European Journal of Taxonomy

European Journal of Taxonomy 205: 1-23
http://dx.doi.org/10.5852/ejt.2016.205

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This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773
www. europeanj ournaloftaxonomy. eu
2016 • Antic D.Z. et al.

Research article

urn:lsid:zoobank.org:pub:B24272FE-1876-40FE-8029-6488D5A31F4E

Taxonomic status of the family Biokoviellidae Mrsic, 1992
(Diplopoda, Chordeumatida): reconsideration,
with a description of one new species

Dragan Z. ANTIC 1 *, Tvrtko DRAZINA 2 3 , Tonci RADA 4 ,

Luka R. LUCIC 5 & Slobodan E. MAKAROV 6

1,5,6 Institute of Zoology, University of Belgrade - Faculty of Biology,

Studentski trg 3, 11000 Belgrade, Serbia.

2 Division of Zoology, Department of Biology, Faculty of Science, University of Zagreb,

Rooseveltov trg 6, 10000 Zagreb, Croatia.

3 Croatian Biospeleological Society, Demetrova 1, 10000 Zagreb, Croatia.

4 Speleological Society “Spiljar”, 21000 Split, Croatia.

* Corresponding author: dragan.antic@bio.bg.ac.rs
2,3 Email: tdrazina@gmail com
4 Email: tonci.radja@gmail.com
5 Email: luka@bio.bg.ac.rs
6 Email: slobodan@bio.bg.ac.rs

1 um:lsid:zoobank.org:author:A3774DC0-0710-4F91-8F36-F18620D694E0

2,3 um:lsid:zoobank.org:author:47C87F36-806F-4F6C-950D-C9ElD48CC747

4 urn:lsid:zoobank.org: author: 06AB80C6-0EBA-4201-96A1-3BA64EE3F717

5 urn:lsid:zoobank.org:author:D5F49D5B-6BCC-42B4-A000-BE23920C811F

6 urn:lsid:zoobank.org:author:D2745110-BE94-4FBF-9D36-5ACC9045B3C3

Abstract. A new troglobitic species of the previously monotypic genus Biokoviella Mrsic, 1992,
B. mosorensis Antic & Drazina sp. nov., is described from caves on Mt. Mosor, Croatia. In addition
to this, B. mauriesi Mrsic, 1992, is partially re-described, and the taxonomic status of the family
Biokoviellidae is re-considered. The genus Biokoviella is placed in the subfamily Biolcoviellinae Mrsic,
1992 stat. nov. within the family Anthogonidae Ribaut, 1913. The relationship of the genus Biokoviella
with other anthogonids is briefly discussed, and a distribution map of the genus is presented. Notes on
ecology and coinhabitants of the genus Biokoviella, and new data on some Balkan anthogonids are also
included in the paper.

Keywords. Biokoviella , new species, caves, Croatia, Balkan Peninsula.

Antic D.Z., Drazina T., Rada T., Lucic L.R. & Makarov S.E. 2016. Taxonomic status of the family Biokoviellidae
Mrsic, 1992 (Diplopoda, Chordeumatida): reconsideration, with a description of one new species. European
Journal of Taxonomy 205: 1-23. http://dx.doi.org/10.5852/eit.2016.2Q5

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European Journal of Taxonomy 205 : 1-23 ( 2016 )

Introduction

A little more than two decades ago, Mrsic (1992) described the new monotypic genus Biokoviella
Mrsic, 1992 from caves on Mt. Biokovo in Croatia. For this taxon, he established the new monotypic
family Biokoviellidae, indicated similarities with representatives of the families Chamaesomatidae and
Anthogonidae, and placed the new family in the superfamily Cleidogonoidea sensu Mauries (1982).
He retained this opinion in his paper from 1994 (Mrsic 1994: 249). After that, in Shear’s paper from
2000 on the higher classification of Chordeumatida, the family Biokoviellidae was not assigned to any
superfamily, not being mentioned at all; however, it was included by Shelley et al. (2000) and Shelley
(2002) in the superfamily Cleidogonoidea sensu Shear (2000), a somewhat extended superfamily
concept based on Hoffman’s classification from 1980, but significantly different from that of Mauries
(1982). The indicated classification of Cleidogonoidea as given by Shelley (2002) was followed by
Shear (2011). In Chapter 16 of a recently published taxonomic overview of Diplopoda by Enghoff et al.
(2015), the family Biokoviellidae is likewise placed in the superfamily Cleidogonoidea sensu Shear
(2000) and Shelley (2002).

In our opinion, great similarities exist between some of the representatives of the family Anthogonidae
sensu Antic et al. (2015) and the family Biokoviellidae (see Discussion). After examining a topotype
male of B. mauriesi Mrsic, 1992, as well as a new Biokoviella species, we conclude that the family
Biokoviellidae deserves subfamily level only, and at the family level we synonymize this taxon
with the family Anthogonidae. In this rearrangement, the family Anthogonidae should be placed in
the superfamily Anthroleucosomatoidea, together with Anthroleucosomatidae and Vandeleumatidae
according to Enghoff et al. (2015). These authors included the family Acherosomatidae, together with
Anthroleucosomatidae and Vandeleumatidae, in Anthroleucosomatoidea. This chapter was probably
prepared before publication of the paper by Antic et al. (2015), where we included the subfamily
Acherosomatinae in the family Anthogonidae. In the present paper, we describe one new Biokoviella
species from Mt. Mosor in Croatia.

