New records of marine Tardigrada in the Mediterranean Sea

~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 .
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*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
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BiogeoLgraph>- . . . . . . .
Acknowledgements . . . . . . .
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Kckrences . . . . . . . . . . . . . . . . . . . . . . . . . . .
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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 [email protected] 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. Benthos survey in the Chafarinas islands was possible through a
collaboration between ICONA (Institute of Nature Conservation)and the University
of Valencia (Spain).The first author was supported through Doctoral (Formacibn de
Personal Investigador) and post-doctoral fellowships from the Spanish Ministry of
Science and Education. Grateful thanks are extended to Dr Reinhardt M.
Kristensen and one anonymous referee for their comments on the manuscript.
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