ONE NEW MEMBER OF THE SUBTERRANEAN FAMILY NIPHARGIDAE FROM SPAIN, NIPHARGUS SPIRITUS, SP. N. (CONTRIBUTION TO THE KNOWLEDGE OF THE AMPHIPODA 291) ABSTRACT From the subterranean waters of the northern Spain, Fuente Aizpara, on the northern slope of Aizpara

From the subterranean waters of the northern Spain, Fuente Aizpara, on the northern slope of Aizpara Mt., Zugarramurdi, Navarra, one new species of the family Niphargidae (Crustacea, Amphipoda, Gammaridea) is described and figured, Niphargus. spiritus, sp. n. This species is characterized by presence of scarce number of additional spines on some of pereopod-dactyls, elongated inner ramus of uropod 1 and distal article of uropod 3 in males, long spines on telson, numerous setae along outer margin of dactylus on gnathopods 1 and 2, by presence of only 1-2 setae on maxilla 1 inner plate, etc. The relation between N. spiritus and other known species of this genus known from Spain and France is discussed.


INTRODUCTION
The genus Niphargus Schiödte, 1849 (Amphipoda, family Niphargidae) is widely spread over Europe and Near East, on west to Great Britain and France, but in Spain is present in the northern part of Iberian Peninsula only, mainly towards French border.
The number of known taxa of genus Niphargus in Spain is very scarce. Margalef (1952) described a new taxon Niphargus ciliatus cismontanus, ssp.n. from Guipuzcoa (Aranzazu: cueva de Guesaltza). G. Karaman mentioned (1986a) Niphargus delamarei Ruffo 1954 from Mosquera cave (= Cova de la Mosquera), Beuda, Gerona province, and later (2015a) described Niphargus notenboomius, sp. n, and N. laisi geronensis, ssp. n.. He cited (2015b) Niphargus gallicus Schell., 1935 andN. delamarei Ruffo, 1954 for several new Spanish localities. Our recent study of material from Spain, collected by Prof. Dr. Jos Notenboom and other collectors, indicated the presence of some other taxa of this genus in Spain, related to the known species from France. Here is presented the results of study of part of these samples.  3E). Epimeral plates 1-2 poorly angular, with marked ventroposterior corner spine-like seta and convex posterior margin bearing a row of short setae; epimeral plate 3 angular, with marked ventroposterior corner by one spine-like seta and almost straight posterior margin bearing a row of short setae ( fig. 5D). Epimeral plate 2 with 1 subventral spine, epimeral plate 3 with 2 subventral spines ( fig. 5D).
Head with short rostrum and short subrounded lateral cephalic lobes, ventroanterior sinus is developed, eyes absent ( fig. 1A).
Pereopod 4: pilosity of article 2 likes that in pereopod 3; articles 3 and 4 at posterior margin with bunch of longer setae ( fig. 1G). Articles 4-6 of unequal length (ratio: 50:34:44); article 4 along posterior margin with 4 bunches o setae (the longest setae are longer than diameter of article itself); article 5 at posterior margin with short spines and setae; article 6 along posterior margin with bunches of short spines and setae. Dactylus much shorter than article 6 (ratio: 19:44), along inner margin with 1-2 strong spines ( fig. 1H, I), at outer margin with one median plumose seta; nail nearly as long as pedestal.
Pereopods 5-7 moderately strong, Pereopod 5 remarkably shorter than pereopods 6 and 7; article 2 longer than broad (ratio: 76:46), along anterior margin with several longer spine-like setae, along posterior almost straight margin with nearly 11 short setae, ventroposterior lobe small ( fig. 4A). Articles 4-6 of unequal length (ratio: 51:55:62), article 4 along anterior margin with 4 bunches of long setae (the longest setae slightly longer than diameter of article itself), along posterior margin with 3 short spine-like setae (fig. 4A); article 5 at anterior margin with 3 groups of spines and longer spine-like setae, along posterior margin with 3 groups of short spines; article 6 along both margins with bunches of short spines accompanied at anterior margin by single setae. Article 6 is slightly shorter than article 2 (ratio: 76:62), with distal bunch of long setae exceeding half of article 6-length.
Dactylus moderately slender, much shorter than article 6 (ratio: 21:62), at inner margin with one strong spine and seta near basis of the nail, along outer margin with one median plumose seta (  Dactylus is moderately slender, much shorter than article 6 (ratio: 30:95), along inner margin with one strong spine and seta near basis of the nail, along outer margin with one median plumose seta ( fig. 4E) and 0-1 pair of simple setae (fig. 4F); nail is shorter than pedestal (ratio: 25:50 or 28:46).
Epimeral plates are hardly more angular than these in male. Epimeral plate 1 with ventroposterior corner marked by one spine-like seta, posterior slightly convex margin is provided with 5-6 short setae ( fig. 7E); epimeral plate 2 with marked ventroposterior corner by one spine-like seta and less convex posterior margin bearing 6-7 short setae. Epimeral plate 3 distinctly angular with marked ventroposterior corner by one spine-like seta, posterior margin slightly inclined and almost straight, bearing several short setae. Epimeral plate 2 with one subventral spine, epimeral plate 3 with 2 subventral spines ( fig. 7E).
