NE: Helgoland (Schulz, 1935; Grell, 1936), Roscoff and Brest, France (Swedmark, 1956; Swedmark & Teissier, 1967; Renaud-Mornant, 1971b). This is the first record for the Mediterranean Sea, The species shows a Western Atlantic and Mediterranean distribution. Actinarctus physophorus Grimaldi De Zio et al., 1982 Actinarctus physophom Grimaldi De Zio et al., 1982; Grimaldi De Zio & D’Addabbo Gallo, 1987. Material examined. Congreso Island ‘Amphioxus’ gravels in stations F06 and F07 (3 males). Bugnosis. AU taxonomic characters according to the original description (Grimaldi De Zio et al., 1982). Ecology. Sublittoral biodetritic sediment from a cave environment and sublittoral coralligenous sediment between -20 and -200 m deep (Grimaldi De Zio et al., 1982; Grimaldi De Zio & D’Addabbo Gallo, 1987). In the Chafarinas Islands Actinarctus physophorus was only found in the sublittoral, from the community of ‘Amphioxus’ gravel. The species is sublittoral and interstitial. Geographical distribution. Mediterranean Sea: Tyrrhenian Sea and Ionian Sea (Grimaldi De Zio et al., 1982; Grimaldi De Zio & D’Addabbo Gallo, 1987). May be a Mediterranean endemic. This work is the first record for the Alboran Sea and the southwestern Mediterranean area. Family Batillipedidae Richters, 1909 Arthrotardigrada with legs bearing four or six digits with distal adhesive discs; claws absent. Genus Batillipes Richters, 1909 Batillipedidae with six digits ending with a distal disc. Cephalic cirri simple but well developed. Secondary clavae, if present, are domed papillae. Caudal projection present or absent. Batillipeef marcellii Morone de Lucia et al., 1988 Material examined. Chafarinas archipelago. Sublittoral sand, -20 m deep, from the station G17 (7 females, 4 males, 1 preadult). LhgiOs-k.All characters in according with the original description (Morone de Lucia et al., 1988). Ecology. Coralligenous sediment, -1 7 to -43 m deep, Podonia meadows, -5 to -7 m deep (Morone de Lucia et al., 1988). In the Chafarinas archipelago Batillipes marcellii inhabited the interstitid system in large extensions of fine sands. The relationship I62 S. VIIA1,OKt\-hIORENOAND S. DE ZIO GRIbIALDI between sediment Wanulometry and abundance of Batillz$es species is more evident in the sublittoral than in the midlittoral (Villora-Moreno & De Zio Grimaldi, 1993). Geqra~hicaldistribution. Ma); be a Mediterranean endemic. Reported from Tyrrhenian Sea (Morone de Lucia et a/., 1988). This work is the first record for the Alboran Sea and the southwestern Mediterranean area. Family Echiniscoididae Kristensen & Hallas, 1980 Unplated marine Echiniscoidea without toes. Cephalic papillae dome-shaped or indistinct. Other cephalic appendages and leg appendages small (reduced). Cirri A and E similar. 'l'he fourth leg appendage is a papilla, as is the clava. Genus Echini.mides Plate, 1889 (emend. Kristensen BL Hallas, 1980) Echiniscoididae with more than four claws on each leg in adult specimens. Claws are without spurs. l'idal or halophilous forms. EchinZrcoides sigiJmundi hkpaniensis Kristensen & Hallas, 1980 i21akrid examined. Isabel I1 Island: midlittoral pool from station IS 10c2 (1 specimen). Rey Francisco Island: midlittoral pool from station RE06c3 (1 specimen). I~zagnosis.Sculptured cuticle and claws as in original description (Kristensen & Hallas, 1980). E C O ~ Q'The . subspecies was found in the midlittoral barnacles Eliminius modestus and Chtlzamnlus stdlatus, and from stones in the mid-tidal zone of a sandy beach (Kristensen & Hallas, 1980). No data are available after the original description. In the Chafarinac archipelago this species was found free in the coarse sediment from midlittoral pools. Geugrnjdzical dishibutzon Atlantic NE, in Galicia, Spain (Kristensen & Hallas, 1980). This is the first record for the Mediterranean Sea. The subspecies shows a Western Atlantic and Mediterranean distribution. CONCLUDING COhlhlENTS Including the new records described in the present paper, 56 species of marine Tardigrada belonging to 2 1 genera have now been reported from the Mediterranean Sea. This number of genera and species represents about half of the marine Tardigrada known worldwide. The fifteen species of marine Tardigrada found in the Alboran Sea are of taxonomic, systematic, ecological or biogeographical interest. Tavonomic and systaatic remarks 'l'he majority of all species recorded from Chafarinas archipelago belong to the family Halechiniscidae. Three families, Stygarctidae, Batillipedidae and Echiniscoi- MEDITERRANEAN TARDIGRADA 163 didae, each with one