Research Article ISSN 2336-9744 (Online) | ISSN 2337-0173 (Print) the Journal Is Available on Line At

Research Article ISSN 2336-9744 (online) | ISSN 2337-0173 (print) The journal is available on line at www.biotaxa.org/em

https://zoobank.org/urn:lsid:zoobank.org:pub:7A76F10A-148C-477F-B978-B88C16F49EF3

One new species of the family Niphargidae from Bosnia and Herzegovina (Contribution to the Knowledge of the Amphipoda 289)

GORDAN S. KARAMAN

Montenegrin Academy of Sciences and Arts, 20000 Podgorica, Montenegro E-mail: [email protected]

Received 10 March 2016 │ Accepted 24 April 2016 │ Published online 26 April 2016.

Abstract From the subterranean waters of the Velika Bukovička špilja Cave near Bukovica village (Tomislavgrad region, Bosnia and Herzegovina) is described and figured one new species of the family Niphargidae (Amphipoda, Gammaridea), Niphargus bukovicensis, sp. n. This species is characterized by presence of elevated number of spines on dactylus of pereopods 6 and 7 only, by numerous setae along dactylus outer margin of gnathopods 1-2, pointed epimeral plate 3 and by elongated inner and outer ramus of uropod 3 in males. The taxonomical relation of this species regarding other similar members of genus Niphargus Schiödte, 1849, is discussed.

Key words: taxonomy, new species, Amphipoda, Niphargus bukovicensis, Bosnia and Herzegovina, cave.

Introduction

The fauna of the Amphipoda in Bosnia and Herzegovina has been studied by various authors and over 60 species of various families (Gammaridae, Niphargidae, Bogidiellidae, Hadziidae, Typhlogammaridae, Crangonyctidae, Talitridae, etc.) have been established, including nearly 30 members of the genus Niphargus Schiödte, 1849 (G. Karaman, 2010); over half of known Niphargus taxa in Bosnia and Herzegovina are endemic for this and adjacent regions. During the recent investigations of caves in Bosnia and Herzegovina, organized by the speleological Society Mijatovi Dvori in Tomislavgrad, in association with the speleologists from Bosnia and Herzegovina, Croatia, Montenegro and other countries, various samples of Amphipoda have been collected and gave me at disposition for study. During the study of this material, one new species of the family NIphargidae (Amphipoda Gammaridea) is established and described here. The existence of over 300 known taxa of genus Niphargus from Europe, Asia Minor and Near East request more and more detailed descriptions of each species to make possible recognition of each species and its variability, and eventual comparison of morphological characters with results of genetic investigations of these species.

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NEW SPECIES OF THE FAMILY NIPHARGIDAE FROM BOSNIA AND HERZEGOVINA

Material and Methods

The sample was collected in the subterranean water by hand-net and catching by hands. The collected specimens were preserved in 70% ethanol. Specimens were examined and dissected using a Wild M 20 stereomicroscope and drawn using a camera lucida attachment. The animals were temporarily mounted in the mixture of glycerin and water for dissection and drawing of body parts, and later transferred on slides with liquid of Faure for final preservation. The body length of examined specimens was measured by tracing individual’s mid-trunk lengths (tip of the rostrum to end of telson) and drawings were made using a camera lucida and inked manually. Some morphological terminology and setae formulae follow G. Karaman’s terminology (Karaman, G., 1969; 1970; 2012b) for palpus article 3 of mandible [A-setae= setae on outer face; B-setae= setae on inner face of article; D-setae= short setae along ventral lateral margin of article; F-setae= long setae at distal part of article], and distal corner of propodus of gnathopods 1 and 2 [S-spine= corner palmar spine on outer face; L-spines= slender serrate spines sitting near S-spine on outer face; M-setae= row of long facial submarginal setae on outer face; R-spine= subcorner spine on inner face]. This study is based on the morphological, ecological and zoogeographical data.

Taxonomical part

Family NIPHARGIDAE

Niphargus bukovicensis sp. n. Figures 1-9

Material examined: Bosnia and Herzegovina: S-7234- Velika Bukovička špilja Cave [Velika Bukovača Cave] near Bukovica village (Tomislavgrad reg.), coordinates: X=4837278, y= 6434669, z= 987] including source going out of the cave, 5.9.2014, many exp. (leg. Gordan Karaman & Božana Karaman).

Diagnosis Body very slender, with short antennae and legs, and with long uropod 3. Epimeral plates 1 and 2 with almost subrounded ventroposterior corner, epimeral plate 3 with pointed corner and concave posterior margin. Maxilla 1 inner plate with 2-3 setae, outer plate with 7 spines (most of them with one lateral tooth each); Inner plate of maxilliped short, with 3 distal spines. Antenna 2 flagellum longer than last peduncular article. Coxae 1-4 short, coxa 1 not produced, coxa 4 unlobed. Propodus of gnathopods 1-2 smaller than corresponding coxae, trapezoid, its dactylus with row of setae along outer margin. Dactylus of pereopods 2-5 stout, with one strong spine at inner margin. Dactylus of pereopod 6 with 1-2 spines at inner margin; dactylus of pereopod 7 in larger specimens with 2-3, rarely only one spine at inner margin. Pereopods 5-7 with article 2 longer than broad, without ventroposterior lobe. Pleopods with 2 retinacula, peduncle of pleopods 1-3 scarcely setose. Urosomal segment 1 at each dorsolateral side with seta, urosomal segment 2 on each dorsolateral side with setae or spines. Uropod 3 in males with elongated peduncle and elongated inner ramus; outer ramus 2-articulated, second article in males long, in females shortened, but always much longer than diameter of article itself. Telson slightly longer than broad, incised nearly 2/3 of telson-length, each lobe with distal, lateral and facial spines.

