The first record of the genus Vejdovskybathynella Serban and Leclerc, 1984 (Syncarida, Bathynellacea, Bathynellidae) in the Iberian Peninsula: three new species

Three new species of the genus Vejdovskybathynella are described from caves in northern Spain. It is the first time that this genus has been documented in the Iberian Peninsula. Vejdovskybathynella edelweiss n. sp. has several unique features: the setal formula of the antenna (0/2+exp/2+0/1+0/0+0/2+2/4); the form of the segments and claws of the mandibular palp of the mandible of the male; exopod of the female thoracopod VIII similar in size to the endopod (exopod is always longer than the endopod); small frontal projection of the basipod of the male thoracopod VIII, large or very large is common; the form and size of the lobes of the male thoracopod VIII is unique; the second spine of the furcal rami 2.5 times longer than the first spine. Vejdovskybathynella caroloi n. sp. has a unique male thoracopod VIII with a spur in the exopod that is a feature unknown in the genus, and additionally, this species has the smallest endopod of the thoracopod VIII male of the entire genus; the presence of three aesthetascs in segments VI and VII of the antennule had not been observed in this genus until now; the form of the segments of the mandibular palp of the mandible of the male are unique; the distal spine of the sympod of the uropod is larger than in other species, whereas in the rest of the species of the genus they are equal in size. Vejdovskybathynella pascalis n. sp. is the smallest species of the genus and has two and three aesthetascs in segments VI and VII of the antennule, respectively; it is the only species of the genus that has one seta on segment I and two setae on segment IV of the antenna; it has a unique male thoracopod VIII with a small spur on the frontal projection of the basipod and with the endopod larger than all other species in spite of its small size; the basal spine of the sympod of the uropod is smaller than the other three.


Introduction
The Family Bathynellidae Grobben, 1905 is widespread in Europe (12 genera and 35 species) (Camacho 2006) but has been scarcely studied in the Iberian Peninsula. Some older records mention the presence of the genus Bathynella Vejdovsky, 1882 in Portugal, of caves. The fixative used with the specimens is 4% buffered formalin and they are stored in alcohol (70%).
A complete dissection of all anatomical parts of all type series was made and kept as permanent preparations (special metal slides, glycerine gelatine stained with methylene blue as the mounting medium). Anatomical examinations were performed using an oil immersion lens (1006) of a Zeiss interference microscope with a drawing tube. The descriptions are based on the type series. The material is deposited in the Museo Nacional de Ciencias Naturales, Madrid (MNCN).
The terminology proposed by Serban (1972 and following papers) was used to name Md. and Th. VIII male. Serban worked in depth with the family Bathynellidae (Serban et al. 1971;Serban and Leclerc 1984;Serban 1989aSerban , 1989bSerban , 1992; he revised and described many genera and his terminology here considered is the most accurate and intuitive.
According to Serban (2000), it was assumed that most of France and the Iberian Peninsula is populated by representatives of the subfamily Gallobathynellinae. This subfamily consists of four tribes (the differences between these four tribes are shown in Table II), two subtribes, 10 genera, and 22 species including the three new species described here. The majority of the species occur in France, two are known from Italy, one from Switzerland, one from Germany, and five, including the three new species, from Spain. Table I. Differences amongst the three subfamilies of the Family Bathynellidae: Gallobathynellinae Serban, Coineau, and Delamare Deboutteville, 1971;Bathynellinae Grobben, 1904; and Austrobathynellinae Delamare Serban, 1973 (modified from Serban 1989b Table III shows the systematic data for this subfamily and details the similarities between the species as well as the type locality for each.

