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Endogean and cavernicolous Coleoptera of the Balkans. XVIII. Strong radiation in caves of the Central Dinarides: seven new species of Thaumastocephalus Poggi et al., 2001 (Staphylinidae: Pselaphinae)

Authors:
  • Natural History Museum, Prague
  • Croatian Biospeleological Society

Abstract and Figures

Seven new species of the cavernicolous and anophthalmous genus Thaumastocephalus Poggi, Nonveiller, Colla, Pavićević & T. Rađa, 2001 are described: T. bilandzijae sp. n., T. kirini sp. n., T. marsici sp. n., T. rujnicensis sp. n., T. slavkoi sp. n. and T. troglavi sp. n. from Croatia and T. dahnae sp. n. from Bosnia and Herzegovina. Aedeagi of all species are illustrated. A key to all species is provided. The records of all specimens of the genus treated here are given, and their distributions are discussed and shown on maps. The distribution of all genera of cavernicolous Pselaphinae in the Dinarides is discussed.
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Accepted by Z.-W. Yin: 19 Nov. 2018; published: 19 Feb. 2019
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
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Copyright © 2019 Magnolia Press
Zootaxa 4559 (1): 090
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Article
https://doi.org/10.11646/zootaxa.4559.1.3
http://zoobank.org/urn:lsid:zoobank.org:pub:F051C9DE-CCB7-454E-8184-B6240049F004
Endogean and cavernicolous Coleoptera of the Balkans. XVIII.
Strong radiation in caves of the Central Dinarides: seven new species of
Thaumastocephalus Poggi et al., 2001 (Staphylinidae: Pselaphinae)
PETER HLAVÁČ
1
, PETRA BREGOVIĆ
2,3
& BRANKO JALŽ
4
1
Department of Entomology, National Museum, Natural History Museum, Cirkusová 1740, CZ-193 00 Praha 9—Horní Počernice,
Czech Republic. E-mail. peterclaviger@gmail.com
2
Croatian Biospeleological Society, Demetrova 1, 10 000 Zagreb, Croatia. E-mail: bregovicpetra@gmail.com
3
SubBioLab, Department of Biology, Biotechnical Faculty, University of Ljubljana, Jamnikarjeva 101, 1000 Ljubljana, Slovenia
4
Croatian Biospeleological Society, Demetrova 1, 10 000 Zagreb, Croatia. E-mail: jalzicbranko@gmail.com
Abstract
Seven new species of the cavernicolous and anophthalmous genus Thaumastocephalus Poggi, Nonveiller, Colla, Pa-
vićević & T. Rađa, 2001 are described: T. bilandzijae sp. n., T. kirin i sp. n., T. marsici sp. n., T. rujnicensis sp. n., T. slavkoi
sp. n. and T. troglavi sp. n. from Croatia and T. dahnae sp. n. from Bosnia and Herzegovina. Aedeagi of all species are
illustrated. A key to all species is provided. The records of all specimens of the genus treated here are given, and their
distributions are discussed and shown on maps. The distribution of all genera of cavernicolous Pselaphinae in the Di-
narides is discussed.
Key words: Thaumastocephalini, taxonomy, biospeleology, Croatia, Bosnia and Herzegovina, zoogeography
Introduction
Thaumastocephalus Poggi, Nonveiller, Colla, Pavićević & T. Rađa, 2001 is an extraordinary cavernicolous genus
of the monogeneric tribe Thaumastocephalini, that was recently described from a series of specimens collected in
caves of the Kozjak and Mosor Mountains in the Croatian Dinarides. The cave Mala Birnjača, which is situated on
the slopes of Kozjak Mountain (Jalžić et al. 2010), is the type locality of T. folliculipalpus Poggi, Nonveiller, Colla,
Pavićević & T. Rađa, the only described species of the genus. However, the paratype female specimens from
Vranjača cave, Mosor Mountain, belong to a different, still undescribed species (Besuchet, pers. comm.).
Subsequent research by members of the Croatian Biospeleological Society (CBSS) revealed that
Thaumastocephalus is a common genus that is widely distributed in the caves of the Central Dinarides. The aim of
this paper is to describe seven new species, as well as to provide the first discussion about the biogeography of the
genus Thaumastocephalus.
Material and methods
Specimens prepared for the morphological study were examined using a Leica S8APO stereoscopic microscope
with diffuse lighting at magnifications of up to 128×. Habitus images were taken in situ with a Canon EOS 80D in
the combination with a Canon MP-E65 1-5x macro lens.
The aedeagus was studied using a Zeiss transmitted-light microscope at magnifications of up to 500X. The
aedeagus was dissected and preserved in Euparal on a plastic card pinned together with the specimen. All drawings
were made using a drawing tube and digitized using Corel DRAW X7.
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The head capsule length was measured from the occipital constriction to the anterior margin of the frontal
rostrum; which is an anterior part of the head extending beyond the antennal insertions and raised above the plane
of the clypeus and labrum; elytral length was measured along the suture; width always refers to the maximum
width of a given structure, e.g. head, pronotum, elytra. The body length is the combined length of the head,
pronotum, elytra, and abdomen. All measurements and ratios given in descriptions refer to the holotype.
Label data is cited verbatim. All labels of the studied material are printed. All type specimens were provided
with the following red printed label: HOLOTYPE or PARATYPE, generic and specific name of the taxon, P.
Hlaváč det., 2018.
The material is deposited in the following collections: Croatian Natural History Museum, Zagreb (CNHM), the
private collection of Peter Hlaváč (CPH, Prague, The Czech Republic), and the private collection of Jan Lakota
(CJL, Ružomberok, Slovakia).
Taxonomy
Thaumastocephalus Poggi, Nonveiller, Colla, Pavićević & T. Rađa
Thaumastocephalus Poggi, Nonveiller, Colla, Pavićević & T. Rađa, 2001: 3. Type species: Thaumastocephalus folliculipalpus
Poggi, Nonveiller, Colla, Pavićević & T. Rađa, 2001
Diagnosis. Large, body length 1.8–2.2 mm, cavernicolous, anophthalmous and depigmented pselaphines placed in
the supertribe Batrisitae. Thaumastocephalus can be readily separated from all other genera of Batrisitae by having
a pedunculate gular process, and by the 2
nd
and 3
rd
palpomeres (Fig. 9) bearing on their external sides a thin
filament ending in a spherical appendix (Poggi et al. 2001). The placement of Thaumastocephalus within the
Batrisini is still a subject of debate but according to the extensive study of DNA of a large number of Pselaphinae it
seems to be well seated in Batrisitae (Parker pers. comm.).
Thaumastocephalus bilandzijae Hlaváč, Bregović & Jalžić, sp. n.