Description. Male 20.0 mm (holotype): Body very slender, metasomal segments 1-3 with 3-4 short dorsoposterior marginal setae each (fig. 1G). Epimeral plates 1 and 2 with subrounded ventroposterior corner and convex posterior margin bearing 6-7 short setae each (fig. 1G). Epimeral plate 3 with strongly acute ventroposterior corner and sinusoid posterior margin bearing 5-6 short setae (fig. 1G). Epimeral plate 2 with 3 subventral spines, epimeral plate 3 with 2 subventral spines.

KARAMAN

Figure 1. Niphargus bukovicensis, sp. n., Velika Bukovička špilja Cave, male 20.0 mm: A= antenna 1; B= accessory flagellum; C= aesthetasc; D= antenna 2; E= head; F= mandible palpus, inner face; G= epimeral plates 1-3.

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NEW SPECIES OF THE FAMILY NIPHARGIDAE FROM BOSNIA AND HERZEGOVINA

Figure 2. Niphargus bukovicensis, sp. n., Velika Bukovička špilja Cave, male 20.0 mm: A-B= gnathopod 1, outer face; C= distal corner of gnathopod 1 propodus [S= corner spine; L= lateral spines; R= subcorner spine; M= facial M-setae]; D-E= gnathopod 2, outer face; F= distal corner of gnathopod 2 propodus [S= corner spine; L= lateral spines; R= subcorner spine].

KARAMAN

Figure 3. Niphargus bukovicensis, sp. n., Velika Bukovička špilja Cave, male 20.0 mm: A-B= pereopod 3; C-D= pereopod 4; E-labium; F= maxilliped; G= maxilla 2.

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NEW SPECIES OF THE FAMILY NIPHARGIDAE FROM BOSNIA AND HERZEGOVINA

Figure 4. Niphargus bukovicensis, sp. n., Velika Bukovička špilja Cave, male 20.0 mm: A-B= pereopod 5; C-D-F= left pereopod 6; E= dactylus of right pereopod 6; G-H-I= pereopod 7.

KARAMAN

Figure 5. Niphargus bukovicensis, sp. n., Velika Bukovička špilja Cave, male 20.0 mm: A= peduncle of pleopod 1; B= peduncle of pleopod 2; C= peduncle of pleopod 3; D= maxilla 1; E= urosome with uropods 1-2; F= left uropod 3; G= right uropod 3.

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NEW SPECIES OF THE FAMILY NIPHARGIDAE FROM BOSNIA AND HERZEGOVINA

Figure 6. Niphargus bukovicensis, sp. n., Velika Bukovička špilja Cave, male 20.0 mm: A= labrum; B= telson. Male 15.0 mm: C= antenna 2. Female 13.0 mm: D-E= pereopod 3; F-G= pereopod 4.