Head like that in male. Antenna 1 slightly longer than half of body-length (ratio: 49:78), main flagellum with 25 articles. Antenna 2 like that in male, flagellum consisting of 12 articles.
Maxilliped: inner plate with 3 distal spines; palpus article 3 at outer margin with one median and one distal bunch of setae; article 4 at inner margin with 2 setae near basis of the nail, at outer margin with one median seta.
Gnathopod 2: article 2 along anterior margin with row of long setae, along posterior margin with several bunches of long setae; article 4 with posterodistal bunch of long setae; article 5 is shorter than propodus (ratio: 40:45), along anterior margin with 2 bunches of setae ( fig. 6C). Propodus trapezoid, slightly longer than broad (ratio: 87:78), along posterior margin with 9 transverse rows of setae ( fig. 6D). Palm inclined slightly less than half of propodus-length, convex, defined on outer face by one corner S-spine accompanied laterally by 3 slender L-spines and 5 facial setae, on inner face by one subcorner R-spine. Dactylus reaching posterior margin of propodus, along outer margin provided with 10 single or paired setae, on inner margin with row of short setae ( fig. 6D).
Pereopods 3 and 4 are moderately slender. Pereopod 3: article 2 along anterior margins with long proximal setae and short distal setae; along posterior margin appear long setae in proximal part and shorter setae in distal part (fig. 7A); articles 3 and 4 at posterior margin with long setae (the longest setae much longer than diameter of articles themselves). Articles 4-6 of different length (ratio: 55:33:40). Article 4 along posterior margin with 4 groups of setae (the longest setae are much longer than diameter of article itself). Article 5 at posterior margin with 3 bunches of spines and short setae; article 6 along posterior margin with 5 bunches of short spines and setae. Dactylus much shorter than article 6 (ratio: 20:40), along inner margin with 2 strong spines, along outer margin with one median plumose seta (fig. 7B); nail shorter than pedestal (ratio: 32:37).
Dactylus is much shorter than article 6 (ratio: 31:96), along inner margin with one strong spine and seta near basis of the nail, along outer margin with one median plumose seta ( fig. 8 I); nail shorter than pedestal (ratio: 37:69).
Pleopods 1-3 with 2 retinacula each. Peduncle of pleopod 1 with 2 distal setae along anterior margin ( Uropod 1: peduncle longer than rami, with dorsoexternal row of spines; at dorsointernal margin with one spine and one seta (except distal spine) ( fig. 6E). Outer ramus is slightly shorter than inner ramus, bearing several lateral and distal strong spines and one lateral pair of short simple setae ( fig. 6E); inner ramus with several lateral and distal strong spines and bearing 2 median and 3 distal short simple setae.
Uropod 3: peduncle longer than broad (ratio: 44:23), bearing distal spines ( fig. 7F). Inner ramus much shorter than peduncle, bearing 3 distal spines ( fig.  7F). Outer ramus 2-articulated: first article along outer margin with 3 bunches of strong spines and one bunch of short simple setae, along inner margin with 5 bunches of strong spines; 3 longer plumose setae are attached near bunches of spines ( fig. 7F); second article much shorter than first one (ratio: 50:130) bearing along both margins the bunches of simple setae; the tip is pointed and naked Telson as long as broad, incised nearly 2/3 of telson-length ( fig. 5L); each lobe with 3 distal and one outer marginal very long spine; one short spine is attached in the middle of outer margin of one lobe; a pair of short plumose setae is attaches near the middle of outer margin in each lobe ( fig. 5L).
The one normal and often one additional spine are present on some of pereopod-dactyls [on pereopod 3, pereopod 4, pereopod 5], but pereopods 6 and 7 were always with only one spine at inner margin. Some specimens are without this additional spine. Evidently, the presence of additional spine on some dactyls of pereopods 3-5 is characteristic for this species, despite the fact that some specimens are without it.
Article 2 of pereopods 5-7 is with well to poorly marked ventroposterior lobe. Urosomal segment 1 in males and females is always with 1 seta only on each dorsolateral side. Urosomal segment 2 on each dorsolateral side in males is with 2 setae or one spine and seta, in females with one spine and one seta, or 2 spines. Telson is with long spines. Females have usually poorly more angular epimeral plate 3 than the males. Peduncle of uropod 1 along dorsointernal margin is with 1-2 median setae or one spine and one seta (except distal spine). The further discovery of this species in other localities will show the limits of variability of this species. HOLOTYPE: male 12.0 mm; paratype female 7.8 mm. Holotype and paratypes are deposited in Karaman`s Collection in Podgorica, Montenegro. DISTRIBUTION: Known from type-locality only. DERIVATIO NOMINIS. The name "spiritus" arrives from the Latin word "spiritus", adequate word "spiritus" in English. Fig 7. Niphargus spiritus, sp. n., Fuente Aizpara, female 7.8 mm (paratype). A-B= pereopod 3; C-D= pereopod 4; E= epimeral plates 1-3; F= uropod 3.