Materials and methods

Specimens preserved in 70% ethanol were examined under a Carl Zeiss Jena Technival 2 binocular
stereo microscope and a Carl Zeiss Stemi 2000-c binocular stereo microscope with an AxioCamMRc
camera. All taxonomically important structures were dissected and mounted in glycerin for temporary
microscope preparations and observed under a Carl Zeiss Axioscope 40 microscope. Drawings of
gonopods were executed using a computer monitor and pictures made with a Canon PowerShot A80
digital camera connected to an Axioscope 40 microscope. Some relevant structures were investigated
with a JEOL JSM-6460LV scanning electron microscope (University Center for Electron Microscopy,
Department of Biology and Ecology, University of Novi Sad, Serbia).

The distribution map was created using QGIS version 2.12.0 and Adobe Photoshop CS6. The final
images were processed with Adobe Photoshop CS6. The coordinates given in the location data are based
on WGS84.

Descriptions of the new species follow a pattern proposed for Chordeumatida by Spelda (2001). The
terminology used to denote the different parts of gonopods follows the pattern proposed by Mrsic (1992),
with some modifications.

Abbreviations for morphological terms used in descriptions

a = anterior coxal process

al, a2 = branches of anterior coxal process

b = bristle apparatus

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ANTIC D.Z. et al.. Taxonomic status of Biolcoviellidae

bu

CIX

f

1

lv

m

MA

MIX

mv

op

P

PIX

s

sp

ss

t

ts

V

bursa

(macrochaetal index) distance between exterior and median macrochaetae/distance between

interior and median macrochaetae

pseudoflagellum

rounded lobe

lateral valve

medial process

(macrochaetal angle) angle between the arm created by the median and exterior macrochaetae
and the arm formed by the median and interior macrochaetae

(median index) distance between interior macrochaetae and axial suture/distance
between interior and median macrochaetae
mesal valve
operculum

posterior coxal process

(paratergal index) (width of metazonite - width of prozonite)/(2 x length of paratergum)

sternal plate

lateral sternal process

shield-like structure of lateral valve

telopodal remnant

tooth-like structure of lateral valve

syncoxal vesicle

Other abbreviations used

CBSS

IZB

NHMSC

SEM

Croatian Biospeleological Society, Zagreb, Croatia

Institute of Zoology, University of Belgrade - Faculty of Biology, Belgrade, Serbia
Natural History Museum, Split, Croatia
scanning electron microscope

Results

Taxonomy

Class Diplopoda Blainville-Gervais, 1844
Order Chordeumatida Pocock, 1894

Family Anthogonidae Ribaut, 1913

Anthroleucosomatidae Verhoeff, 1899: 130. (pro parte)
Acherosomatinae Verhoeff, 1930: 4.

Haasiinae Hoffman, 1980: 137.

Biokoviellidae Mrsic, 1992: 53. syn. nov.

Subfamily Biokoviellinae stat. nov.

Nomen translatum ex family Biokoviellidae Mrsic, 1992

Genus Biokoviella Mrsic, 1992

This genus was established by Mrsic (1992) for the type and until now only species, B. mauriesi (from
caves and pits on Mt. Biokovo, Dalmatia, Croatia). The name is regarded as a feminine noun.

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European Journal of Taxonomy 205 : 1-23 ( 2016 )

Type species

Biokoviella mauriesi Mrsic, 1992, by monotypy.

Diagnosis

This genus clearly differs from all other anthogonids in the presence of one pair of bristle apparatuses
in combination with one pair of bristle-clad pseudoflagella on the anterior gonopods, while the posterior
gonopods consist of coxal vesicles fused in the form of a medial process and telopodal remnants; without
lateral coxal processes.

Etymology

The genus is named after its type locality, Mt. Biokovo.

Included species

Biokoviella mauriesi Mrsic, 1992
Biokoviella mosorensis Antic & Drazina sp. nov.

Biokoviella mauriesi Mrsic, 1992
Figs 1-3, 9A, 10A

Diagnosis

Differs from B. mosorensis sp. nov. in larger body size, anterior gonopods that are more robust and
several other details of their structure. Tateral valve of vulva with tooth-like subtriangular structures
posteriorly.

Etymology

The species was named in honor of the well-known world expert in diplopodology Dr. Jean-Paul Mauries.

Material examined (total: 13 c?c?, 7 55, 9 juveniles)

Topotype

CROATIA: StaraLedenica Cave, 43.318° N, 17.050° E, 1350 m a.s.l., Biokovo Mountain, Dalmatia,

16 Jun. 2002, leg. R. Ozimec (IZB).

Other material (total: 12 $$, 7 $$, 9 juveniles)

CROATIA, Dalmatia, Mt. Biokovo: 4 Jamakraj Stare Skole Pit, Tokva, 12. Jul. 1989, leg. B. Jalzic
(CBSS); 2 SS, Jama Cavlenovaca Pit, 18 Jun. 2002, leg. R. Ozimec (CBSS); 2 SS, 2 $?, 2 juveniles,
Jama pod Sv. Jurom Pit, below Sv. Jure Peak, 20 Jun. 2002, leg. R. Ozimec (CBSS); 1 §, Kasogijeva
Jama Pit, below Sv. Jure Peak, 21 Jul. 2002, leg. M. Tukic (CBSS); 1 §, 1 juvenile, Lovricia Jama 2
Pit, 26 Jul. 2002, leg. J. Bedek (CBSS); 1 1 juvenile, Jezero Tedenica Cave, Strazbenica, Kaoci,