KARAMAN

Urosomal segment 1 on each dorsolateral side with 1 seta; urosomal segment 2 on each dorsolateral side with 2 setae; urosomal segment 3 naked (fig. 5E). Urosomal segment 1 at each ventroposterior corner with on weak short spine (fig. 5E). Head with short rostrum and short subrounded lateral cephalic lobes, ventroanterior excavation well developed eyes absent (fig. 1E). Antenna 1 shorter than half of the body (ratio: 8:20), peduncular articles 1-3 progressively shorter (ratio: 68:53:29), scarcely setose (fig. 1A); main flagellum scarcely setose, consisting of 26 articles (most of them with one short aesthetasc) (fig. 1C); accessory flagellum 2-articulated, much shorter than last peduncular article (fig. 1A, B). Antenna 2: peduncular article 3 short, with distoventral bunch of setae; peduncular articles 4 and 5 of equal length; article 4 at ventral margin with 3-4 bunches of setae (the longest setae exceeding the diameter of article itself (fig. 1D), at dorsal margin with 4 bunches of short setae; article 5 at ventral margin with 5 bunches of long setae (setae remarkably exceeding the diameter of article itself), at dorsal margin with several bunches of short setae; flagellum slender, with missing distal part, probably longer than last peduncular article (fig. 1D) (see sub Variability, fig. 6C). Antennal gland cone short (fig. 1D, 6C). Mouthparts well developed. Labrum broader than long, with convex distal margin (fig. 6A). Labium much broader than long, inner lobes well developed; outer lobes subrounded distally (fig. 3E). Mandibles: molar triturative, right mandible molar with 1 long lateral seta. Left mandible: incisor with 5 teeth and lacinia mobilis with 4 teeth and 6-7 rakers. Right mandible: incisor with 4 teeth and lacinia mobilis bifurcate, with several teeth, accompanied by 6-7 rakers. Mandibular palpus 3-articulated: first article naked (fig. 1F); second article hardly shorter than third article and provided with 10 setae (fig. 1F); third article subfalciform, with nearly 26 D-setae and 6 distal E-setae, on outer face is attached one bunch of 8 A-setae (fig. 1F), on inner face are attached 4 bunches of B-setae (2-2-1-1). Maxilla 1: inner plate with 3 distal setae (fig. 5D); outer plate with 7 spines [6 spines with one lateral tooth, one spine with 3 lateral teeth); palpus 2-articulated, almost reaching distal tip of outer plate spines, and provided with 5 distal setae (fig. 5D). Maxilla 2: both plates with distolateral setae only (fig. 3G). Maxilliped: inner plate short, with 3 distal pointed spines mixed with single setae (fig. 3F); outer plate reaching nearly half of palpus article 2 and provided with row of distomarginal lateromarginal spines; palpus article 3 at outer margin with one median and one distal bunch of setae; palpus article 4 at outer margin with 2 median setae, at inner margin with 2 setae near basis of the nail (fig. 3F), nail is much shorter than pedestal. Coxae 1-4 short. Coxa 1 broader than long (ratio: 54:37), with subrounded ventroanterior corner and bearing nearly 10 marginal setae (fig. 2A). Coxa 2 much broader than long (ratio: 59:47), with almost subrounded ventral margin and provided with nearly 10 marginal setae (fig. 2D). Coxa 3 broader than long (ratio: 63:51), with slightly convex ventral margin bearing nearly 8 short marginal setae (fig. 3A). Coxa 4 much broader than long (ratio: 65:49), central ventral margin almost straight, bearing nearly 8 marginal setae, ventroposterior lobe absent (fig. 3C). Coxae 5-7 short. Coxa 5 much broader than long (ratio: 80:41), anterior lobe as long as coxa 4, and provided with several marginal setae (fig. 4A). Coxa 6 smaller than coxa 5, bilobed, broader than long (ratio: 63:36), with several short setae on anterior and posterior lobe (fig. 4C). Coxa 7 entire, much broader than long (ratio: 62:28), with one posterior seta (fig. 4G). Gnathopods 1 and 2 relatively small, with propodus not larger than corresponding coxa (fig. 2A, D). Gnathopod 1: article 2 stout, along anterior margin with row of long setae, along posterior margin with bunches of long setae (fig. 2A); article 3 at posterior margin with one bunch of setae; article 5 shorter than propodus (ratio: 36:49), at anterior margin with one distal bunch of setae (fig. 2A). Propodus trapezoid, slightly longer than broad (ratio: 97:82), along posterior margin with 8 transverse rows of setae (fig. 2B); palm poorly convex, inclined nearly half of propodus-length, defined on outer face by one corner S-spine accompanied laterally by 2 L-spines and 3 facial long M-setae (fig. 2C), on outer face by one subcorner R- spine (fig. 2C). Dactylus reaching posterior margin of propodus, along outer margin with 9-11 single median setae, along inner margin with 7-9 short setae (fig. 2B). Gnathopod 2: article 2 along anterior and posterior margin with numerous long setae; article 3 at posterior margin with one bunch of setae; article 5 shorter than propodus (ratio: 40:50), along anterior margin with one median and one distal bunch of setae (fig. 2D). Propodus trapezoid, only slightly longer than broad (ratio: 98:90), along posterior margin with 9 transverse rows of setae (fig. 2E); palm slightly

