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A New Species of Typhlatya (Crustacea: Decapoda: Atyidae) from Anchialine Caves on the French Mediterranean Coast

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A New Species of Typhlatya (Crustacea: Decapoda: Atyidae) from Anchialine Caves on the French Mediterranean Coast Blackwell Science, LtdOxford, UKZOJZoological Journal of the Linnean Society0024-4082The Lin- nean Society of London, 2005? 2005 1443 387414 Original Article NEW ANCHIALINE ATYID FROM MEDITERRANEAN CAVESD. JAUME and F. BRÉHIER Zoological Journal of the Linnean Society, 2005, 144, 387–414. With 19 figures A new species of Typhlatya (Crustacea: Decapoda: Atyidae) from anchialine caves on the French Mediterranean coast DAMIÀ JAUME1* and FRANCK BRÉHIER2 1IMEDEA (CSIC-UIB), Instituto Mediterráneo de Estudios Avanzados, C/. Miquel Marquès 21, E-07190 Esporles (Mallorca), Spain 2Alas, F-09800 Balaguères, France Received June 2004; accepted for publication March 2005 A new species of the thermophylic Tethyan relict prawn Typhlatya is described from two anchialine caves near Per- pignan (southern France). The new species is closely related to a congener known only from a freshwater cave at Cas- tellón (eastern Spain), about 400 km to the south-west, differing apparently only in the size and shape of the rostrum and the armature of the dactylus of the fifth pereiopod. Based on palaeogeographical evidence and assuming a sister- group relationship between both species, we suggest that their common ancestor could not be older than early Pliocene in age, and that it was already a stygobiont taxon adapted to live in shallow-water marine crevicular habitats. This ancestor would have vanished from the western Mediterranean after the cooling associated with the onset of northern Hemisphere glaciation, about 3 Mya, as documented for other Mediterranean marine taxa. Indeed, the genus is completely stygobiont and does not occur in fluvial environments. The Pyrenees represent a watershed boundary that eliminates the possibility of the derivation of one species from the other by active dispersal after establishment in continental waters. © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 387–414. ADDITIONAL KEYWORDS: biogeography – France – stygofauna – systematics – Tethyan relicts – vicariance. INTRODUCTION France to the Black Sea coast of the Republic of Georgia. In 1973 the French biologist J. Y. Bertrand collected a Two other epigean, occulate atyid genera occur in single specimen of a blind atyid prawn in the cave of the area and are endemic to it: the first is Dugastella Font Estramar, a small cave opening just above a Bouvier, 1912, with two species, one restricted to spring of the same name in Massif des Corbières Morocco, the other to coastal marshes of the Spanish (Pyrénées-orientales, France). He identified it as Tro- Levant; the second is the circum-Mediterranean glocaris sp., presumably with reference to the pres- Atyaephyra desmaresti (Millet, 1931), with three rec- ence of Troglocaris inermis Fage, 1937 in the nearby ognized subspecies (d’Udekem d’Acoz, 1999). A third karsts of Hérault, where the stygofauna is similar to genus, Caridina H. Milne Edwards, 1837, broadly that of Corbières (Bertrand, 1974). Hitherto Troglo- spread along the Indo-Pacific, Asia and Africa south caris Dormitzer, 1853, together with the monotypic to the Sahara, has been reported from close to the Spelaeocaris pretneri Matjasic, 1956 from caves in Mediterranean Levantine Region (Mediterranean Herzegovina, were the only stygobiont atyid genera Nile; Suez Canal; Syria; see d’Udekem d’Acoz, 1999), known from the peri-Mediterranean region. Troglo- but not from the Mediterranean proper. caris, with nine recognized species (see d’Udekem The Mediterranean was apparently devoid of stygo- d’Acoz, 1999) is limited to coastal regions on the north- biont atyids belonging to thalassoid genera displaying ern rim of the Mediterranean, from the Gulf of Lyon in extreme disjunct distributions, such as those known in oceanic island groups of both the Atlantic and Indo- Pacific. That was the picture until the remarkable dis- *Corresponding author. E-mail: [email protected] covery of a new species of Typhlatya Creaser, 1936 in © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 387–414 387 388 D. JAUME and F. BRÉHIER Spain by Sanz & Platvoet (1995), who also transferred somites, and telson. Material is deposited in the the monotypic Spelaeocaris to Typhlatya. This repre- Crustacea collection of the Departement des Milieux sented the first undeniable relict Tethyan distribution et Peuplements Marins, Muséum national d’Histoire known among European decapods. Later, based on the naturelle, Paris (MNHN). emended diagnoses of atyid genera presented by Holthuis (1993), d’Udekem d’Acoz (1999) added another Eurasiatic Typhlatya to the list, by transfer- THE CAVES ring the Georgian Troglocaris schmidti jusbaschjani Font Estramar is an important artesian spring that Birstein, 1948 to Typhlatya (as