20 Jul. 2003, leg. R. Ozimec (CBSS); 1 juvenile, Ted 1 Pit, below Sv. Jure Peak, 21 Jul. 2003, leg.
J. Bedek (CBSS); 1 <$, Crna Tedenica Cave, below Sv. Jure Peak, 24 Oct. 2006, leg. M. Tukic (CBSS); 1
5, same data except 18. Jun. 2011, leg. A. Kirin (CBSS); 1 S, 1 §, 3 juveniles, Velika Macka Pit, Velilca
Macka, 22 Jun. 2008, leg. M. Tukic (CBSS); 1 juvenile, same data except leg. S. Nizetic (CBSS); 1
1 5, Tovricia 3 Pit, Tipi Docic, 30 May 2015, leg. T. Delic (IZB).

Type locality

Stara Tedenica Cave, Mt. Biokovo, Dalmatia, Croatia.

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ANTIC D.Z. etal.. Taxonomic status of Biokoviellidae

Fig. 1 . Biokoviella mauriesi Mrsic, 1992, S, topotype, pleurotergite 15. A. Posterior view. B. Prozonite,
dorsal view. C. Metazonite, dorsal view. D. Lateral view. E. Dorsal view.

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European Journal of Taxonomy 205 : 1-23 ( 2016 )

Redescription
Body indices

Pleurotergite 15 (Fig. 1). CIX ~ 0.8; MIX ~ 0.8; PIX = 0.25; MA~ 160°.

Fig. 2. Biokoviella mauriesi Mrsic, 1992, topotype, gonopods. A. Anterior gonopods, anterior
view. B. Anterior gonopods, lateral view. C-D. Anterior and posterior gonopods, postero-distal view.
E. Anterior gonopods, postero-distal view of the medial part. F. Anterior gonopods, lateral view of the
posterior process and bristle apparatus. Abbreviations: see Material and methods.

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ANTIC D.Z. et al ., Taxonomic status of Biokoviellidae

Redescription of male sexual characters

Male sexual characters. Teg pairs 3-7 are somewhat larger than the rest of the legs, leg pairs 3 and 4
the most robust; without any peculiarities (see Discussion).

Anteriorgonopods (Fig. 2). Anterior coxal processes (a) [= angiocoxites sensuyhsit (1992)] completely
separated from each other, rising from the wide subquadrangular sternal plate (s); first A (base) very
broad, remaining 2 A very slender and long, curved caudad, apically with two branches (al and a2)

Fig. 3. Biokoviella mauriesi Mrsic, 1992, $ from the Tovricia 3 Pit, vulvae. A. Posterior view.
B. Tateral view. C. Tatero-distal view. D. Anterior view. E. Antero-distal view. F. Distal view.
Abbreviations: see Material and methods.

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European Journal of Taxonomy 205 : 1-23 ( 2016 )

opposite to each other. Posteriorly, anterior coxal processes equipped with bristle apparatuses (b) [= oval
coxal intermediate process with bristles sensu Mrsic (1992)]. Posterior coxal processes (p) [= syncoxal
processes sensu Mrsic (1992)] well-developed, broad at the base, “C’-curved in posterior view, apically
divided into two parts, very wide in lateral view. The most posterior parts of the anterior gonopods are
a pair of pseudoflagella (f) covered by bristles over almost their whole length, with tassels at the top
(see Discussion). These pseudoflagella starting from the well-developed, rounded lobes (1). The only
unpaired part of the anterior gonopods, apart from the sternum, is the rounded syncoxal vesicle (v) [= a
small intermediate (medial) bridge sensu Mrsic (1992)] situated between the coxal processes (deflated
on the SEM photos). There are no traces of telopodites.

Posterior gonopods (Figs 2C-D, 9A). Very reduced in both size and structure. Consist of coxal vesicles
fused in the form of a medial process (m) apically divided. Taterally with telopodal remnants (t). Tateral
sternal processes (sp) high and subtriangular.

Descriptions of vulvae

Vulvae (Fig. 3). Somewhat elongated from lateral view. Mesal (mv) and lateral (lv) valves of bursae
(bu) distally with long setae. Several setae are present on the lateral side of the lateral valvae. Fateral
valvae posteriorly with tooth-like subtriangular structures (ts), which only partly cover the mesal valvae,
so the bursa apertures can be seen from posterior view. Operculum (op) with long setae.

Distribution

Croatia: known only from several caves on Mt. Biokovo (Fig. 11, red triangles).

Habitat

Biokoviella mauriesi is limited to deep and cold caves on Mt. Biokovo with temperatures ranging from
0 to 7°C. The entrances of these caves are situated on an elevated plateau stretching from above 1300 m
a.s.l. to the highest parts of Biokovo.

Biokoviella mosorensis Antic & Drazina sp. nov.
um:lsid:zoobank.org:act:6270477C-2572-4DBC-80A3-937677DE8949

Figs 4-8, 9B, 10B

Diagnosis

The new species can be clearly separated from B. mauriesi on the basis of the smaller body size, the
presence of anterior gonopods that are much more gracile and differences in several other details of their
structure. Fateral valve of vulva with shield-like structures posteriorly.

Etymology

The new species is named after Mt. Mosor (Middle Dalmatia, Croatia), its terra typica.