Ecol. Mont., 6, 2016, 15-33 23

NEW SPECIES OF THE FAMILY NIPHARGIDAE FROM BOSNIA AND HERZEGOVINA convex, inclined hardly over half of propodus-length, defined on outer face by one corner S-spine accompanied laterally by 2 L-spines and 4 facial long M-setae, on inner face by one subcorner R-spine (fig. 2F). Dactylus reaching posterior margin of propodus, along outer margin with 10 single or paired median setae, along inner margin with 6-8 short setae (fig. 2E). Pereopods 3 and 4 relatively short. Pereopod 3: article 2 at anteroproximal margin with long setae, at anterodistal margin with short setae (fig. 3A), posterior margin with row of long setae. Articles 4-6 of unequal length (ratio: 56:40:45); article 4 at posterior margin with 5 bunches of setae (the longest setae exceeding the diameter if the article itself), along anterior margin with 3 bunches of setae; article 5 at posterior margin with 3 bunches of longer setae, along anterior margin with 2 bunches of short setae; article 6 at posterior margin with 5 bunches of short spines or single short setae, along anterior margin with 3 bunches of setae. Dactylus strong, much shorter than article 6 (ratio: 15:45), at inner margin with one strong spine and one short seta near basis of the nail (fig. 3B), along outer margin with one short median plumose seta; nail shorter than pedestal (ratio: 29:36). Pereopod 4: article 2 at anterodistal margin with row of short setae and at anteroproximal margin with several long setae (fig. 3C); posterior margin of article 2 with row of long setae; articles 4-6 of unequal length (ratio: 51:38:44); article 4 along posterior margin with 4 bunches of setae (the longest setae are nearly as long as diameter of the article), along anterior margin with 3 groups of setae; article 5 along posterior margin with 2 bunches of short setae mixed with 2 strong spines. Article 6 along posterior margin with 5 groups of short spines or setae, along anterior margin with 3 bunches of setae. Dactylus strong, much shorter than article 6 (ratio: 16:44), along inner margin with one strong spine and short seta near basis of the nail, along outer margin with one median plumose seta (fig. 3D); nail shorter than pedestal (ratio: 27:36) (fig. 3D). Pereopods 5-7 stout and relatively short, pereopod 7 reaching posterior margin of epimeral plate 2 only. Pereopod 5: article 2 rather longer than broad (ratio: 77:50), along anterior rather convex margin appears a row of 7 single or group of spine-like setae (fig. 4A), along posterior poorly concave margin are attached nearly 17 short setae, ventroposterior dilatation not forming the lobe (fig. 4A). Articles 4-6 of unequal length (ratio: 51:53:53); article 4 at anterior margin with 4 bunches of short setae, along posterior margin with 3 groups of single spines or setae; articles 5 and 6 along both margins with several bunches of short spines mixed with single short setae. Article 2 is longer than article 6 (ratio: 77:53). Dactylus is short and strong, much shorter than article 6 (ratio: 17:53), along inner margin with one strong spine and short seta near basis of the nail, along outer margin with one median plumose seta (fig. 4B); nail shorter than pedestal (ratio: 27:42). Pereopod 6 is remarkably longer than pereopod 5; article 2 much longer than broad (ratio: 88:50), along anterior slightly convex margin appears a row of 5 single spine-like setae and distal group of setae, along posterior almost straight or hardly concave margin appear nearly 13 short setae, ventroposterior dilatation not forming the lobe (fig. 4C). Articles 4-6 of unequal length (ratio: 60:75:75) (fig. 4C, D); article 4 at anterior margin with single or paired short setae, along posterior margin 3 groups of short spines; articles 5 and 6 along both margins with several bunches of short spines mixed with single short seta. Article 2 is longer than article 6 (ratio: 88:75). Dactylus much shorter than article 6 (ratio: 23:75), at inner margin with 1-2 strong spines and one short seta, at outer margin with one median plumose seta (fig. 4E, F). Nail shorter than pedestal: ratio: 27:50, dactylus with one spine, or: ratio: 33:56, dactylus with 2 spines. Pereopod 7: article 2 much longer than broad (ratio: 94:52), along anterior poorly convex margin with 5 groups of spine-like setae, along posterior margin with nearly 14 short setae, ventroposterior lobe absent (fig. 4G); articles 4-6 of unequal length (ratio: 56:75:82); article 4 at anterior margin with 3 groups of short setae, along posterior margin with 3 single spines; articles 5 and 6 along both margins with single or bunches of short spines mixed with single short setae (fig. 4H); article 2 is longer than article 6 (ratio: 94:82). Dactylus short and strong, much shorter than article 6 (ratio: 14:82), at inner margin with 4 spines, at outer margin with one median plumose seta (fig. 4 I); nail shorter than pedestal (ratio: 30:64). Pleopods 1-3 with 2 retinacula each. Peduncle of pleopod 1 at anterodistal margin with 4 setae (fig. 5A); peduncle of pleopods 2 with one anterodistal seta (fig. 5B); peduncle of pleopod 3 at posterior margin with 2 median setae (fig. 5C). Uropod 1 slender, peduncle without distal tubercle, bearing dorsoexternal row of spines and dorsointernal row of setae (except distal spine); inner ramus elongated, as long as peduncle, slightly curved distally, bearing one lateral and 3-4 distal short spines, as well as 5 bunches of simple short setae (fig. 5E);

KARAMAN

outer ramus shorter than half of inner ramus, bearing 4 distal short spines and 3 lateral bunches of short setae. Uropod 2 short, peduncle with dorsal spines (fig. 5E); inner ramus is poorly longer than outer ramus, both rami with single short lateral and distal spines (fig. 5E). Uropod 3 long, with elongated peduncle and inner ramus. Left uropod 3: peduncle much longer than broad (ratio: 54:18) bearing one lateral spine and lateral and distal short setae or spines (fig. 5F). Inner ramus slightly shorter than peduncle (ratio: 40:54), bearing several lateral spines and lateral and distal short simple setae (fig. 5F). Outer ramus 2-articulated, slender: first article poorly longer than second article (ratio: 133:120), along outer margin with 6 bunches of short simple setae, along inner margin with bunches of single spines and short simple setae accompanied with longer plumose setae (fig. 5F); second article along both margins and tip provided with bunches of simple setae. Right uropod 3: peduncle like that of left uropod 3. Inner ramus nearly as long as peduncle, bearing 3 lateral short spines and several lateral and distal short simple setae (fig. 5G). Outer ramus 2-articulated, first article slightly longer than second article (ratio:136:113), along inner ramus with 7 groups of short spines and 1-2 simple short setae, accompanied by single longer plumose setae (fig. 5G), along outer margin appear 5 groups of short simple setae; second article along both margins and tip with several groups of short simple setae. Telson slightly longer than broad (ratio: 79:70), incised nearly 2/3 of telson-length; lobes slightly tapering distally, provided with 3 short distal spines and 2 facial spines; along outer margin is attached one spine and 0-1 seta, along inner margin of each lobe appear 2-3 short spines; a pair of short plumose setae is attached near the middle of outer margin of each lobe (fig. 6B). Coxal gills are moderately large, ovoid, not exceeding corresponding leg article 2 (figs. 2D, 3A, C; 4A, C).