Family BATHYNELLIDAE Grobben, 1905
Subfamily GALLOBATHYNELLINAE Serban, Coineau, and Delamare Deboutteville, 1971 Subfamily diagnosis (translated from Serban et al. 1971;Serban 1989aSerban , 1989b Antennule (A. I) six or seven segments; antenna (A. II), three or five segments on the endopod; the apical part of paragnaths with thin setae; prehensile mandibular palp of the mandible (Md.), with one to three segments and both sexes with similar or different claws on the terminal segment; pars incisiva (or incisor process) (''pars incisive'') of mandible with two teeth; processus incisivus accesorius (or molar process) of Md. made up of two parts, the pars molaris normally has two simple teeth or denticles on margins; endopod with three segments on thoracopods (Th.) II, IV, V, and VI or with four segments on all the pairs; male Th. VII without coxal endite. Penial region (''región pénienne'') of Th. VIII with one lobe and with an apical developed frontal projection (Fr. prj.) (''prolongement rostral'') or one lobe and two similar in length lobes or one lobe, one frontal projection and one large Table II. Differences amongst the tribes of the Subfamily Gallobathynellinae: Gallobathynellini Serban, Coineau, and Delamare Debutteville, 1971;Pseudobathynellini Serban, 1989;Sardobathynellini Serban, 1992;Vejdovskybathynellini Serban, 1989(modified from Serban 1989a Table III. Systematic account and distribution (type locality of the species) of the Subfamily Gallobathynellinae Serban, Coineau, and Delamare Deboutteville, 1971.
Tribe VEJDOVSKYBATHYNELLINI Serban, 1989 Tribe diagnosis (translated from Serban 1989aSerban , 1989b A. I and A. II with usual structure. Md.: mandibular palp with three segments with the setae longer in the males than in the females. Th. I-VII: endopod with four segments on all the pairs. Male Th. VIII: penial region (''región pénienne'') with one lobe that shows an elongated frontal projection (''prolongement rostral'') and an outer lobe (''lobe externe'') shorter than the projection, vertical basipod with a very prominent frontal crest (Fr. crt.) (''crête rostral'') and with a distal region curved on the external side, forming a spur (''eperon'') (S. fr. crt.); endopod is a single segment, lengthened and shorter than the exopod. Female Th. VIII with habitual structure and coxal seta elongated.
Type locality: Cubilla Cave, Ogarrio, Cantabria, Spain a Species not confirmed. basis has a vertical position and a prominent frontal crest (Fr. crt.). Female Th. VIII: with a long lateral internal seta on the coxopod; exopod very simplified. Uropod: sympod with four spines and endopod with two or three spines. Furcal rami: the second spine is the longest.
Vejdovskybathynella edelweiss n. sp. (Figures 1-4 The details of the locality type and the chemical and physical nature of the sampling points can be obtained from Camacho et al. (2006).

Description
Body. Total length of holotype (male) 0.81 mm and paratype (female) 0.76 mm. The sizes of the specimens in the different populations found are shown in Table IV. The greatest male found measured 0.90 mm and the smallest 0.58 mm; the largest female measured 1.3 mm and the smallest 0.81 mm. Body form can be observed in Figure 1. All drawings are of the holotype (male) and the allotype (female, Th. VIII and Md.).
Antennule (A. I) (Figure 2A). Seven segments; length of first three segments similar to the first of the other four segments; segments 4 and 5 smaller than the others; setation as in Figure 2A; two aesthetascs on each of segments 6 and 7. A. I slightly longer than the antenna.
Paragnath ( Figure 2E). Lengthened and with a projection in the distal part and thin setation.
Mandible (Md.) ( Figure 2C, K). Palp with three segments, terminal segment with one plumose claw and another smooth and expanded in the top (see Figure 2C) and segment 2 with a special form expanded at the distal end; in the females segment 2 is more or less cylindrical, without expansions, and claws of segment 3 are equal and smooth ( Figure 2D). Masticatory part ( Figure 2K): pars molaris with two main teeth, the first (the nearest to processus incisivus accessorius) simple, the second tooth with one small tooth on inner side and one small tooth in the outer side (secondary formatons); incisor process (pars incisiva) of two teeth.
Maxillule (Mx. I) ( Figure 2F). Proximal endite with four setae; distal endite with six teeth, four with spines (denticles) and two smooth, and with three plumose setae on the outer margin.
Female thoracopod VIII (Th. VIII) ( Figure 2J). Coxa with one smooth lateral seta; large epipod, longer than the basipod; endopod with single segment with two apical setae, one of these longer and thicker than the other; exopod slightly more slender than endopod, of similar length and with two apical smooth setae of different lengths.
Uropods ( Figure 4F). Sympod slightly longer than endopod, twice longer than broad and with four equal spines; endopod almost twice as long as the exopod, with three strong claws, the most distal is the longest (twice the length of the second), terminally with two setae (one of these very long) and with two shorter ones located dorsolaterally; exopod with two terminal and two medial setae.
Furcal rami ( Figure 4E). Almost square, bearing five spines; long dorsal spine, almost twothirds the length of second spine, which is 2.5 times longer than the other three spines, which are all almost of the same size.

Etymology
The species name is derived from the Edelweiss Speleological Club from Burgos, which has been working in the karst of Ojo Guareñ a for over 50 years. The name is in gratitude to their members for finding this new species.