Material studied (total: 14 $$, 17 $$, 4 juveniles)

Holotype

CROATIA: c?, Balica Spilja Cave (= Kraljeva Pec Cave), 43.573° N, 16.570° E, 404 m a.s.l., Balici,
village of Dugopolje, Mt. Mosor, Middle Dalmatia, Croatia, 7 Sep. 2015, leg. D. Antic & T. Rada
(NHMSC).

Paratypes (total: 10 10 4 juveniles)

CROATIA: 9 7 §§, 2 juveniles (IZB, NHMSC), same data as holotype; 1 S, 3 $?, 2 juveniles,

same data except 28 Apr. 2012, leg. M. Pavlek (CBSS).

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ANTIC D.Z. etal.. Taxonomic status of Biokoviellidae

Fig. 4. Biokoviella mosorensis Antic & Drazina sp. nov., S, paratype. A. Pleurotergite 15, posterior
view. B. Pleurotergite 15, prozonite and metazonite, dorsal view. C. Pleurotergite 15, metaznonite,
dorsal view. D. Pleurotergites 14 and 15, lateral view. E. Pleurotergites 14 and 15, dorsal view.

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European Journal of Taxonomy 205 : 1-23 ( 2016 )

Other material (total: 3 SS, 7 5 5)

CROATIA, Mt. Mosor: 1 §, Ledenica pod Jabukovcem Pit, Jabukovac, 23 Jun. 2011, leg. P. Bregovic
(CBSS); 1 5, Maklutaca Spilja Cave, Cevrljin Klen, Kute, 24 Jun. 2011, leg. R. Balcovic (CBSS); 1 S,
Velilca Gajna Pit, Gornje Sitno, 5 May 2012, leg. R. Cvitanic (CBSS); 2 5 §§, same data except

7 Jun. 2015, leg. B. Jalzic (CBSS).

Fig. 5. Biokoviella mosorensis Antic & Drazina sp. nov., paratype, gonopods. A. Anterior gonopods,
anterior view. B. Anterior gonopods, antero-lateral view. C. Anterior gonopods, lateral view. D. Anterior
gonopods, latero-distal view. E. Anterior gonopods, distal view. F. Anterior and posterior gonopods,
posterior view. Abbreviations: see Material and methods.

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ANTIC D.Z. et al ., Taxonomic status of Biokoviellidae

Type locality

CROATIA: Balica Spilja Cave (= Kraljeva Pec Cave), Balici, village of Dugopolje, Mt. Mosor, Middle
Dalmatia.

Fig. 6. Biokoviella mosorensis Antic & Drazina sp. nov., paratype, gonopods. A. Anterior gonopods,
posterior view. B. Anterior gonopods, postero-distal view of vesicle. C. Anterior gonopods, lateral view
of bristle apparatus. D. Anterior gonopods, latero-distal view of posterior coxal process and vesicle.
E. Anterior gonopods, lateral view of the apex of posterior coxal processes. F. Posterior gonopods,
postero-distal view. Abbreviations: see Material and methods.

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European Journal of Taxonomy 205 : 1-23 ( 2016 )

Fig. 7. Biokoviella mosorensis Antic & Drazina sp. nov., holotype, anterior gonopods. A-B. Anterior
view. C-D. Posterior view. E-F. Lateral view. Abbreviations: see Material and methods. Scale line =
0.2 mm .

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ANTIC D.Z. et al., Taxonomic status of Biokoviellidae

Description

Segments. Body with 30 segments (including telson) in adults. Juveniles with 28 and 26 segments
(including telson), respectively.

Measurements. Males 9—12 mm long, vertical diameter of the largest pleurotergite 0.9-1 mm Females
10.5-13 mm long, vertical diameter of the largest pleurotergite 1-1.05 mm. Holotype male 10.5 mm
long, vertical diameter of its largest pleurotergite 1 mm.

Coloration. Pigmentless, whitish (Fig. 10B).

Head. Setose. Tabrum with three medial labral teeth and with 4+4 labral and 1+1 supralabral setae.
Gnathochilarium normal. Ante nn ae elongated, 2.4 mm long in holotype male. Length of antennomeres
(in mm): I (0.1), II (0.27), III (0.58), IV (0.3), V (0.7), VI (0.22), VII (0.2) and VIII (0.03). Length/
breadth ratios of antennomeres I-VII: I (1), II (2.1), III (5.3), IV (2.7), V (5.4), VI (1.5) and VII (2).
Antennomeres II, IV, V, VI and VII with one, three, one, four and one macrochaetae, respectively. Blind.

Collum. Narrower than head, with six macrochaetae.

Body segments (Fig. 4). Without lateral keels, metazonites with only small lateral swellings. Macrochaetae
short, trichoid. CIX (pleurotergite 15) ~ 0.9; MIX (pleurotergite 15) ~ 1; PIX (pleurotergite 15) = 0.6;
MA (pleurotergite 15) ~ 140°.

Telson. Epiproct with a pair of spinnerets and six trichoid setae arranged in two rows (2+2 marginal and
1+1 paramedial setae). Hypoproct with two trichoid setae. Paraprocts with 3+3 marginal trichoid setae.

Walking legs. Elongated. Leg pairs 1 and 2 with tarsal combs in both sexes.