Female 13.0 mm with setose oostegites and 17 eggs (paratype): Body moderately slender, metasomal segments 1-3 at dorsoposterior margin with 3-5 dorsoposterior short setae (fig. 9B). Epimeral plates 1 and 2 with poorly marked ventroposterior corner defined by 1 strong spine-like seta, and by convex posterior margin bearing several short setae each. Epimeral plate 3 sharply pointed and produced, with remarkably inclined posterior margin bearing 4-5 short setae (fig. 9B). Epimeral plate 2 is provided with 2 subventral spines, epimeral plate 3 is provided with 3 subventral spines (fig. 9B). Urosomal segment 1 on each dorsolateral side with one seta, urosomal segment 2 on each dorsolateral side with 2 spines; urosomal segment 3 naked. Urosomal segment 1 on each posteroventral corner with one spine-like seta near basis of uropod 1 peduncle (fig. 9C). Antenna 1 reaching half of body-length, main flagellum consisting of 27-28 articles. Antenna 2: articles 4 and 5 like these in male, flagellum longer than last peduncular article and consisting of 13 articles. Mouthparts mainly like these in male. Mandible palpus article 2 with 10-12 setae, palpus article 3 with up to 26 D-setae and 5-6 distal E- setae, on outer face with row of 6-7 A-setae, on inner face with 6 B-setae (2-2-1-1). Maxilla 1 inner plate with 2-3 setae, outer plate with 7 spines (5 spines with 1 lateral tooth, one spine with 1-2 teeth, 1 spine with 3 lateral teeth, palpus nearly reaching tip of outer plate spines, bearing 5-6 distal setae. Maxilliped inner plate with 3 distal spines; palpus article 3 at outer margin with one median and one distal group of setae; palpus article 4 at inner margin with 2 setae near basis of the nail, along outer margin with 1-2 median setae. Coxae are rather longer than these in male. Coxa 1 is rather broader than long (ratio: 50:44), with subrounded ventroanterior corner and bearing nearly 14 short marginal setae (fig. 7A). Coxa 2 hardly longer than broad (ratio: 56:52), along ventral margin with nearly 12 marginal setae (fig. 7D). Coxa 3 is longer than broad (ratio: 63:56), at ventral margin with nearly 16 short setae (fig. 6D). Coxa 4 poorly longer than broad (ratio: 60:57), along ventral margin with nearly 13 short setae, ventroposterior lobe is not developed (fig. 6F). Coxa 5 is broader than long (ratio: 73:50), provided with several setae only, anterior lobe nearly as long as coxa 4 (fig. 8A). Coxa 6 remarkably smaller than coxa 5, broader than long (ratio: 53:39) (fig. 8C). Coxa 7 distinctly broader than long (ratio: 55:30), entire, with convex ventral margin (fig. 8E). Gnathopods 1 and 2 relatively small, with propodus poorly smaller than corresponding coxa (fig. 7A, D). Gnathopod 1: article 2 along anterior and posterior margin with numerous long setae; article 3 at

