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: karaman@t-com.me
Received 10 March 2016 Accepted 24 April 2016 Published online 26 April 2016.

Abstract
From the subterranean waters of the Velika Bukovika pilja 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 Schidte, 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
Schidte, 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 individuals 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. Karamans 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 Bukovika pilja Cave [Velika Bukovaa
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 & Boana 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.

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Figure 1. Niphargus bukovicensis, sp. n., Velika Bukovika pilja 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 Bukovika pilja 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].
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Figure 3. Niphargus bukovicensis, sp. n., Velika Bukovika pilja 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 Bukovika pilja 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.
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KARAMAN

Figure 5. Niphargus bukovicensis, sp. n., Velika Bukovika pilja 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 Bukovika pilja 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.

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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 Rspine (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
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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);
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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 Esetae, 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
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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.
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Figure 7. Niphargus bukovicensis, sp. n., Velika Bukovika pilja 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.

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

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NEW SPECIES OF THE FAMILY NIPHARGIDAE FROM BOSNIA AND HERZEGOVINA

Figure 8. Niphargus bukovicensis, sp. n., Velika Bukovika pilja 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.

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KARAMAN

Figure 9. Niphargus bukovicensis, sp. n., Velika Bukovika pilja 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.
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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 Bukovika pilja Cave, Bukovica village, Tomislavgrad region, Bosnia and
Herzegovina.
Distribution: Known from type locality only.

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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 Schferna 1922 [loc. typ.: Ilida spring in Sarajevo, Bosnia and
Herzegovina], and additional taxa: dalmatinus Schferna 1922 [loc. typ.: spring Biba at coast of Vransko
jezero Lake near ibenik, Dalmatia), pannonicus S. Karaman, 1950b [loc. typ.: well in Slavonia, Croatia]
and montenigrinus G. Karaman 1962 [loc. typ.: abljak, 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 Katel 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.: Straie 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 Bukovika pilja 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,

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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 Bukovika pilja Cave near village Bukovica during the International Speleo-diving Expedition Ponor
Kovai Riina 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.

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