Male sexual characters. Leg pairs 3-7 somewhat larger than the rest of the legs, leg pairs 3 and 4
the most robust; without any peculiarities. Leg pairs 10 and 11 with coxal glands; leg pair 11 with
posteriorly directed coxal horn.

Anterior gonopods (Figs 5, 6A-E, 7). Anterior coxal processes (a) [= angiocoxites sensu Mrsic (1992)]
completely separated from each other, rising from the very wide sternal plate (s); first A (base) very
broad, remaining 2 A very gracile, long, thin and curved caudad, apically with two branches (al and
a2) opposite to each other. Posteriorly anterior coxal processes with bristle apparatuses (b) [= oval
coxal intermediate process with bristles sensu Mrsic (1992)]. Posterior coxal processes (p) [= syncoxal
processes sensu Mrsic (1992)] well-developed, broad at the base, “S”-curved from posterior view, with
only a small apical notch. The most posterior parts of the anterior gonopods are a pair of pseudoflagella
(f), naked over their whole length, with tassels at the top. These pseudoflagella starting from the well-
developed, rounded lobes (1). The only unpaired part of the anterior gonopods, apart from the sternum,
is the rounded syncoxal vesicle (v) [= small intermediate (medial) bridge sensu Mrsic (1992)] situated
between the coxal processes (deflated on the SEM photos). There are no traces of telopodites.

Posterior gonopods (Figs 6F, 9B). Very reduced in both size and structure. Consisting of coxal vesicles
fused in the form of a medial process (m) divided halfway. Laterally with telopodal remnants (t). Lateral
sternal processes (sp) low and rounded.

Vulvae (Fig. 8). Subquadrangular from lateral view. Mesal (mv) and lateral (lv) valvae of bursae
(bu) distally with long setae. Lateral valvae posteriorly with shield-like structures (ss) which almost
completely cover the mesal valvae, so that bursa apertures can’t be seen from posterior view. Operculum
(op) with long setae.

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European Journal of Taxonomy 205 : 1-23 ( 2016 )

Fig. 8. Biokoviella mosorensis Antic & Drazina sp. nov., paratvpe, vulvae. A. Posterior view. B. Lateral
view. C. Anterior view. D. Antero-distal view. E. Distal view. F. Postero-distal view. Abbreviations: see
Material and methods.

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ANTIC D.Z. et al.. Taxonomic status of Biokoviellidae

Distribution

Croatia, known only from several caves on Mt. Mosor (Fig. 11, blue circles).

Habitat

Balica Spilja is a cave with a huge entrance enabling light to penetrate through almost the whole cave.
Only the innermost part is in compete darkness and with various speleothems. Mean air temperature in
the cave is 6.3°C and humidity is high (99.6%). Altogether, the cave is 194 m long and 50 m deep. Balica

Fig. 9. Posterior gonopods. A. Biokoviella mauriesi Mrsic, 1992, S from the Tovricia 3 Pit. B. Biokoviella
mosorensis Antic & Drazina sp. nov., S, holotype. Abbreviations: see Material and methods. Scale line
= 0.2 mm

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European Journal of Taxonomy 205 : 1-23 ( 2016 )

Fig. 10. Habitus. A. Biokoviella mauriesi Mrsic, 1992, topotype (photo by H. Bilandzija).
B. Biokoviella mosorensis Antic & Drazina sp. nov., S and $ from the type serie (photo by D.A.).
Without scale.

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ANTIC D.Z. et al. , Taxonomic status of Biokoviellidae

V

Spilja is a very important biospeleological locality, being locus typicus for eight different taxa (Bedek
et al. 2006; Makarov et al. 2011): Folkia boudewijni Deeleman-Reinhold, 1993, Troglohyphantes
giromettai (Kulczynski, 1914) and Troglohyphantes strandi Absolon & Kratochvil, 1932 (all Araneae);
Neobisium dalmatinum Beier, 1938 (Pseudoscorpiones); Massarilatzelia dugopoljica Makarov & Rada,
2011 (Diplopoda); and Neotrechus ganglbaueri bluehweissi (Hoffmann, 1913), Haplotropidius taxi
subinflatus (Apfelbeck, 1907) and Spelaites grabowskii Apfelbeck, 1907 (all Coleoptera). Apart from
the aforementioned taxa, the cave is also inhabited by another interesting troglobiont or trogophile fauna,
including such species as: Oroniscus dalmaticus Strouhal, 1937, Alpioniscus balthasari (Frankenberger,
1937) and Armadillidium sp. (all Isopoda, Bedek et al. 2011 and CBSS); Apfelbeckia insculpta (L. Koch,
1867) and Brachydesmus subterraneus Heller, 1858 (both Diplopoda, present study); Nesticus eremita
Simon, 1879, Rhode sp., Stygopholcus absoloni (Kulczynski, 1914) and Barusia maheni (Kratochvil
& Miller, 1939) (all Aranea, CBSS); and Duvalius novaki (Muller, 1911) (Coleoptera, Pretner 1973).
Biokoviella mosorensis sp. nov. is found in the middle part of the cave, walking on rocks or large stones
on surfaces facing the dark.