Ecol. Mont., 6, 2016, 15-33 25

NEW SPECIES OF THE FAMILY NIPHARGIDAE FROM BOSNIA AND HERZEGOVINA posterior margin with one bunch of setae. Article 5 shorter than propodus (ratio: 35:39), along anterior margin with 1-2 groups of setae (fig. 7A). Propodus trapezoid, hardly longer than broad (ratio: 80:85, or: 81:77), along posterior margin with 8 transverse groups of setae (fig. 7B, C). Palm poorly convex, inclined nearly half of propodus-length, defined on outer face by one corner S-spine accompanied by 2 L-spines and 4 long facial M-setae (fig. 7B, C), on inner face by one subcorner R-spine. Dactylus nearly reaching posterior margin of propodus, along outer margin with 5-9 long median setae, along inner margin with row of 5-9 short setae. Gnathopod 2: article 2 along anterior margin with row of nearly 10 shorter setae, along posterior margin with several bunches of long setae (fig. 7D); article 3 at posterior margin with one bunch of setae; article 5 nearly as long as propodus, along anterior margin with 2 bunches of setae. Propodus trapezoid, nearly as long as broad as or hardly broader than long, along posterior margin with 11 transverse groups of setae (fig. 7E). Palm slightly convex, inclined nearly half of propodus-length, defined on outer face by 1 S spine accompanied laterally by 2 L-spines and 4 long facial M-setae (fig. 7E), on inner face by one R-spine. Dactylus reaching posterior margin of propodus, along outer margin with row of 9 long setae, along inner margin with row of short setae. Pereopods 3 and 4 moderately setose. Pereopod 3: anteroproximal margin of article 2 with several long setae, at anterodistal margin with row of short setae, along posterior margin with numerous long setae (fig. 6D). Articles 4-6 of unequal length (ratio: 50:40:41); article 4 at posterior margin with 3 bunches of setae (the longest setae exceeding the diameter of article itself), along anterior margin with 3 bunches of shorter setae. Article 5 at posterior margin with one median spine and 3 bunches of setae (fig. 6D); article 6 along posterior margin with 5 single or paired short spines and single short seta, along anterior margin with 3 bunches of short setae. Dactylus much shorter than article 6 (ratio: 17:41), along inner margin with one spine and short seta near basis of the nail, along outer margin with one median plumose seta (fig. 6E); nail is shorter than pedestal (ratio: 24:30). Pereopod 4: article 2 at anterodistal margin with row of shorter setae and at anteroproximal margin with several long setae, and with numerous long setae along posterior margin (fig. 6F); articles 4-6 of unequal length (ratio: 45:34:41); article 2 at posterior margin with 3 bunches of setae as long as or poorly longer than diameter of article itself, along anterior margin with 3 groups of setae; article 5 at posterior margin with 2 spines and several short setae; article 6 along posterior margin with 4-5 groups of short setae or single spines. Dactylus much shorter than article 6 (ratio: 13:41), along inner margin with one strong spine and short seta near the basis of the nail, at outer margin with one median plumose seta (fig. 6G); nail shorter than pedestal (ratio. 24:26). Pereopod 5 remarkably shorter than pereopods 6 and 7, with article 2 longer than broad (ratio: 69:42), along anterior margin with 5 groups of setae, along posterior margin with nearly 18 short setae, ventroposterior dilatation not forming the lobe (fig. 8A). Articles 4-6 of unequal length (ratio: 45:49:57); article 4 at anterior margin with 4 groups of setae, along posterior margin with 3 spines; article 5 at both margins with groups of spines and setae; article 6 along anterior margin with 4 bunches of short spines and single setae, along posterior margin with setae only. Article 2 is slightly longer than article 6 (ratio: 69:57). Dactylus much shorter than article 6 (ratio: 17:57), along inner margin with 1 spine and short seta, at outer margin with one median plumose seta (fig. 8B); nail shorter than pedestal (ratio: 27:36). Pereopod 6: article 2 much longer than broad (ratio: 85:46), along anterior poorly convex margin with 6 groups of spine-like setae, along posterior almost straight margin with nearly 15 short setae, ventroposterior dilatation not forming the lobe (fig. 8C). Articles 4-6 of unequal length (ratio: 60:70:83); article 4 at anterior margin with 4 groups of short setae, along posterior margin with 4 groups of spines and one long seta; article 5 and article 6 along both margins with groups of short spines accompanied sometimes with setae. Article 2 is almost as long as article 6. Dactylus much shorter than article 6 (ratio: 22:83), along inner margin with 1 spine and 1 short seta, at outer margin with 1 median plumose seta (fig. 8D); nail is shorter than pedestal (ratio: 28:53). Pereopod 7: article 2 much longer than broad (ratio: 90:49), along anterior poorly convex margin with 7-8 groups of spine-like setae, along posterior almost straight margin, with nearly 14 short setae ventroposterior dilatation not fully lobed (fig. 8E), Articles 4-6 of unequal length (ratio: 56:67:90); article 4 at anterior margin with 3 groups of setae, along posterior margin with 3 groups of spines; articles 5 and 6 along both margins with groups of spines; article 2 is nearly as long as article 6 (fig. 8F). Dactylus much shorter than article 6 (ratio: 36:60), at inner margin with 2 spines (fig. 8G), or: with ratio: 28:60, and at inner margin with 3 spines and one seta (fig. 8H), at outer margin with one median plumose seta.

KARAMAN

Figure 7. Niphargus bukovicensis, sp. n., Velika Bukovička špilja Cave, female 13.0 mm: A-B= left gnathopod 1, outer face; C= right gnathopod 1 propodus, outer face; D-E= gnathopod 2, outer face.

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NEW SPECIES OF THE FAMILY NIPHARGIDAE FROM BOSNIA AND HERZEGOVINA

Figure 8. Niphargus bukovicensis, sp. n., Velika Bukovička špilja Cave, female 13.0 mm: A-B= pereopod 5; C-D= pereopod 6; E-F-G= left pereopod 7; H= dactylus of right pereopod 7.

KARAMAN

Figure 9. Niphargus bukovicensis, sp. n., Velika Bukovička špilja Cave, female 13.0 mm: A= telson; B= epimeral plates 1-3; C= uropod 1; D= uropod 2; E= uropod 3. Female 12.0 mm: F= gnathopod 1 propodus, outer face; G= gnathopod 2 propodus, outer face.