Description
Body. Total length of the holotype (male) 0.61 mm and paratype 0.60 mm. The sizes of the species in the different populations found are shown in Table IV. All drawings are of the holotype (male) and the allotype (female, Th. VIII and Md.).
A. I ( Figure 5A). Seven segments; length of first three segments similar to the first of the other four segments; setation as in Figure 3A; on segments 6 and 7 three aesthetascs. A. I slightly longer than the antenna.
Paragnath ( Figure 5D). Lengthened and with a rounded projection in the distal part, and with setation.  Md. (Figure 5E, F). Palp with three segments, terminal segment with two claws with setation as shown in the figure, and first two segments thick and with rounded aspect; in the females segments 1 and 2 are normal, the second longer, without expansions, and claws of segment 3 equal and with few setae. Masticatory part ( Figure 5F): pars molaris with two main teeth, the first (the nearest to processus incisivus accessorius) bidentate, the second tooth with three small teeth (secondary formations) on each side; incisor process (pars incisiva) of two teeth.
Mx. I ( Figure 5G). Proximal endite with four setae; distal endite with six teeth, five with spines (denticles), and with three plumose setae on the outer margin.
Male Th. VIII ( Figure 5K, L). Outer lobe (O. lb.) very large, the distal edge almost reaches the edge of the frontal projection; frontal projection (Fr. prj.) with transverse circular section; vertical basipod (Bsp.) not very long and almost square; frontal crest (Fr. crt.) of basipod prominent and with a very evident spur (S. fr. crt.); exopod heavy, with a spur (see Figure 5K), with four setae on the distal end. Very small endopod, one-seventh the size of the exopod, with two smooth terminal setae.
Female Th. VIII ( Figure 5M). Coxa with one smooth lateral seta; large epipod, longer than basipod; endopod singly segmented with two plumose apical setae, one of these longer than the other; exopod longer than the endopod, and with two apical smooth setae of different lengths.
First pleopods ( Figure 5N). Two segments, first segment with one seta; second segment with five setae.
Uropods ( Figure 6I). Sympod slightly longer than endopod, twice longer than broad and with four spines, distal longer than the others; endopod twice as long as the exopod, with three strong claws, the most distal slightly longer, terminally with two setae (one of these very long) and with two shorter ones dorsolaterally; exopod with two terminal and two medial setae.
Furcal rami ( Figure 6H). Almost square, bearing five spines; second spine is four times longer than the other four spines, which are all almost of the same size including the dorsal spine.

Etymology
The species name is derived from Carolo, who has worked for more than 23 years with the author and is a best friend, sampling companion, and husband to the author.

Remarks
Vejdovskybathynella caroloi n. sp. has a combination of unique characters (Tables V, VI); it has a unique male thoracopod VIII, with a spur in the exopod that is a feature otherwise unknown to the genus (see Table VI); the basipod is small and the outer lobe and outer protuberance are similar in size; the frontal projection is as short as the basipod; the endopod is the smallest of the whole genus. The presence of three aesthetascs on segments VI and VII of the antennule also was not previously known in this genus; the form of the segments of the mandibular palp of the mandible of the male are unique; the setal formula of the endopod of all thoracopods is unique (see Table V), as having three setae on the basipod of Th. I only occurs in the French species, whereas the Spanish species only has two (Table V) and Th. VI-VII are like those of V. pascalis n. sp.; the distal spine of the sympod of the uropod is larger than the others, whereas in the rest of the species they are all equal in size. The setal formula of A. II is similar to the French species. The male is medium sized and the female is large. A. I ( Figure 7A). Seven segments; length of the first three segments similar to that of four remaining segments; setation as in Figure 7A; two and three aesthetascs on segments 6 and 7, respectively. A. I longer than A. II.
Paragnath ( Figure 7D). Lengthened and with a projection similar to a tooth in the distal part and with setation.
Md. (Figure 7G, M). Palp with three segments, terminal segment with two claws of similar length and with setation in proximal part, and segment 2 thick, almost square; in the females the segments are normal, the second longer, and claws of segment 3 are unequal and with few setae on the distal end ( Figure 7E). The masticatory part of the Md. of the male broke during the dissection and for this reason there is no drawing. Pars molaris with two main teeth in the female ( Figure 7F, M), the first with a small tooth (secondary  formation) near to processus incisivus accessorius, the second tooth with a small tooth (secondary formation) in the outer side; incisor process (pars incisiva) with two teeth.
Th. I-VII ( Figure 8A-G). Th. I smaller than others; Th. II-V similar in size and Th. VI-VII longer than others. Th. I ( Figure 8A) without epipod; coxa with a long plumose seta; basipod with two plumose setae. Basipod of Th. II ( Figure 8B) with two smooth setae and with only one smooth seta on the rest of the thoracopods. Exopod with one segment on all thoracopods, of equal length to the endopod in Th. I-V, and shorter in Th. VI ( Figure 8F) and VII ( Figure 8G) Female Th. VIII ( Figure 7L). Coxa broke in the dissection; epipod long, longer than basipod; endopod small, a single segment with two smooth apical setae; exopod twice as long as endopod, and with two smooth apical setae of different lengths, one of these very long.
Uropods ( Figure 8H). Sympod slightly longer than endopod, a little longer than wide and with four spines, the basal shorter than the others; endopod 1.5 times longer than exopod, with two strong claws, distal twice as long as the other, terminally with two setae (one of these very long) and with two located dorsolaterally; exopod with two terminal and two medial setae.
Furcal rami ( Figure 8I). Almost square, bearing five spines; second spine twice as long as the other three spines, which are shorter than the dorsal spine.