(Figs 1, 11)
Etymology. The species is named after biospeleologist Helena Bilandžija, a member of the Croatian
Biospeleological Society (CBSS) and collector of the holotype specimen.
Material studied. HOLOTYPE: ♂: CROATIA: Dugopolje, Kolić, Peć u Čulinovim raljevinama (cave),
6.I.2009, H. Bilandžija lgt. (CNHM). PARATYPES (2♀): same locality as the holotype but collected on 5.XI.2016
by A. Kirin (CNHM, CPH).
Description. Body shiny (Fig. 11), reddish-brown, covered with very sparse, uneven, golden setae; legs,
antennae and maxillary palpi of same colour, length 1.82 mm, maximum width of elytra 0.63 mm. Head 1.22 times
as long as wide, 0.84 times narrower than pronotum, rostrum 2.17 times as wide as long. Antennae about 0.68 mm
long, scape twice as long as pedicel, antennomeres III–VII of same length, VIII slightly shorter than VII and as
long as IX, antennomere X about as long as VII, terminal antennomere four times as long as X, relative length of
antennomeres: 1.00 / 0.50 / 0.33 / 0.33 / 0.33 / 0.33 / 0.25 / 0.25 / 0.30 / 1.17. Pronotum 1.16 times as long as wide.
Elytra 1.32 times as wide as long and 1.28 times as long as pronotum. Abdomen 1.31 times as long as elytra. Legs
long, protibiae modified, depressed in apical half, with median, well-defined sharp spine, mesotibiae narrowed and
slightly bent at apex, metatibiae simple. Aedeagus (Fig. 1) 0.33 mm long, with basal bulb about 1.54 times as long
as apical part, right paramere more robust than left, both with three preapical setae, internal sac with two
asymmetric structures, dorsal diaphragm present.
Sexual dimorphism. Females lacking modification of pro- and mesotibiae.
Differential diagnosis. T. biland z i j ae is readily separated from all other known species of the genus by: 1)
small size, length under 1.85 mm, 2) antennae short, only 0.68 mm long, and 3) scape which is only twice as long
as pedicel.
Distribution and habitat. T. bilandzijae is known only from the cave Peć u Čulinovim raljevinama (Figs 10,
12), situated at 425 m a.s.l., near the village Kolić in the municipality Dugopolje (Fig. 30, red square). The cave has
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two separate chambers, and is about 25 m long and 15 m deep (Jalžić et al. 2010). The air temperature measured in
the cave on 25th June 2011 was 13 °C. The cave is the type locality for Coleoptera—Duvalius novaki novaki
(Müller, 1911). The following fauna is also recorded from the cave: Araneae—Barusia maheni (Kratochvíl &
Miller, 1939) and Troglohyphantes strandi Absolon & Kratochvíl, 1932 as noted by Deeleman-Reinhold (1978);
Isopoda—Alpioniscus sp. (det. J. Bedek); Coleoptera—Laemostenus cavicola modestus (Schaufuss, 1862) as noted
by Pretner (1973).
Thaumastocephalus kirini Hlaváč, Bregović & Jalžić, sp. n.
(Figs 2, 14)
Etymology. This species is named after speleologist Alen Kirin, a member of the Croatian Biospeleological
Society (CBSS) and collector of the first specimen from the type locality.
Material studied. HOLOTYPE: ♂: CROATIA: Metković, Desne, Rujnica, Varda. Jamica u docima (cave),
17.VI.2015, B. Jalžić lgt. (CNHM). PARATYPES (7♀): 2♀: same data as holotype; 4♀: same data as holotype but
collected on 30.IV.2010 by A. Kirin (CNHM, CPH); 1♀: same data as holotype but collected on 16.IX.2016 by B.
Jalžić (CNHM).
Description. Body shiny (Fig. 14), reddish-brown, covered with dense, uneven setae, with long setae on
posterior part of elytra, setae lacking on disc of elytra; legs, antennae and maxillary palpi slightly lighter, length
1.96 mm, maximum width of elytra 0.68 mm. Head 1.32 times as long as wide, 0.79 times narrower than pronotum,
rostrum 3.00 times as wide as long. Antennae about 0.82 mm long, scape 2.57 times as long as pedicel,
antennomere III slightly longer than IV, antennomeres IV–VII of same length, VIII slightly shorter than VII and as
long as IX, antennomere X about as long as VII, terminal antennomere 3.10 times as long as X, relative length of
antennomeres: 1.00 / 0.39 / 0.28 / 0.25 / 0.25 / 0.25 / 0.25 / 0.22 / 0.22 / 0.25 / 0.78. Pronotum 1.10 times as long as
wide. Elytra 1.22 times as wide as long and 1.40 times as long as pronotum. Abdomen 1.14 times as long as elytra.
Legs long, protibiae slightly modified, slightly depressed in apical half, median spine minuscule, mesotibiae
simple, metatibiae simple. Aedeagus (Fig. 2) 0.49 mm long, with basal bulb about 1.15 times as long as apical part,
right paramere more robust than left, both with three preapical setae, internal sac with one spiral structure, dorsal
diaphragm present.
Sexual dimorphism. Females lacking modification of protibiae.
Differential diagnosis. T. kir i ni is readily separated from all other known species of the genus by: 1) frontal
tibiae at most with minuscule tooth, 2) body length under 2 mm, 3) terminal antennomere short, less than 4 times as
long as antenommere X, 4) elytra long, more than 1.3 times as long as pronotum, and 5) short rostrum which is
about 3 times as wide as long.
Distribution and habitat. T. k irini is known only from the cave Jamica u docima (Figs 13, 15), situated at 345
m a.s.l. near peak Varda (517 m) and village Desne (Fig. 30, violet star). The cave is a vertical shaft, 10.2 m deep
and 5 m long. The air temperature measured in the cave on 30th April 2010 was 10.1 °C and the relative humidity
was 98 %. The following fauna is also recorded from the cave: Isopoda—Alpioniscus verheoffi (Strouhal, 1938) as
noted by Bedek et al. 2011; Coleoptera—Neotrechus sp., Speonesiotes sp. (det. B. Jalžić), and Bryaxis scapularis
(Reitter, 1881) as noted by Bekchiev and Hlaváč (2016).
Thaumastocephalus marsici Hlaváč, Bregović & Jalžić, sp. n.
(Figs 3, 17)
Etymology. This species is named after the local guide to the cave Boro Maršić and village Maršići located near
the type locality.
Material studied. HOLOTYPE: ♂: CROATIA: Vrgorac, Maršići, Stilja, Jama pod Gažnovcem (cave),
18.IX.2016, A. Kirin & P. Bregović lgt. (CNHM). PARATYPES (1♂, 3♀): same data as holotype (CNHM, CPH).