Ecol. Mont., 6, 2016, 15-33 29

NEW SPECIES OF THE FAMILY NIPHARGIDAE FROM BOSNIA AND HERZEGOVINA

Pleopods 1-3 with 2 retinacula each. Peduncle of pleopods 1 at anterior margin with 3-4 setae; peduncle of pleopods 2 at anterior margin with 1 seta; peduncle of pleopod 3 with one seta at posterior margin. Uropod 1: peduncle with dorsoexternal row of spines and dorsointernal row of setae (fig. 9C); inner ramus slightly shorter than peduncle, bearing 2 lateral and 4-5 distal short spines, accompanied by 2 bunches of simple setae (fig. 9C); outer ramus slightly shorter than inner ramus, bearing 3 lateral and 4-5 distal short spines, accompanied by 2 lateral bunches of simple setae. Uropod 2: peduncle with dorsal spines; inner ramus is only slightly longer than outer ramus and provided with 3 lateral and 4 distal short spines (fig. 9D); on outer ramus appear 3 lateral and 4-5 distal short strong spines (fig. 9D). Uropod 3: peduncle longer than broad (ratio: 46:22), bearing 2 lateral and several distal short spines and short setae (fig. 9E); inner ramus much shorter than peduncle (ratio: 46:18), scale-like, with lateral spine and simple seta and with distal spine and several simple setae (fig. 9E); outer ramus 2-articulated, first article along outer ramus with 4 groups of short spines and single simple setae, along inner margin with 5 groups of short spines and single short simple setae accompanied by single long plumose setae; second article much shorter than first article (ratio: 49:118), along both margins and tip with groups of simple setae (fig. 9E). Telson rather longer than broad (ratio: 82:68), incised nearly 2/3 of telson-length; each lobe with 3-4 distal and 2-3 facial spines; along inner margin of lobes appear 1-2 spines; along outer margin of each lobe are attached 0-1 spine (fig. 9A); a pair of short plumose setae is attached near the middle of outer margin on each lobe. Coxal gills like these in male, not exceeding ventral tip of corresponding article 2 (figs. 6D, F; 7D; 8A C). Oostegites very large, bearing marginal setae (figs. 6D, F; 7D; 8A).

Variability. In our sample, the adult females with setose oostegites were 11.0-14.0 mm long, and males were up to 20.0 mm long. Inner plate of maxilla 1 bearing 2-3 setae, 6 spines of outer plate in maxilla 1 usually with one lateral tooth, rarely one spine is provided with 2 lateral spines, inner spine is always with several lateral teeth. Along inner margin of dactylus on pereopods 3-5 appears always one spine near basis of the nail, in males and females. Dactylus of pereopod 6 is mainly provided at inner margin with one spine, sometimes with 2 spines (male 12.2 mm; female ovig. 13.0 mm). Dactylus of pereopod 7 is in females usually with only one spine near basis of the nail (females ovig. 10-14 mm), or with 2 spines (females ovig. 10.0-12.0 mm). In female of 13.5 mm left pereopod 7 dactylus was with 2 spines and right pereopod 7 dactylus with 3 spines. In male 15.0 mm antenna 2 exceeding the length of last peduncular article and consisting of 13 articles (fig. 6C). All collected males have 2-3 spines at inner margin of pereopod 7 dactylus (males 12.2-20.0 mm). Evidently the males are more inclined to have more spines on dactylus of pereopod 7 than the females, but the increasing number of these spines is not in direct correlation with the size of the specimens. Propodus of gnathopods 1-2 is hardly narrower in female of 12.0 mm: gnathopod 1 propodus is longer than broad (ratio: 87: 76) (fig. 9F), gnathopod 2 propodus is longer than broad also (ratio: 92:85) (fig. 9G). In adult males is inner ramus of uropod 3 elongated, reaching nearly the size of peduncle. Similar shape of uropod 1 is present in Niphargus karamani Schellenberg, 1935 known from Croatian- Slovenian border [loc. typ.: well near Miljana castle (on riverbank of Sutla river, Sava tributary), Croatia [S. Karaman, 1950; G. Karaman, 2014).

Holotype: Male 20.0 mm. Holotype is preserved in Karaman`s Collection in Podgorica, Montenegro

Locus typicus: Velika Bukovička špilja Cave, Bukovica village, Tomislavgrad region, Bosnia and Herzegovina.

Distribution: Known from type locality only.

KARAMAN

Derivatio nominis. The name “bukovicensis” arrives from the name of the locality Bukovica village, where the species was collected.