REMARKS AND AFFINITIES
Niphargus spiritus, sp. n. is very close to Niphargus ciliatus Chevreux, 1906 [Loc. typ.: Grotte de Meailles, Valee de la Vaire, affluent du Var, France]. As the description of this species was scarce, Ginet (1988 and1991) redescribed this species more in detail, and based on this description is evident that N. spiritus, sp. n. is rather similar to N. ciliatus by various characters: elongated uropod 3, elongated inner ramus of uropod 1 in males, shape of pereopods 3-7 except dactyls, by pleopods with 2 retinacula, etc. But N. ciliatus differs from N. spiritus by strong dactylus of pereopods 3-7 bearing higher number of spines each, by inner plate of maxilla 1 bearing 2-4 setae, telson with distal and inner marginal and facial short spines, elevated number of dorsolateral spines on urosomal segments 1 and 2, slightly pointed epimeral plate 3 in male, etc. Margalef (1952) Margalef, 1952], provided with strongly angular or almost pointed epimeral plate 3, dactylus of pereopod 7 with 4 spines, maxilla 1 inner plate with 2 setae, outer plate with 8 spines bearing one lateral tooth; telson with short spines along outer and inner margin, face and tip of each lobe. Margalef (1970) figured partially again this species: epimeral plate 3 poorly angular, nearly like that in our specimens, telson with short spines along inner margin and tip only, and dactylus of pereopod 7 with 6 spines along inner margin.
Chevreux figured entire male of this species only, with rectangular epimeral plate 3, mentioning longer coxae, and elongated uropod 3 in males only, with remarks that "this variety don`t differ significantly from N. plateau". On page 232 (1901) he figured uropod 3 of female from Robine with short distal article of outer ramus. France. Ginet (1991) presented several figures of this species: dactylus of pereopod 3 and pereopod 7 with 3 spines along inner margin, inner plate of maxilliped with 5 spines, urosomal segment 2 with 6 spines on each dorsolateral side, etc. Chevreux & Fage (1925) cited N. robustus and figured male 21 mm from Padirac with large gnathopods 1-2 propodus exceeding the width of corresponding coxae, propodus palm inclined over half of propodus-length, and telson bearing short distal and outer marginal spines. Later nobody redescribed this taxon in detail, but based on known taxonomical characters of N. robustus, our specimens from Fuente Aizpara are not identic with N. robustus.  Chevreux, 1896 [Loc. typ.: Grotte de Arbois, de Beaumeles-Messieurs, de Baume-les-Dames, Jura, France] has long lobed coxa 4, epimeral plates strongly angular to poorly acute, telson with 2 facial and 5-6 distal short spines; maxilla 1 inner plate with 3 setae, dactylus of pereopods 3-7 with one spine, etc.
Niphargus sestoputeanus G. Karaman, 2016 [Loc. typ.: Sesta Godano, Passo del Rastrello, 1000 m a.s.l. (N. of La Spezia) is also provided with additional spines on dactylus of pereopods, but differs from our species by elongated inner ramus of uropod 3 in male, by pointed epimeral plates in males and females, by strong dactylus of pereopods 3-7 bearing higher number of spines, by higher number of setae on maxilla 1 inner plate, etc.
Niphargus puteanus Koch, in Panzer, 1836 [Loc. typ.: Weichelmuhle near Ratisbonne (= Regensburg), Germany] was mentioned by various authors from many localities of Europe (often erroneously), because the good description of this species was not published. Stock (1974) redescribed this species from Weichselmühle, Ratisbonne, Germany]. We compared the specimens from this locality with N. spiritus from Spain, and N. puteanus differs from our species by remarkably higher coxae, by more pointed epimeral plates, by presence of dorsointernal row of spines on uropod 1 peduncle in male, almost quadrate propodus of gnathopod 2, by higher number of setae on maxilla 1 inner plate, shorter spines on telson, etc.
The species Niphargus rhenorhodanensis Schellenberg, 1937 [Loc. typ.: Rumingen, N of Lorrach, Germany], redescribed by Ginet (1985), and N. orbis G. Karaman, 2013 [Loc. typ.: Calizzano, Alpi Ligure, Rio di Valle, Italy] are rather similar to our species, but both of them differ from N. spiritus by absence of additional spines on dactylus of pereopods 3-7. The most of known Niphargus species from France are poorly or partially described, and some of its taxonomical characters are still unknown. By this way to establish the real taxonomical relations between newly described taxa and taxa described one century ago, remains very difficult and rather hazardous. The further studies and redescription of known taxa will help in establishing of the real relationships among various Niphargus taxa and understand the limits of variability of each taxon.

CONCLUSION
The fauna of the genus Niphargus Schiödte, 1849 (Amphipoda Gammaridea, fam. Niphargidae) is very poorly known from the subterranean waters in Spain, and only several species and subspecies of this genus have been described or mentioned from various localities of Spain. Genus Niphargus settled the subterranean waters in northern part of Spain only (Karaman, G. 1986b), replaced in other part of country by members of genus Haploginglymus Mateus & Mateus, 1958. The known members of genus Niphargus from Spain are related to these of France, but scarce description of many taxa from France made very difficult recognition of real relations among all known taxa in France, and consequently, in Spain also.