Description. Body shiny (Fig. 17), reddish-brown, covered with dense, uneven setae, with long setae on
posterior part of elytra, setae lacking on disc of elytra and on disc of first ventrite; legs, antennae and maxillary
palpi slightly lighter, length 1.92 mm, maximum width of elytra 0.68 mm. Head 1.28 times as long as wide, 0.75
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times narrower than pronotum, rostrum 2.33 times as wide as long. Antennae about 0.84 mm long, scape 2.64 times
as long as pedicel, antennomere III slightly longer than IV, antennomeres IV–VII of same length, VIII slightly
shorter than VII and as long as IX, antennomere X about as long as VII, terminal antennomere 3.33 times as long as
X, relative length of antennomeres: 1.00 / 0.38 / 0.30 / 0.24 / 0.24 / 0.24 / 0.24 / 0.22 / 0.22 / 0.24 / 0.81. Pronotum
1.13 times as long as wide. Elytra 1.25 times as wide as long and 1.33 times as long as pronotum. Abdomen 1.17
times as long as elytra. Legs long, protibiae slightly modified, slightly depressed in apical half, median spine
minuscule, mesotibiae simple, metatibiae simple. Aedeagus (Fig. 3) 0.58 mm long, with basal bulb as long as
apical part, right paramere more robust than left, left paramere with four, right with three preapical setae, internal
sac with large asymmetric, bifurcate structure, dorsal diaphragm present.
Sexual dimorphism. Females lacking modification on protibiae.
Differential diagnosis. T. marsici is readily separated from all other known species of the genus by: 1)
protibiae at most with minuscule tooth, 2) body length under 2 mm, 3) terminal antennomere short, less than 4
times as long as antennomere X, 4) elytra long, more than 1.3 times as long as pronotum, and 5) long rostrum that
is about 2.3 times as wide as long.
Distribution and habitat. T. marsici is known only from the cave Jama pod Gažnovcem (synonym Špilja pod
Gažnovcem) (Figs 16, 18), situated at 510 m a.s.l. near village Maršići (Fig. 30, green circle). The cave is small, it
consists of one chamber that is 45 m long and 3.5 m deep. The air temperature measured in the cave on 18th
September 2016 was 11.6 °C and the relative humidity was 99.2 %. The cave is the type locality for Diplopoda—
Typhloiulus opisthonodus Antić, 2018 (see Antić et al. 2018) and Pseudoscorpiones—Neobisium curcici
Dimitrijević, 2016 (Dimitrijević & Rađa 2016). The study of Antić et al. (2018) reports an especially rich
Diplopoda fauna fot this cave—Typhloglomeris sp., Apfelbeckia insculpta (L. Koch, 1867), Dyocerasoma
biokovense Mršić, 1986, Macrochaetosoma troglomontanum Absolon & Lang, 1933 (Antić et al. 2016),
Brachydesmus lapadensis Verhoeff, 1897, B. lobifer Verhoeff, 1897, B. subterraneus Heller, 1858, Metonomastus
albus (Verhoeff, 1901), and Stosatea iadrense (Pregl, 1883), as well as the following fauna: Coleoptera—Bryaxis
scapularis (Reitter, 1881), Phaneropella lesinae (Reitter, 1881), Speonesiotes sp. and Neotrechus sp.; Hemiptera—
Trirhacus cf. biokovensis Dlabola, 1971; and Opiliones—Cyphopthalmus sp. We have also found Tychobythinus
sp. (det. P. Hlaváč) in the cave.
Thaumastocephalus rujnicensis Hlaváč, Bregović & Jalžić, sp. n.
(Fig. 4)
Etymology. This species is named after Rujnica where the cave Bežurova jama is situated.
Material studied. HOLOTYPE: ♂: CROATIA: Metković, Desne, Rujnica, Varda, Bežurova jama (cave),
9.II.2015, B. Jalžić lgt. (CNHM). PARATYPE (1♂): same data as holotype (CPH).
Description. Body shiny, reddish-brown, covered with dense, uneven setae, with long setae on posterior part
of elytra; legs, antennae and maxillary palpi of the same colour, length 1.97 mm, maximum width of elytra 0.66
mm. Head 1.25 times as long as wide, 0.74 times narrower than pronotum, rostrum 2.50 times as wide as long.
Antennae about 0.84 mm long, scape 2.43 times as long as pedicel, antennomere III as long as IV, antennomere IV
1.2 times as long as V, V–VII of same length, VIII slightly shorter than VII and slightly shorter than IX,
antennomere X about as long as VII, terminal antennomere 3.56 times as long as X, relative length of
antennomeres: 1.00 / 0.41 / 0.29 / 0.29 / 0.26 / 0.26 / 0.26 / 0.24 / 0.26 / 0.26 / 0.94. Pronotum 1.07 times as long as
wide. Elytra 1.30 times as wide as long and 1.17 times as long as pronotum. Abdomen 1.26 times as long as elytra.
Legs long, protibiae slightly modified, slightly depressed in apical half, median spine minuscule, mesotibiae
simple, metatibiae simple. Aedeagus (Fig. 4) 0.38 mm long, with basal bulb about 1.45 times as long as apical part,
right paramere more robust than left, both with three preapical setae, internal sac with two asymmetric structures
from which one is spiral, dorsal diaphragm present.
Sexual dimorphism. Females lacking modification of protibiae.
Differential diagnosis. T. rujnicensis is readily separated from all other known species of the genus by: 1)
protibiae at most with minuscule tooth, 2) body length under 2 mm, 3) terminal antennomere short, less than 4
times as long as antennomere X, 4) elytra short, less than 1.2 times as long as pronotum.
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FIGURES 1–4. 1—Thaumatocephalus bilandzijae sp. n., aedeagus, dorsal view (scale: 0.1 mm); 2—Thaumatocephalus kirini
sp. n., aedeagus, dorsal view (scale: 0.2 mm); 3—Thaumatocephalus marsici sp. n., aedeagus, dorsal view (scale: 0.2 mm);
4—Thaumatocephalus rujnicensis sp. n., aedeagus, dorsal view (scale: 0.1 mm).
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FIGURES 5–9. 5—Thaumatocephalus slavkoi sp. n., aedeagus, dorsal view (scale: 0.1 mm); 6—Thaumatocephalus troglavi
sp. n., aedeagus, dorsal view (scale: 0.2 mm); Thaumatocephalus dahnae sp. n., 7—aedeagus, dorsal view (scale: 0.1 mm); 8—
protibiae (scale: 0.1 mm); 9—maxillary palpomere (scale: 0.1 mm).