With a depth of 176 m, Velilca Gajna Pit is one of the deepest pits on Mt. Mosor. Biokoviella mosorensis
sp. nov. was collected near the bottom of this pit, where measured air temperature was 7°C and humidity
90%. Other notable collected species of the pit’s fauna were: Alpioniscus sp. (Isopoda); Brachydesmus
subterraneus Heller, 1858 (Diplopoda); and Duvalius erichsonii (Schaufiiss, 1864), Haplotropidius taxi
(Muller, 1903), Laemostenus cavicola (Schaum, 1858), S. grabowskii and Speoplanes giganteus (J.
Muller, 1911) (all Coleoptera) (Jalzic et al. 1990 and CBSS).

14° E 15° E 16° E 17° E 18° E

Fig. 11. Distribution of the subfamily Biokoviellinae stat. nov.

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European Journal of Taxonomy 205 : 1-23 ( 2016 )

Maklutaca is a cave almost 150 m long and 39 deep (Jalzic etal. 2013). Similar to BalicaSpilj a, Maklutaca
has a huge entrance, with light penetrating through the first half of the cave. Measured air temperature
was 6.7°C. This cave is the type locality of Nicoletiella absoloni absoloni Willmann, 1940, a subspecies
of small troglobitic mite (Acari). The following troglobitic or troglophilic forms were also recorded
from this locality: T. giromettai and T. strandi (both Aranea); A. balthasari (Isopoda, Bedek et al. 2011);
Onychiuridae indet., Heteromurus (Verhoeffiella ) sp. and Pseudosinella sp. (all Collembola, CBSS); and
Neotrechus ganglbaueri (Padewieth, 1891) and H. taxi (Pretner 1973 and CBSS). Furthermore, another
diplopod species, Massarilatzelia dugopoljica Makarov & Rada, 2011, was collected in Maklutaca cave,
which is the second known locality for this Mt. Mosor endemic species (present study).

The Tedenica pod Jabukovcem Cave is situated in a deep dolina, and its specific position enables it
to retain snow and ice up to the summer months. For that reason air temperature is extremely low, on
average around 3.6° C. The cave has a total length of 105 m and depth of 34 m (Jalzic et al. 2013). This
speleological locality is an important locus typicus, with three described taxa: Neobisium maderi Beier,
1938 (Pseudoscorpiones), Traegardhia dalmatina gigantea Willmann, 1941 (Acari) and Speoplanes
giganteus giganteus (J. Muller, 1911) (Coleoptera). Pretner (1973) mentioned two additional species of
Coleoptera for this cave: H. taxi and S. grabowskii.

Notes on the ecology and distribution of Biokoviella

Biokoviella mosorensis sp. nov. inhabits cold caves on Mt. Mosor, those in which the air temperature
does not exceed 7° C and humidity ranges from 95 to 100%. A similar ecological preference is also
characteristic of B. mauriesi. Chordeumatidans are known to tolerate low temperature, and they can be
active in the soil and in rocky crevices below the snow cover (Mauries 1986, 1988). Thus, this group of
diplopods is well adapted for colonization of cold underground habitats.

The subterranean habitats of Mts Biokovo and Mosor have diverse and endemic troglobiotic faunas.
Turbulent geological history and the role of the Dinaric area as a refuge during glacial periods promoted
continuous colonization of suitable underground habitats. A high rate of endemism is present throughout
the entire Dinarides, and Mosor is no exception - even after more than 100 years of speleological and
biospeleological research (Girometta 1914), new species are still being discovered.

New data for some Balkan anthogonids

New data for Egonpretneria vudutschajldi Antic & Drazina, 2015

CROATIA: 2 $$, 1 $, Pecina Cakovec Cave, Javorinske Drage, Mlakva, Perusic, 2 Aug. 2015, leg.
B. Jalzic (CBSS); 2 $$, 1 §, Kalvarija Cave, Buzin, Gospic, 1 Sep. 2015, leg. H. Cvitanovic (CBSS).

New data for Haasia cornuata (Strasser, 1940)

STOVENIA: 1 1 juvenile, Veliki Mizuk Pit, Mokri Potolc, 15 Dec. 2015, leg. T. Delic & S. Polalc

(IZB).

New data for Haasia stenopodium (Strasser, 1966)

CROATIA: 1 (?, 2 $ $, Pepelarica Pit, Middle Velebit, 4 Jul. 2015, leg. G. Rnjak (CBSS); 1 S, 3 ? ?,
Meduza Pit, Crikvena, North Velebit National Park, 4 Aug. 2015, leg. B. Jalzic (CBSS).

STOVENIA: 1 Velika Ledenica u Paradani Pit, Lokve, Batuje, Nova Gorica, 6 Apr. 2014, leg. T.
Delic & S. Borko (IZB); 1 <$, same data except Jun. 2014, leg. S. Borko (IZB); 2 $$, 1 juvenile, Brezno
Treh Src, Cifre, Sneznik, 12 Sep. 2015, leg. T. Delic (IZB); 1 2 9 9, 2 juveniles, Brajnice Pit, Dobec,

Bezuljak, 14 Nov. 2015, leg. T. Delic (IZB).

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ANTIC D.Z. et al. , Taxonomic status of Biokoviellidae

New data for Macrochaetosoma bertiscea Antic & Makarov, 2015

MONTENEGRO: 2 juveniles, Trzy Kopce Pit, Prokletije Mountains, 17 Jul. 2015, leg. M. Petkovic
(IZB).

New data for Macrochaetosoma troglomontanum Absolon & Lang, 1933

CROATIA: 1 4 juveniles. Jama pod Gaznovcem Pit, Stilja, Vrgorac, 3 Aug. 2014, leg. T. Rada (IZB);

3 juveniles, Martina Jama Pit, Stilja, Vrgorac, 9 Aug. 2015, leg. T. Rada (IZB).