Etymology
The species name is derived from the PASCALIS European Project, thanks to which these populations of Syncarida could be studied.

Remarks
Vejdovskybathynella pascalis n. sp. is the smallest species of the genus with a unique combination of characters (Tables V, VI); it has two and three aesthetascs on segments 6 and 7 of the antennule, respectively. To have two aesthetascs on segment 6 is common in the genus (except V. caroloi n. sp. which has three) but to have three in segment 7 is unusual, and only V. caroloi n. sp. shares this character; it is the only species of the genus that has one seta on segment 1 and two setae on segment 4 of the antenna. This is the only Spanish species of this genus that has an exopod of Th. I-V longer than the endopod, and the relation of sizes is not known in the French species. The setal formula of the endopod of the thoracopods is unique and the number of setae on the basipod of the thoracopods is also unique; it has a unique male thoracopod VIII, with a small spur of the frontal crest of the basipod (absent or present, but large in this case, in the other five species of the genus) and with the endopod the largest of all species in spite of its small size. The basal spine of the sympod of the uropod is smaller than the other three; the dorsal spine is longer than the first spine, as only occurs in V. edelweiss n. sp., in the other species the two spines are a similar size.v

Discussion
Tables V and VI show the similarities and differences between the six species of the genus Vejdovskybathynella. In the original description of V. espattyensis Serban and Leclerc, 1984 only the differences with the species type V. balazuci Serban and Leclerc, 1984 were mentioned and no drawings of the majority of the parts of the body were given. For this reason, there is not the same level of detail for comparison as in the other species. For example, Mx. I and Mx. II are unknown in the French species. The largest species are V. leclerci and V. edelweiss n. sp.; the smallest is V. pascalis n. sp. The three French species show more similarity among them than is seen between the Spanish species.
Only V. caroloi n. sp. has three aesthetascs on segment 6 of A. I. Only the species V. pascalis n. sp. has setae on the first segment of A. II; the medial seta of the exopod of A. II is absent in V. leclerci, V. caroloi n. sp., and V. pascalis n. sp.
The sexual dimorphism in the mandibular palp of the Md. is most evident in V. edelweiss n. sp. and V. caroloi n. sp. while in the other species it is very subtle.
The setal formula of the endopod of Th. I-VII in the French species is similar, with only slight differences in the number of setae of Th. III-V (Table V). The Spanish species differ in all setae on the segments of the endopod of Th. I-VII as well as in the number of setae on the basipod of the thoracopods (Table V); only V. edelweiss n. sp. has one seta on the first segment of the endopod of Th. VI and VII. The exopod of the thoracopods is smaller than the endopod on all thoracopods in V. edelweiss n. sp. and V. caroloi, whereas it is larger on Th. I-V in V. pascalis n. sp.
In the female Th. VIII the exopod is larger than the endopod, except in V. edelweiss n. sp. in which they are similar in size.
The male Th. VIII of V. edelweiss n. sp. is more similar to Th. VIII of V. balazuci and V. espattyensis than to the Spanish species, though it has a less-developed frontal projection and a more rectangular basipod than the French species. The male Th. VIII of V. caroloi n. sp. and V. pascalis n. sp. have a more square aspect.
The pleopod of the new Spanish species (unknown in V. pascalis n. sp.) has five setae on the second segment, whereas it has four or six setae in the French species.
Vejdovskybathynella pascalis n. sp. has two claws on the endopod of the uropod as in the French species; in the rest of the new Spanish species there are three claws present.
Vejdovskybathynella edelweiss n. sp. and V. pascalis n. sp. have dorsal spines on the furcal rami that are longer than the first spine, and are similar in the rest of the species; the second spine of the furcal rami is very long in V. balazuci, V. espattyensis, and V. caroloi n. sp. (four times longer than the first spine).
There is little doubt that these three new species found in Spain belong to the genus Vejdovskybathynella described from the south of France and this confirms the presumption of the presence in Spain of other genera of the Family Bathynellidae, than Bathynella, as was postulated by Serban (2000).