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Distribution and habitat. T. rujnicensis is known only from the cave Bežurova jama (Figs 19, 21), situated at
430 m a.s.l., about 200 m away from the cave Jamica u Docima (type locality of T. k irini), near the peak Varda (517
m, Fig. 20) and the village Desne (Fig. 30, yellow circle). The cave is 51 m deep, and the air temperature measured
in the cave on 23rd August 1997 was 10.3 °C. The following fauna is also recorded in the cave: Coleoptera—
Neotrechus sp., Speonesiotes sp. (det. B. Jalžić), and Bryaxis scapularis (Reitter, 1881) as noted by Bekchiev and
Hlaváč (2016).
FIGURES 10–12. 10—entrance to Peć u Čulinovim raljevinama (cave), type locality (photo Marko Lukić); 11—
Thaumatocephalus bilandzijae sp. n., habitus in situ (photo Alen Kirin); 12—topographical map of the type locality (the
finding place of the T. bilandzijae sp. n. is marked with red dots).
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Thaumastocephalus slavkoi Hlaváč, Bregović & Jalžić, sp. n.
(Figs 5, 23)
Etymology. This species is named after Slavko Antunović from the village Antunovići, the local guide to the cave
near the area of Kozica.
Material studied. HOLOTYPE: ♂: CROATIA: Biokovo, Kozica, Jujnovići, Jujnovića špilja (cave),
29.IV.2014, P. Bregović lgt. (CNHM). PARATYPE (1♂): the same data as the holotype but collected on 14.V.2016
by A. Kirin (CPH).
Description. Body shiny (Fig. 23), reddish-brown, covered with dense, uneven setae, with long setae on
posterior part of elytra; legs, antennae and maxillary palpi of the same colour, length 1.97 mm, maximum width of
elytra 0.72 mm. Head 1.23 times as long as wide, 0.79 times narrower than pronotum, rostrum 1.78 times as wide
as long. Antennae about 0.88 mm long, scape 2.19 times as long as pedicel, antennomere III slightly longer than IV,
antennomeres IV–VII of same length, all 1.5 times as long as VIII, antennomeres VIII–X of same length, terminal
antennomere 4.25 times as long as X, relative length of antennomeres: 1.00 / 0.46 / 0.31 / 0.29 / 0.29 / 0.29 / 0.29 /
0.23 / 0.23 / 0.23 / 0.97. Pronotum as long as wide. Elytra 1.37 times as wide as long and 1.30 times as long as
pronotum. Abdomen 1.20 times as long as elytra. Legs long, protibiae slightly modified, slightly depressed in
apical half, median spine minuscule, mesotibiae simple, metatibiae simple. Aedeagus (Fig. 5) 0.35 mm long, with
basal bulb about 2.8 times as long as apical part, right paramere more robust than left, both with three preapical
setae, internal sac with one asymmetric structure, dorsal diaphragm present.
Sexual dimorphism. Female unknown.
Differential diagnosis. T. slavkoi is readily separated from all other known species of the genus by: 1) frontal
tibiae at most with minuscule tooth, 2) body length under 2 mm, 3) terminal antennomere long, 4.25 times as long
as antennomere X, and 4) long rostrum that is only 1.78 times as wide as long.
Distribution and habitat. T. slavkoi is known only from the cave Jujnovića špilja (Figs 22, 24), situated at 410
m a.s.l., near the village Jujnovići, on the slopes of Biokovo Mountain (Fig. 30, red circle). The cave consists of
two chambers, and is 108 m long and 16 m deep. The air temperature measured in the cave on 14th May 2016 was
8.9 °C and on 15th October 2014 was 13.8 °C; the relative humidity was 96.8 %. The following fauna is also
reported from the cave: Isopoda—Alpioniscus sp., Alpioniscus verheoffi (Strouhal, 1938) and Oroniscus
dalmaticus Strouhal, 1937 (det. J. Bedek); Diplopoda—Macrochaetosoma troglomontanum Absolon & Lang, 1933
as noted by Antić et al. 2015; Collembola—Coecobrya cf. tenebricosa (Folsom, 1902) and Heteromurus nitidus
(Templeton, 1835) as noted by Lukić and Deharveng (2008); Coleoptera—Laneyriella staudacheri (Müller, 1934),
Speonesiotes sp., Neotrechus dalmatinus (L. Miller, 1861), Laemostenus cavicola (Schaum, 1858) (see Bregović et
al. 2015) and Speluncarius biokovensis Hlaváč, Lakota & Čeplík, 2016.
Thaumastocephalus troglavi Hlaváč, Bregović & Jalžić, sp. n.
(Figs 6, 26)
Etymology. The species is named after the mountain peak Troglav near the cave Sistem Velika-Velika.
Material studied. HOLOTYPE: ♂: CROATIA: Biokovo, M. Troglav, Sistem Velika-Velika (cave),
22.VI.2017, A. Kirin lgt. (CNHM). PARATYPES 1♂, 2♀: same data as holotype (CNHM, CPH). Other material:
1♂: CROATIA: Biokovo, Župa, Gradska spila (cave), 15.V.2016, V. Sudar lgt (CPH). 1♀: same data as previous
paratype but collected on 23.I.2016 by M. Lukić (CNHM).
Remarks. The specimens from Gradska spila are tentatively attributed to this species as the male has
practically identical aedeagus, but due to the following reasons we did not include them in the type material od T.
troglavi: 1) ecological differences of habitat between Sistem Velika-Velika and Gradska spila (see details further in
text), and 2) different fauna composition in the highest part of Biokovo Mountain (see Discussion). More male
specimens from both caves will be needed to check the identity of specimens from Gradska spila with T. troglavi.
Description. Body shiny (Fig. 26), reddish-brown, covered with dense, uneven setation, with long setae on
posterior part of elytra; legs, antennae and maxillary palpi of the same colour, length 2.07 mm, maximum width of
elytra 0.69 mm. Head 1.25 times as long as wide, 0.71 times narrower than pronotum, rostrum 2.14 times as wide
as long. Antennae about 0.87 mm long, scape 2.38 times as long as pedicel, antennomere III slightly longer than IV,
IV–VII of same length, VIII slightly shorter than VII and as long as IX, X 1.25 times as long as IX, terminal
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antennomere 3.78 times as long as X, relative length of antennomeres: 1.00 / 0.42 / 0.26 / 0.24 / 0.24 / 0.24/ 0.24 /
0.21 / 0.21 / 0.24 / 0.89. Pronotum 1.07 as long as wide. Elytra 1.31 times as wide as long and 1.17 times as long as
pronotum. Abdomen 1.37 times as long as elytra. Legs long, protibiae slightly modified, slightly depressed in
apical half, median spine absent, mesotibiae simple, metatibiae simple. Aedeagus (Fig. 6) 0.59 mm long, with basal
bulb about 1.5 times as long as apical part, right paramere smaller than left, both with three preapical setae, internal
sac with two asymmetric structures, dorsal diaphragm present.