T. jusbaschjani). opens in the limestone of the Massif des Corbières In November 2001, and again in 2002 and 2003, (Early Cretaceous; see Aunay et al., 2002) about during the Pascalis project (see http://www.pascalis- 150 m inland from the inner shore of the Mediterra- project.com/), Font Estramar was repeatedly surveyed nean coastal lagoon of Salses-Leucate (Salses-le- by one of us (FB) in order to catalogue its stygofauna. château; Pyrénées-orientales, south-west France; The prawn recorded there by J. Y. Bertrand was col- Fig. 1). The lagoon, with a surface area of 54 km2 and lected in large numbers. Following detailed compari- a maximum depth of 3.70 m, is separated from the sea son with material from the type locality, it was by a sand bar. Water salinity is slightly lower (22– recognized as a related, new species, distinguishable 35‰, depending on the local conditions) than in the by the ornamentation of the dactylus of the fifth sea nearby (37.1‰). pereiopod and by the shape of the rostrum, from Underwater exploration of the spring started in Typhlatya miravetensis Sanz & Platvoet, 1995, which 1949 and the extent of surveyed passages has is endemic to a single limestone freshwater cave in the increased progressively since then, in line with the Spanish province of Castellón. In 2003 we found the improvements in cave diving techniques. Considering prawn in a second cave, the Grotte des Fées de Leu- the complexity of the cave, the Association de Recher- cate, situated some 7 km to the north-east of Font ches de Font Estramar (ARFE) was created in 1991 to Estramar. coordinate its exploration, leading to the survey of This paper provides a detailed description of the more than 2800 m of underwater passages. The cur- new species, with a special emphasis on the determi- rent explored limit in the cave was reached in 1996 by nation of the segmental homologies of all limbs, in ARFE’s cave diver Cyrille Brandt, who reached a order to provide a better basis for comparison with depth of -164 m at 850 m from the entrance (Brandt, other species of the genus and with related genera. 1997). It has been estimated that the karstification at The paper also constitutes the first record of anchia- the spring area extends 350 m, with an even greater line caves in metropolitan France. potential as the thickness of the limestone is around 1000 m (Ladouche et al., 2004). Font Estramar and Font Dame are the main water MATERIAL AND METHODS sources within the large and complex karstic system of Most of the prawns were caught in traps baited with Corbières d’Opoul (Salvayre, 1978, 2002). The large strips of raw beef and set in the submerged passages associated aquifer derives around 30% of its water of the caves using specialized cave diving techniques. from rainfall, the remainder coming from sinks in the Other specimens were captured by hand, the animals Agly and Verdouble riverbeds (Ladouche et al., 2004). often entering the open water column and being illu- The outflow of Font Estramar has, over the last minated by torchlight. Salinity and temperature pro- 30 years, averaged 2.1 m3 s-1, fluctuating from files of the water column in Grotte des Fées de Leucate 0.8 m3 s-1 to more than 30 m3 s-1 during floods. The were produced with a Seabird SBE19 CTD oceano- water is characterized by a slightly elevated tempera- graphic profiler programmed to take readings every ture and oligohaline state (salinity range 0.2–2.2‰, 0.5 s. The descent of the profiler was controlled to a depending on the spring’s outflow volume; Kiener & rate of c. 10 cm s-1; the device was held out in front by Petit, 1968; Ladouche et al., 2004). Recent studies the diver to avoid disturbing the water column. Speci- (Ladouche et al., 2004) indicate that the salinity mens were fixed in the field in ethanol and treated in derives from the water of the coastal lagoon. The deep the laboratory with hot lactic acid to remove internal intrusion and circulation of this water would explain tissue. Black Chlorazol B cuticular staining was used its elevated temperature, ranging from 15.6 to 18.6 ∞C following the procedure described in Wagner (1994). at the spring’s outlet (Cyrille Brandt, pers. comm.). Drawings were prepared using a camera lucida on an Typhlatya arfeae sp. nov. has been collected in Font Olympus BH-2 microscope equipped with Nomarski Estramar from the entrance to the so-called Puits du differential interference contrast. Body measurements Silence (see map in Brandt, 1997), although it has also were derived from the sum of the maximum dorsal been observed by divers further within the cave. The dimensions of the carapace (rostrum included), free species seems to be more abundant in the Galerie © 2005 The Linnean Society of London, Zoological Journal of the Linnean Society, 2005, 144, 387–414 NEW ANCHIALINE ATYID FROM MEDITERRANEAN CAVES 389 Figure 1. Location of the two caves harbouring Typhlatya arfeae sp. nov. in south-eastern France plus the single locality of the closely related T.
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