MONTENEGRO: 1 $, Jama u Dubokomdolu Pit, Njegusi, no dates or leg. (IZB); 2 $ §, Ericova Jama
Pit, Krivosije Mt., Crkvice, 20 Sep, 2006, leg. M. Perreau (IZB); 1 9, Opasna Jama Pit, Kucka Korita,
21 Jun. 2015, leg. M. Petkovic (IZB).

Notes on the distribution of the Macrochaetosomatinae

In a recent review of the family Anthogonidae, Antic et al. (2015) in their key to the anthogonid genera
stated that the genus Macrochaetosoma Absolon & Lang, 1933 is distributed in Croatia, Bosnia and
Herzegovina, Montenegro and Serbia (?). In writing that review, we unfortunately overlooked a paper
mentioning M. troglomontanum from caves in Albania (Stoev 2001).

Discussion

In their review of the family Anthogonidae Antic et al. (2015) recognized three subfamilies viz.,
Anthogoninae, Acherosomatinae and Macrochaetosomatinae, with 30 species. To the family
Anthogonidae, we now add one more subfamily, Biokoviellinae, with two species.

As stated earlier, Mrsic established a new monotypic family, Biokoviellidae, based on specimens from
caves on Mt. Biokovo, even though he acknowledged some similarities with the families Anthogonidae
and Chamaesomatidae. After examining the topotype male of B. mauriesi and specimens of B. mosorensis
sp. nov., we recognized some mistakes in the descriptions of Biokoviella and found enough evidence
to support inclusion of this genus in the family Anthogonidae and consequently synonymization of
the family Biokoviellidae with the family Anthogonidae. On the other hand, new observation on the
posterior gonopods show clear differences of the genus Biokoviella from all other anthogonid genera
and supports recognition of the Biokoviellinae as a fourth subfamily within the family Anthogonidae.

According to Antic et al. (2015), the main taxonomic character of the family Anthogonidae is the
presence of bristle apparatuses, a bristle-clad pseudoflagellum and several pseudoflagella on coxites
of the anterior gonopods. These characters (bristle apparatuses and bristle-clad pseudoflagellum) can
be seen in the genus Biokoviella. In his paper from 1992, Mrsic mentioned the presence of bristled
oval inner processes at the base of the posterior part of the anterior coxal processes and listed these
structures as one of the similarities with the subfamily Acherosomatinae (Haasiinae sensu Mrsic 1992).
Such structures indeed exist in both Biokoviella species and are much hairier than in Mrsic’s drawing
(see Mrsic 1992: 57, fig. 2), especially in the new species (Fig. 7). These structures are very similar
to, and definitely homologous with bristle apparatuses in the subfamily Macrochaetosomatinae, being
placed in both taxa in the base of the posterior part of the anterior coxal processes (see Antic et al. 2015:
figs 56-66, 76, 79, 84). Also, well-developed and high anterior coxal processes are present in some
representatives of the subfamily Anthogoninae and especially the subfamily Acherosomatinae.

According to Mrsic (1992), the presence of bristle-clad pseudoflagella in Biokoviella links it with
the subfamily Scutogoninae (fam. Chamaesomatidae). One of the main characters of Scutogoninae
is the presence of a single pair of flagella on the posterior side of the anterior gonopods. Such

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European Journal of Taxonomy 205 : 1-23 ( 2016 )

flagella in Scutogoninae are very different from the bristle-clad pseudoflagella in Biokoviella. It is
therefore unnecessary to comment on the possibility of including the genus Biokoviella in the family
Chamaesomatidae. Thus, Biokoviella cannot be linked with scutogonins. On the other hand, more
similar, but still different, bristle-clad pseudoflagella are present in some Anthogoninae [Escualdosoma
gourbaultae (Mauries, 1965) and Vascanthogona vicenteae Mauries & Barraqueta, 1985]. After
examining the topotype male of B. mauriesi and a male from another locality, we found some differences
compared to the drawing by Mrsic (1992), who presented these pseudoflagella as mascara brush
(see Mrsic 1992: 57, fig. 2), i.e., one rod with a bristle on the top. Our examination shows that these
pseudoflagella are indeed equipped with bristles on the top, in the form of a tassel not a brush, but also
with hairs over their entire length (Fig. 2E). On the other hand, in the new species, pseudoflagella are
equipped with an apical tassel only.

The only unpaired structure on the anterior gonopods actually is not a bridge between the posterior coxal
processes, but a rounded syncoxal vesicle, cited by Mrsic (1992) as a “small intermediate bridge or little
intermediate (medial) bridge (brace)”. This vesicle starts from the base of the anterior gonopods, between
the coxal processes. Clear evidence that this structure represents a vesicular structure is presented on
the SEM photographs, where the vesicle in question is obviously deflated (Figs 2A, C-E, 6B-C, 6E)
in contrast to anterior gonopods in situ, where the structure has globular shape (Fig. 7A-D, or see
Mrsic 1992: 57, fig. 2G). Such structures are also present in some other anthogonids and are especially
developed in both species of Egonpretneria Strasser, 1966 (see Antic et al. 2015: 156, figs 12 and 16,
sco; and Mrsic 1992: 72, figs 2 and 7B). This structure probably represents fused coxal vesicles. On the
other hand, in Macrochaetosomatinae coxal vesicles on the anterior gonopods are not fused but paired,
and they are situated on the posterior part of those gonopods (see Antic et al. 2015: 167, fig. 60, cv; 172,
figs 79 and 84, cv).