Sexual dimorphism. Females lacking modification of protibiae.
Differential diagnosis. T. troglavi is readily separated from all other known species of the genus by: 1)
protibiae at most with minuscule tooth and 2) body length over 2.5 mm.
Distribution and habitat. T. tro glavi is known from the cave system Velika-Velika (Figs 25, 27), situated at
1375 m a.s.l. near the peak M. Troglav on Biokovo Mountain (Fig. 30, dark blue circle). The cave is 135 m long
and 48 m deep, and it was discovered during 1
st
Biospeleological Expedition—Biokovo 2017 (Sudar et al. 2017).
The air temperature measured in the cave on 1st June 2017 was 2.2 °C and the relative humidity was 99 %. The
following fauna is also recorded in the cave: Diplopoda—Biokoviella mauriesi Mršić, 1992 (det. T. Dražina);
Coleoptera—Pygoxyon sp. (det. P. Hlaváč) and Speoplanes giganteus biocovensis Müller, 1934 (det. P. Bregović).
The second possible location of T. troglavi is the cave Gradska spila (see above Remarks) situated at 750 m a.s.l.,
near the village Župa, on the slopes of Biokovo Mountain (Fig. 30, light blue circle). The cave is small, 26.5 m long
and 1.8 m deep. The air temperature measured in the cave on 15th May 2016 was 10.1 °C and the relative humidity
was 97.9 %. The following fauna is also recorded from the cave: Isopoda—Alpioniscus sp., Alpioniscus verheoffi
(Strouhal, 1938) (det. J. Bedek) and Oroniscus dalmaticus Strouhal, 1937 as noted by Bedek et al. 2011;
Collembola—Heteromurus nitidus Templeton, 1835, Orchesella sp., Verhoeffiella sp. and Tomocerus terrestralis
Stach, 1922 (Lukić & Deharveng 2008); Coleoptera—Neotrechus dalmatinus (L. Miller, 1861), Speonesiotes sp.
and Speluncarius cf. biokovensis Hlaváč, Lakota & Čeplík, 2016 (det. P. Bregović).
Thaumastocephalus dahnae Hlaváč, Bregović & Jalžić, sp. n.
(Figs 7–9, 29)
Etymology. This species is named after the type locality, the cave Dahna, where the holotype specimen was
collected.
Material studied. HOLOTYPE: ♂: BOSNIA AND HERZEGOVINA: Tomislavgrad, Omerovići, Dahna
(cave), 29.III.2012, M. Vasić lgt. (CNHM).
Description. Body shiny (Fig. 29), yellowish-brown, abdomen darker, covered with dense, uneven setae, with
long setae on posterior part of elytra; legs, antennae and maxillary palpi of same colour, length 1.95 mm, maximum
width of elytra 0.66 mm. Head 1.30 times as long as wide, 0.83 times narrower than pronotum, rostrum twice as
wide as long. Antennae about 0.79 mm long, scape twice as long as pedicel, antennomere III 1.5 times as long as
IV, V slightly longer than IV, V–VII of same length, VIII slightly shorter than VII, VIII–X of same length, terminal
antennomere 3.75 times as long as X, relative length of antennomeres: 1.00 / 0.50 / 0.33 / 0.27 / 0.30 / 0.30 / 0.30 /
0.27 / 0.27 / 0.27 / 1.00. Pronotum 1.08 as long as wide. Elytra 1.16 times as wide as long and 1.46 times as long as
pronotum. Abdomen 1.05 times as long as elytra. Legs long, protibiae slightly modified (Fig. 8), slightly depressed
in apical half, median spine minuscule, mesotibiae simple, metatibiae simple. Aedeagus (Fig. 7) 0.35 mm long,
with basal bulb about 1.35 times as long as apical part, right paramere more robust than left, both with three
preapical setae, internal sac with one large asymmetric structure, dorsal diaphragm present.
Sexual dimorphism. Female unknown.
Differential diagnosis. T. dahnae and T. folliculipalpis are readily separated from all other known species of
the genus by the modified protibiae bearing a well-defined median tooth (Fig. 8); from the latter species T. dahnae
can be easily distinguished by its smaller size, 1.95 mm versus 2.2 mm, and by the head that is 1.3 times as long as
wide versus more than 1.4 times as long as wide in the latter species.
Distribution and habitat. T. dahnae is known only from type locality, the cave Dahna (Fig. 28), situated at
935 m a.s.l., near the village Omerovići in the western part of Duvanjsko polje (Fig. 30, red triangle). The cave is
about 1100 m long. The air temperature measured on 29
th
March 2012 was 9.1 °C; and the relative humidity was
92.2 % at the entrance chamber of the cave, and 6.4 °C in the deeper parts of the cave. The cave is the type locality
for Coleoptera—Leptomeson dombrowskii pubipenne (Müller, 1941), and it is also known in the literature as the
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cave Dana pećina or Bezdana pećina (Lukić Bilela & Ozimec 2013, Pretner 2011). The following fauna is also
recorded from the cave: Isopoda—Alpioniscus heroldi (Verhoeff, 1931) (det J. Bedek); Coleoptera—
Haplotropidius pubescens as noted by Pretner (2011).
Key to males of Thaumastocephalus
1 Small species, body length under 1.85 mm, aedeagus (Fig. 1). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. bilandzijae Poggi et al.
- Larger species, body length more 1.90 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Protibiae in males modified, with well-defined median tooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
- Protibiae in males slightly modified, median tooth minuscule or absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
3 Large species, body length 2.2 mm, head much longer than wide, ratio: 1.43–1.47, aedeagus (see Poggi et al. 2001: Fig. 10). .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. folliculipalpis Poggi et al.
- Small species, body length about 1.95 mm, head shorter than wide, ratio: 1.30, aedeagus (Fig. 7) . . . . . . . . . . T. dahnae sp. n.
4 Larger species, body length over 2.5 mm, aedeagus (Fig. 6) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .T. troglavi sp. n.
- Smalle species, body length under 2.0 mm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
5 Antennae short, less than 0.85 mm long, scape more than 2.3 times as long as pedicel, terminal antennomere short, less than
4.00 times as long as antennomere X . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
- Antennae long, about 0.88 mm long, scape less than 2.2 times as long as pedicel, terminal antennomere long, 4.25 times as
long as antennomere X, rostrum long, only 1.78 times as wide as long, aedeagus (Fig. 5) . . . . . . . . . . . . . . . . . T. slavkoi sp. n.