Remarks and affinities

Niphargus bukovicensis is characterized by presence of additional spines on dactylus of pereopods 6 and 7. The presence of additional spines on dactylus of various pereopods is present in various taxa of genus Niphargus: in some cases these presence of additional spines can be more occasional than regular (Niphargus navotinus G. Karaman, 2014a) or always occasional (Niphargus longicaudatus Costa, 1851), or can be more or less regular (N. stankoi G. Karaman, 1974, N. radzai G. Karaman, 2014a, N. castellanus S. Karaman, 1960, etc.). Based on the presence of additional spines on dactylus of some pereopods, N. bukovicensis is rather similar to the members of the Supraniphargus Group of species (S. Karaman, 1950), characterized by elevated number of spines on some or all pereopods. On the other hands, the existence in our sample of adult specimens with and without additional spines on dactylus of pereopods 6 and 7, show the transitive position of Supraniphargus group regarding some other groups of Niphargus taxa. Niphargus illidzensis illidzensis Schäferna 1922 [loc. typ.: Ilidža spring in Sarajevo, Bosnia and Herzegovina], and additional taxa: dalmatinus Schäferna 1922 [loc. typ.: spring Biba at coast of Vransko jezero Lake near Šibenik, Dalmatia), pannonicus S. Karaman, 1950b [loc. typ.: well in Slavonia, Croatia] and montenigrinus G. Karaman 1962 [loc. typ.: Źabljak, Montenegro] differs from N. bukovicensis by presence of additional spines on all pereopods and by various combination of other taxonomical characters). Niphargus zavalanus S. Karaman 1950c (loc. typ.: Zavala, Herzegovina) bearing pereopod 3-5 with one spine, pereopod 6 usually also with one, rarely 2 spines, only on P7 with 2-3, rarely 4 spines. But it differs from our species by short uropod 3, narrow and poorly spinose telson, etc. Niphargus castellanus S. Karaman 1960 [loc. typ.: spring in Blace above Kaštel Stari near Split, Croatia] has elevated number of spines on pereopods 3-4, and only one spine on dactylus of pereopods 5-7, short and broad article 2 of pereopods 5-7, less spinose telson, etc. Niphargus radzai G. Karaman, 2014a [loc. typ.: Spring above “P.D. Sv. Jakov”, Ravno Vrdovo, Dinara Mt., Croatia] has elevated number of spines on dactylus of pereopods 3-4 only, subrounded epimeral plates 1-3 etc. N. slovenicus S. Karaman, 1950b [loc. typ.: Stražišče near Kranj, Slovenia], differs by pereopods 3-5 provided with 2-4 spines, but dactylus of pereopods 6-7 is provided with 1 spine, etc. The species N. smederevanus S. Karaman, 1950a [loc. typ.: Smederevo, Serbia] is close to N. bukovicensis by many characters (maxilla 1, maxilliped, spinose telson, dactylus of gnathopods 1-2, etc.), but differs from later by presence of additional spines on dactylus of pereopods 3 and 4 only, etc. In the central Europe is present Niphargus tatrensis Wrzesniowski 1890- Complex of taxa [loc. typ.: Zakopane, Poland], characterized also by presence of some additional spines of some of pereopod dactyl. Various forms and taxa of this complex have been described; most of them probably are a distinct taxa. This Complex of taxa is still poorly known and its relation to the western Balkan taxa and to N. illidzensis Complex is poorly known and needs detailed studies. Near this complex is also Niphargus stankoi G. Karaman, 1974 known from Macedonia [loc. typ.: Bukovo], but this species differs remarkably from N. bukovicensis by elevated number of spines on dactylus of pereopods 3-7, by uropod 1, maxilla 1, etc. Further studies on all members of species with additional spines on dactylus of pereopods will put more light on the taxonomical value of this taxonomical character in various members of Supraniphargus Complex and its relation to other Niphargus taxa.

Conclusions

The discovery of new species Niphargus bukovicensis, sp. n. in the Velika Bukovička špilja Cave in Bosnia and Herzegovina show the existence of high number of endemic Niphargus taxa in the subterranean waters of Bosnia and Herzegovina and its connections to the taxa of adjacent regions, thanks to the geological,

Ecol. Mont., 6, 2016, 15-33 31

NEW SPECIES OF THE FAMILY NIPHARGIDAE FROM BOSNIA AND HERZEGOVINA ecological, zoogeographical and other events in the past on this territory, and the present condition of this region. The members of the family Niphargidae are still in the process of speciation and splitting into numerous more or less distinct populations. Among them, some of these populations developed complete reproductive isolation (genetic or geographic) forming a distinct species. On the other hands, many populations are still in the process of isolation and transition, still in some manner connected together. In this light, to fix the boundaries between single taxa is very difficult and undergoes often towards subjective criteria. Neither pure morphological studies, nor pure genetical studies can resolve alone this problem; it is necessary to use of combination of various methods to resolve this problem, leaving still questionable results. We don`t know the real value of character appearance of additional spines on dactylus, as a character used to separate the single taxa, but in any case, this character must be present in the genetic code of these populations, and maybe next deep genetical and molecular studies will recognize it as a distinct character. The number of splitting populations into a distinct taxa will depend on how deep we will go in establishing of differences among them, and of our decisions where to put the border under which the further splitting of a distinct taxa begin to be useless (G. Karaman 2012a: 33).

Acknowledgements I am indebted to Mg. Roman Ozimec, biologist and speleologist from Zagreb, and other speleologists, who helped us to visit the Velika Bukovička špilja Cave near village Bukovica during the International Speleo-diving Expedition Ponor Kovači –Ričina Spring 2014, organized by Speleological Society Mijatovi Dvori from Tomislavgrad, and collect the material used in this study.Many thanks to the Montenegrin academy of sciences and arts in Podgorica for help in realization of this work.

References

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