The similarity between the genus Biokoviella and Anthogonidae that Mrsic (1992) saw also applied to
the posterior gonopods. It is probable that he had in mind similarity of the medial process on the posterior
gonopods. He stated that the posterior gonopods “[...] are merely sternite remnants, with no coxal part at
all” and that “In the middle the medial sternite process is situated”. The medial process of the posterior
gonopods in the subfamilies Acherosomatinae and Macrochaetosomatinae is indeed of sternal origin
(see Antic et al. 2015). As a result of examining the posterior gonopods of both Biokoviella species
under a scanning electron microscope, we think that Mrsic was wrong when he wrote that the medial
process of the posterior gonopods is of sternal origin. To judge from the SEM and light microscope
photos (Figs 2C-D, 5F, 6F, 9), it seems that this medial process is actually of coxal rather than sternal
origin and probably represents partly fused coxal vesicles, but completely without the lateral coxal
processes seen in other anthogonids. Also, one more point about structure of the posterior gonopods
was omitted by Mrsic. These are telopodal remnants located lateral to the medial process in both species
(Fig. 9, t). Within the Anthogonidae, telopodal remnants are present in only two representatives of the
Macrochaetosomatinae, and they are situated on the very robust lateral coxal processes (see Antic et al.
2015: 172, figs 77-78, 82-83). Generally, the posterior gonopods of biokoviellines are very reduced in
size in comparison to the anterior gonopods or posterior gonopods of other anthogonids.

All the above-mentioned similarities between the genus Biokoviella and some representatives of
Anthogonidae with respect to structure of the anterior gonopods argue in favor of including it in the
indicated family, but on the other hand, structure of the posterior gonopods clearly separates the genus
from all other anthogonids. We therefore think that inclusion of the subfamily Biokoviellinae as a
member of Anthogonidae is justified.

Apart from features of the anterior and posterior gonopods, one more sexual character should be
discussed. This character relates to the structure of leg pairs 3-7 in males. Mrsic (1992: 54) stated that

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ANTIC D.Z. et al. , Taxonomic status of Biokoviellidae

these pairs of legs “[...] are equipped with torsal papillae”. He probably had in mind the tarsal papillae
present in some chordeumatidans. In examining these pairs of legs in our specimens of both B. mauriesi
and B. mosorensis sp. nov., we could not find any traces of tarsal papillae. Inasmuch as Mrsic (1992) in
the same paper dealt with the family Heterolatzeliidae, a small Dinaric group with tarsal papillae on leg
pairs 3-7 in males, it seems likely that he may have accidentally mixed up the legs from specimens of
these two groups. The absence of tarsal papillae in Biokoviella is one more reason for inclusion of this
genus in the family Anthogonidae.

The family Anthogonidae now includes four subfamilies: Anthogoninae, Acherosomatinae,
Biolcoviellinae and Macrochaetosomatinae. All of these subfamilies are characterized by the presence
of bristle apparatuses and/or bristle-clad pseudoflagella as parts of the anterior gonopods. On the other
hand, they all possess some apomorphies that separate them from each other, and they would surely be
recognized as separate families by some diplopodologist, a viewpoint to which we do not have any major
objections. But in our opinion, it is better for now to leave four subfamilies within Anthogonidae and
gradually reduce, if possible, the number of families within the extremely chaotic order Chordeumatida.

Acknowledgments

We are grateful to Roman Ozimec, Branlco Jalzic, Marko Lukic, Jana Bedelc, Alen Kirin, Petra Bregovic,
Robert Bakovic, Ratko Cvitanic, Martina Pavlek, Goran Rnjak, Slaven Nizetic and Hrvoje Cvitanovic

_ V

(all from Croatia); Teo Delic, Spela Borko & Slavko Polak (all from Slovenia); Matija Petkovic (Serbia)
and Michel Perreau (France) for collecting and making material considered in this study available
to us. The authors would like to thank Milos Bokorov (University Center for Electron Microscopy,
Department of Biology and Ecology, University of Novi Sad, Serbia) for the SEM photographs. Helena
Bilandzija (Croatia) lent us her wonderful photo of living Biokoviella mauriesi. Determinations of
the accompanying fauna from Mt. Mosor caves were generously given by specialists for individual
groups and we are sincerely grateful to them: P. Bregovic and B. Jalzic for Coleoptera, M. Lukic for
Collembola, M. Pavlek for Araneae and J. Bedelc for Isopoda. The Biokovo Nature Park supported
the collecting of diplopods and other forms of cave fauna through different projects. The team leader
for most biospeleological research on Mt. Biokovo was R. Ozimec, and T.D. is extremely grateful
to him for his enthusiasm in biospeleological research and for his support in the work on diplopod
taxonomy. Dr. Henrik Enghoff and one anonymous referee provided helpful co mm ents and suggestions
on the manuscript. This study was partly supported by the Serbian Ministry of Education, Science and
Technology (Grant No. 173038).

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Manuscript received: 22 December 2015
Manuscript accepted: 24 February 2016
Published on: 17 June 2016
Topic editor: Rudy Jocque
Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the
EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;
Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, London, United
Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of
Denmark, Copenhagen, Denmark.

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