species, represent the remainder of the Tardigrada. The genera Halechiniscus and Florarctus have the highest diversity, with four and five species, respectively. Some species of these genera have an additional taxonomic interest. The new species of Halechiniscus, for which we have presented a preliminary diagnosis, shows affinities with the group H. macrocephalus, H. tuleari and H. paratuleari. The high species heterogeneity observed within the genus Halechinkcus, i.e. from H. guiteli to H. macrocephalus, may be diminished after the discovery of this new species because of its intermediate position between the two extreme clusters of Halechinkcus species. Before the present study only two species of the genus Florarctus, F. hulingsi and F. cinctus, were known in the Mediterranean Sea (Renaud-Mornant, 1976a). Florarctus stellatus, F. asper and F. acer are new records for the Mediterranean. Likewise, the male of the former species was previously unknown to science. Florarctus stellatus displays the sexual dimorphism observed in other species of this genus. The body size is smaller in the male, however, the clavae are longer than in the female. Ecological records Different sedimentary environments, ranging from 0 to 60 m depth, were surveyed: mud, sand, muddy sand, ‘Amphioxus’ gravel, heterometric and biogenic sediment in mixed environments (seagrass meadows and littoral ponds), and midlittoral sand or gravel. Tardigrada were not present in all sedimentary environments. Communities of pure mud and sandy mud were devoid of Tardigrada. Batill$es marcellii was the only tardigrade species identified in the community of very fine sands (in agreement with reported preferences of the family Batillipedidae). Fourteen species were identified in the communities of mixed environments. In these communities the high heterogenity of interstitial environments provides numerous microhabitats that support a high biodiversity. Halechiniscus greueni was the most abundant species. ‘Amphioxus’gravel showed a high diversity of meiofauna groups and tardigrade species. Florarctus stellatus and Halechiniscus greueni reached their higher densities in this community. Midlittoral pools and very shallow gravels and sands showed an outstanding richness of tardigrades, with the species Spraconyx craticulus, Spgarctus bra&pus, Florarctus hulingsi, F. cinctus, Halechiniscus remanei and Echiniscoides sigismundi hispaniensis. Many species have been recorded for the first time in different sedimentary environments: Spgarctus bradypus and Florarctus hulingsi in midlittoral pools and very shallow gravels; Halechinixus pdectus, Florarctus acer, F. cinctus and F. stellatus from pools in a mixed rocky environment or between Posidonia oceanica meadows; Halechinkcus greueni, Florarctus stellatus and Actinarctusphysophorus, in ‘Amphioxus’gravels; and finally, Florarctus asper from Coastal Detritic. Biogeography The majority of marine Tardigrada show a geographical distribution highly correlated with that of Tardigrada-workers.To date we can only review the available data concerning the distribution of marine Tardigrada. From a biogeographical 164 S. VILLORA-MORENO AND S. DE ZIO GFUMALDI point of view, marine Tardigrada from the Chafarinas Islands may be divided into five groups: (1) Five new records for the Mediterranean Sea: three species were only known from NE-Atlantic Ocean and two species had an Indo-Pacific distribution. (2) Three probably endemic species: Batillipes marcellii, Actinarctus physophorus, Halechiniscus nsp. (3) Four species have been recorded exclusively from the Mediterranean and NEAtlantic Ocean: Halechinism greveni, Florarctus acer, Actinarctus do?yphorus and Echiniscoides sigbmundi hispanimk. (4)Three species have been recorded exclusively from the Mediterranean and Indo-Pacific Ocean: Florarctus asper, F. hulingi and F. stellatus. (5) Five species show a circumglobal* or Mediterranean- NE Atlantic -1ndoPacific distribution: Sggarctw bradypd, Halechiniscus pgectus, H. remanet*, Florarctus cinctus, and Spracoyx craticuld. Even though more information is now available than in the past, the data are still too scarce and scattered to allow the geographical distribution of the different species to be determined. Since some species, such as Batillipes mirus or Echiniscoides szgzsmundi, have often been found and determined by specialists of other groups, many records must be re-evaluated. ACKNOWLEDGEMENTS We are grateful to Dr Manuel Garcia-Carrascosa, coordinator of the Chafarinas- 91 Expedition. 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