6 Elytra short, less than 1.2 times as long as pronotum, aedeagus (Fig. 4) . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. rujnicensis sp. n.
- Elytra long, more than 1.3 times as long as pronotum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Rostrum long, about 2.3 times as wide as long, aedeagus (Fig. 3). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. marsici sp. n.
- Rostrum short and wide, about 3.0 times as wide as long, aedeagus (Fig. 2). . . . . . . . . . . . . . . . . . . . . . . . . . . . . .T. kirin i sp. n.
Further material of Thaumastocephalus available for this study
Remarks. Six additional and potentially new species from six different caves have been discovered but are not
decribed here, because only females were available. Another six species exist in the Museum of Geneva. We have
three of them and they are listed in this work (see below). There is an agreement that these six species will be
described in the future (Cuccodoro pers. comm.).
CROATIA: Biokovo: 1♀: Dedići, Rastovac, Špilja u Radinovcima (cave), 1.V.2014, B. Jalžić & P. Bregović
lgt. (CNHM). 1♀: the same locality but collected on 16.X.2014 by J. Lakota (CJL). 2♀: Kozica, Jujnovići, Spila 2
(cave), 14.V.2016, B. Jalžić lgt. (CNHM). 2♀, 2♂: Župa, Matijaševa špilja (cave), 15.V.2004, B. Jalžić lgt
(CNHM, CPH). 1 ex: the same locality but collected on 15.V.2016 by V. Sudar (deposited in 95.5 % ethanol, CPH).
2♂: Zagvozd, Brnasi, Špilja u Gaju (cave), 16.V.2016, A. Ćukušić lgt. (CNHM, CPH). 1♂: Gornje Igrane, Sošići,
Saranač, Jama iznad Saranača (cave), 17.V.2016, T. Rožman lgt. (CNHM). 1♀: the same locality but collected on
17.V.2016 by A. Čukušić (CNHM). Mosor: 1♀: Balići, Dugopolje, Balićeva špilja (cave), 28.IV.2012, T. Čuković
lgt. (CNHM). 2♀: Omiš, Podgrade, Šćadin, Muzejska jama (cave), 2.VII.2014, R. Cvitanić lgt. (CNHM). 1♀:
Split, Dugopolje, Žmirino gumno, Žmirina jama (cave), 29.IV.2012, A. Čukušić lgt. (CNHM). 1♂: Dugopolje,
Kotlenice, Vranjača špilja (cave), 26.X.2007, B. Jalžić lgt. (CNHM). Other: 1♀: Sinj, Neorić, Sutina, Bazine,
Mala špilja (cave): 14.VI.2011, H. Bilandžija & B. Jalžić lgt. (CNHM). 1♀: the same locality but collected on
21.V.2016 by N. Kuharić (CNHM).
Discussion
The Balkan Peninsula, together with the Pyrenees and the Caucasus, is the most important hotspot of biodiversity
(Kryštufek & Reed 2004) in the western Palaearctic region. This is particularly so for the subterranean fauna,
where the Balkan Peninsula has practically no competition. There is no other place on our planet with such a great
concentration of terrestrial subterranean beetles (Hlaváč et al. 2017). In total, 1427 species and subspecies
attributed into 219 genera and 714 subgenera have been described (Hlaváč et al. 2017 and subsequent descriptions,
Hlaváč pers. database). The Dinarides in the western Balkan Peninsula is the most species-rich region for both
aquatic and terrestrial obligate hypogean taxa (Sket et al. 2004). To date, 791 species and subspecies (Hlaváč pers.
database) have been described from this area, which is 55.5% of all subterranean Coleoptera known from the
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Balkan Peninsula. The Dinarides can be biogeographically divided into three bio-geographical subregions:
Northern, Central, and Southern Dinarides (Hlaváč et al. 2008, 2017). The Southern Dinarides is the most species-
rich with 429 taxa recorded, followed by the Northern Dinarides with 200 taxa, while the Central Dinarides hosts
162 taxa. The subregion of the Central Dinarides extends from the Una River to the Neretva River, an area of about
230 km in length and up to 140 km in width. Due to the low number of Pselaphinae (only twelve species of
subterranean Pselaphinae are known from the Central Dinarides, see Hlaváč et al. 2017), but a high number of
other subterranean beetles are recorded from this area, the chances are high that more new species will be found.
The Pselaphinae have almost 12,000 named species and subspecies (Newton pers. comm.) and is one of the
largest taxa of Coleoptera. Pselaphinae in numbers of species occupy after Carabidae: Trechinae and Leiodidae:
Cholevinae: Leptoderini third place amongst cave-dwelling Coleoptera. More than 250 species of Pselaphinae are
cavernicolous (Poggi et al. 1996, subsequent descriptions), and about one-quarter of these (63 species), are known
from the Balkan Peninsula, with the majority (60 species) from the Dinarides. Out of the Balkan Peninsula there
are other important areas where cavernicolous Pselaphinae were and still are regularly discovered. In the western
Palaearctic region this is Turkey, Caucasus (Hlaváč et al. 1999; Hlaváč & Faille 2018), Italy (Poggi 1982, 1990,
1992) and Pyrenees (Jeannel 1914; Besuchet 1974, 1985). Many species occur also in USA (Park 1951, 1960; Barr
1962; 1974; 1987; Besuchet 1982; Carlton & Cox 1999; Carlton 2008; 2012; Caterino & Vásquez-Vélez 2017), in
Japan (Tanokuchi 1985; Nomura 2012), and many new discoveries has been recently carried out in China (Nomura
& Wang 1991; Yin, Li & Zhao 2011; Yin & Li 2015; Yin, Nomura & Li 2015; Yin, Jiang & Steiner 2016; Yin &
Zhou 2018). Three cavernicolous Pselaphinae are known from Africa (Leleup 1956; Jeannel 1950) but none from
Australia and New Zealand. The study of neotropical cavernicolous Pselaphinae has been also recently initiated in
Brazil (Asenjo, Lopes-Ferreira & de Almeida Zanpaulo 2017; Asenjo, de Almeida Zanpaulo & Lopes-Ferreira
2018).
Most cavernicolous Pselaphinae belong to two supertribes, Batrisitae (Amauropini, Batrisini and
Thaumastocephalini) and Goniaceritae (Bythinini). Since the Amauropini have been tentatively rooted in the
Batrisini (Parker & Owens 2018), the tribes Batrisini and Bythinini hold the vast majority of cavernicolous
Pselaphinae globally, and this is also reflected in the cavernicolous pselaphine fauna of the Dinarides.
There are two dominant groups of strictly cavernicolous Pselaphinae in the Dinarides: the bythinine genus
Machaerites L. Miller, 1855 with 14 species and 1 subspecies, and the amauropine genus Seracamaurops Winkler
with 14 described species. The first genus is exclusively found in caves and pits of the Northern Dinarides in
Slovenia and Croatia, while the second genus is found in the Southern Dinarides, in Croatia, Bosnia and
Herzegovina, and Montenegro. These two parts of the Dinarides are recognized as local hotspots of subterranean
beetles (Zagmajster et al. 2008), due to habitat heterogeneity and historical climate stability (Bregović &
Zagmajster 2016). The cavernicolous species of Bryaxis Kugelann, 1794 (11 species) and Tychobythinus
Ganglbauer, 1896 (11 species), both Holarctic genera of Bythinini with numerous species, are found in all Dinaric
subregions. All other genera are endemic to a particular biogeographical subregion of the Dinarides, the most
species-rich being the amauropine genus Troglamaurops Ganglbauer 1903 (3 species) and the bythinine genus
Nonveilleria Pavićević & Besuchet, 2003 (2 species). All other genera, Thaumastocephalus as well as
Biokovobythus Pavićević & Ozimec, 2014, Bythoxenus Motschulsky, 1860, Gasparobythus Poggi, 1992, Grguria
Pavićević & Ozimec, 2014, Melledobythus Hlaváč, Nakládal & Jalžić, 2014, and Pauperobythus Nonveiller,
Pavićević & Ozimec, 2002 (all Bythinini) are monospecific, with the two remaining species, Protamaurops
montenegrinus (Székessy, 1943) and Decatocerus unicornis Winkler, 1913, the only cavernicolous species of their
otherwise endogean genera.
A logical question is: whether there is a pselaphine genus with a strong radiation in the Central Dinarides,
comparable to Machaerites in the Northern Dinarides and Seracamaurops in the Southern Dinarides? This paper
may provide the answer that there is such a genus, Thaumastocephalus. The discovery of the genus
Thaumastocephalus and the establishment of a new tribe for this genus was a significant staphylinidological and
coleopterological event, but more surprising is the fact that Thaumastocephalus is a relatively common
cavernicolous pselaphine that is widely distributed in the caves and pits of nearly the whole region of the Central
Dinarides. Together with seven species named here and six new species awaiting description in the museum of
Geneva (Cuccodoro, pers. comm.), there are at least 13 species of Thaumastocephalus, or even more due to a
further six caves from which we have sampled only females.
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FIGURES 13–15. 13—entrance to Jamica u docima (cave), type locality (photo Branko Jalžić); 14—Thaumatocephalus kirini
sp. n., habitus in situ (photo Boris Krstinić); 15—topographical map of the type locality (the finding place of the T. kirini sp. n.
is marked with red dots).
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FIGURES 16–18. 16—entrance to Jama pod Gažnovcem (cave), type locality (photo Petra Bregović); 17—Thaumatocephalus
marsici sp. n., habitus in situ (photo Petra Kutleša); 18—topographical map of the type locality (the finding place of the T.
marsici sp. n. is marked with red dots).
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FIGURES 19–21. 19—entrance to Bežurova jama (cave), type locality (photo Branko Jalžić); 20—peak Varda where the cave
is situated (photo Branko Jalžić); 21—topographical map of the type locality (the finding place of the T. rujnicensis sp. n. is
marked with red dots).
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FIGURES 22–24. 22—entrance to Jujnovića špilja (cave), type locality (photo Petra Bregović); 23—Thaumatocephalus
slavkoi sp. n., habitus in situ (photo Petra Bregović); 24—topographical map of the type locality (the finding place of the T.
slavkoi sp. n. is marked with red dots).
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FIGURES 25–27. 25—entrance to Sistem Velika -Velika (cave), type locality (photo Petra Bregović); 26—Thaumatocephalus
troglavi sp. n., habitus in situ (photo Petra Bregović); 27—topographical map of the type locality (the finding place of the T.
troglavi sp. n. is marked with red dots).
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FIGURES 28–29. 28—entrance to Dahna (cave), type locality (photo Jana Bedek); 29—Thaumatocephalus dahnae sp. n.,
habitus in situ (photo Jana Bedek).
FIGURE 30. Distribution map of the species of the genus Thaumastocephalus mentioned in this work.
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The discovery of two new species of Thaumastocephalus in Biokovo, T. slavkoi and T. troglavi (Fig. 30, red
and dark blue circles), is not surprising, because the subterranean fauna of this area has still not been systematically
investigated (Ozimec 2008). Only one other cavernicolous Pselaphinae has been recently described from this area,
Biokovobythus radici Pavićević & Ozimec, 2013. The fourth cavernicolous Pselaphinae recorded for this area is
Tychobythinus neumanni (Müller, 1909) (see Hlaváč et al. 2008, 2017). With altogether 21 subterranean
Coleoptera species (Hlaváč et al. 2017), and two more described here, Biokovo is the outstanding local hotspot of
subterranean biodiversity in the Dinarides. While this is a relatively high mountain of the biogeographical region of
the Central Dinarides, the species turnover of one genus along an elevational gradient (as seen in T. slavkoi and T.
troglavi) is not the first such case reported from this mountain. It is also known for a genus of Cholevinae, the
genus Leptomeson (Giachino et al. 2011). The locality of T. marsici is very near Biokovo (Fig. 30, green circle),
but it seems that the karst area in the proximity of Biokovo holds a unique subterranean fauna, which is also
supported by other subterranean groups (Antić et al. 2018, Dimitrijević & Rađa 2016). The type locality of T.
bilandzijae is situated in the lowlands near Mosor (Fig. 30, red square), and nearby there are two localities of T.
folliculipalpus: cave Vranjača and cave Trojama, both on Mosor (Ozimec et al. 2009, Poggi et al. 2001). Although
the female from cave Vranjača belongs to a new, still undescribed species (Besuchet, pers. comm.), it is not clear
that the specimen from cave Trojama will also belong to this new species. With the present state of our knowledge,
the distribution of T. folliculipalpus seems to be restricted to the Kozjak and Mosor (Fig. 30, green triangle).
However, further material of male specimens from Mosor will be needed to confirm its relationship with T.
bilandzijae or with a new, still undescribed species. The karst area near Metković, from where two new species
have been described, T. rujnicensis and T. kirini, is situated at the eastern border of the Central Dinarides (Fig. 30,
yellow circle and violet star). This is the first report of cavernicolous Pselaphinae from this area, whereas T. dahnae
is the first report of cavernicolous Pselaphinae from karst area near Tomislavgrad (Fig. 30, red triangle). We are
sure that further research in the Dinarides will yield more new taxa in the future, and reveal a fuller picture of the
biogeography of the genus.
Acknowledgements
We thank Jon Cooter (Oxford, England) for reading and correcting the manuscript. We also thank the following
reviewers of this paper: Donald S. Chandler, Michael S. Caterino and Joe Parker. The fieldwork was conducted
under permission issued by the Croatian nature conservation authorities (Ministry of Culture and Ministry of
Environmental Protection and Energy; license numbers: 532-08-02-03/1-07-2; 538-08-01-01/3-09-03; 538-08-01-
01/3-10-02; 532-08-01-01/1-11-02, 517-07-1-1-1-12-2; 517-07-1-1-1-13-4; 517-07-1-1-1-15-6; 517-07-1-1-16-4;
517-07-1-1-1-17-4). The authors wish to thank all collectors of the specimens: H. Bilandžija, R. Cvitanić, A.
Ćukušić, T. Čuković, J. Lakota, M. Lukić, A. Kirin, N. Kuharić, T. Rožman, V. Sudar and M. Vasić. Also, we are
grateful to all other members of Croatian Biospeleological Society who helped during fieldwork; T. Rožman for
improving quality of the images; R. Baković, A. Ćukušić, D. Lacković, M. Lukić, G. Rnjak, T. Rožman, V. Sudar,
P. Visković for preparing the topographical maps of the caves; and finally, the biospeleologists for helping with
distribution records and determination of the cave fauna—J. Bedek for Isopoda, T. Dražina for Diplopoda, M.
Lukić for Collembola, and M. Pavlek for Araneae. The fieldtrip to Bosnia and Herzegovina was partly funded by
the Whitley Fund for Nature (project leader Jana Bedek). We also wish to thank the Speleological Club Mijatovi
dvori from Tomislavgrad who helped with the fieldtrip to the cave Dahna. This work was supported by the Ministry
of Culture of the Czech Republic (DKRVO 2019-2023/5.I.a, National Museum, 00023272).
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... S3b), and many Euplectitae have asymmetrical aedeagi, including the type genus of the supertribe, Euplectus Leach and its relatives (e.g., Hlaváč & Nakládal 2014;Jałoszyński & Nomura 2021). The 'lack of parameres' in Thaumastocephalus was verified by Hlaváč, Bregović & Jalžić (2019), who described seven new species of this genus (all from caves in the Dinarides) and clearly illustrated asymmetrical, Euplectitae-form aedeagi with apically or subapically setose parameres. On the other hand, the condition of parameres in Batrisitae is highly variable, as Chandler (2001) mentions that "elongate paramere(s) [are] often fused to median lobe, one or both parameres may be fused or lost", and therefore the lack or presence of parameres is a poor indication for placing a genus either in Batrisitae or Euplectitae. ...
... Aedeagus (illustrated in Poggi et al. 2001 andHlaváč et al. 2019) elongate, with weakly sclerotized walls and asymmetrical median lobe bearing circular or oval dorsal diaphragm; endophallic sclerites elongate and asymmetric, parameres asymmetric, long and broad, with subapical and/or apical setae. ...
Article
Full-text available
A new genus and species, Percussiopalpus inusitatus Hlaváč & Jałoszyński gen. n. et sp. n. of the tribe Thaumastocephalini is described and illustrated, based on specimens collected in the Asian part of Turkey (Balıkesir Province). The discovery of a free-living pselaphine species morphologically closely resembling the obligate cavernicolous Thaumastocephalus Poggi et al. prompted a detailed morphological study of both genera, which results in transferring Thaumastocephalini from Batrisitae to Euplectitae.
... Comprising over 10,000 described species (Yin et al., 2019) and showing astounding morphological diversity, pselaphines have conquered a plethora of terrestrial habitats. They are most abundant in the upper soil layers (Chandler, 2001;Park, 1942Park, , 1947, but many groups have adapted to highly specialized conditions of deep soil (e.g., all Mayetiini and Imirini, some Bythinoplectini, Trichonychini, Trogastrini, Bythinini and Amauropini; Besuchet, 1980;Coiffait, 1955Coiffait, , 1958Jeannel, 1948), caves (e.g., all Thaumastocephalini, many Batrisitae, and some Metopiasini, Goniaceritae and Tyrini; Asenjo et al., 2017;Carlton, 2012;Hlav ač et al., 2019;Yin, 2020), or the similarly unique and demanding environment of ant colonies (all Clavigeritae, and many species in most tribes; e.g., Jałoszy nski et al., 2020; Luo et al., 2021). Among Pselaphinae, there are also highly interesting, relatively small lineages that include beetles morphologically rather uniform, and yet showing apparently various degrees or stages of adaptation to life in deeper soil layers and caves. ...
... A derived character state of B. saulcyi is the greatly reduced condition of the compound eyes. This is likely an autapomorphy of the genus, even though reduction of light sense organs occurs frequently in Pselaphinae, especially in cave-dwelling species and inquilines (e.g., Asenjo et al., 2017;Hlav ač et al., 2019;Jałoszy nski et al., 2020). ...
Article
The pselaphine Bergrothia saulcyi shows features seemingly linked with life in deep soil layers, such as greatly reduced and non-functional compound eyes, a sensorium of long tactile setae, long appendages, and flightlessness. However, the tiny beetles occur in forest leaf litter, together with a community of beetles with wings and well-developed eyes. We hypothesize that B. saulcyi moves into deep soil under dry conditions, and returns to upper layers when humidity increases again. Despite the evolutionary cost of a reduced dispersal capacity, this life strategy may be more efficient and less hazardous than moving to different habitats using flight and the visual sense in an environment periodically drying out. We also discuss cephalic features with potential phylogenetic relevance. Plesiomorphies of B. saulcyi include the presence of anterior tentorial arms, well-developed labral retractors, and a full set of extrinsic maxillary and premental muscles. Apomorphic cephalic features support cla-des Protopselaphinae + Pselaphinae, and Pselaphinae. A conspicuous derived condition , the clypeo-ocular carina, is a possible synapomorphy of Batrisitae and genera assigned to Goniaceritae. A complex triple set of cephalic glands found in B. saulcyi is similar to a complex identified in the strict myrmecophile Claviger testaceus (Clavigeritae). It is conceivable that glands linked with food uptake in free-living pselaphines were genetically reprogrammed in ancestors of inquilines, to enable them to appease the host ants. We suggest that behavioral studies are necessary to understand the poorly known life habits of B. saulcyi. Additional information is required to explain why a species with irreversibly reduced visual sense and other adaptations typical of endogean or cave-dwelling beetles was only collected from the upper leaf litter layer. K E Y W O R D S 3D reconstruction, micro/anophthalmy, micro-CT, musculature, rove beetles
... Comprising over 10,000 described species (Yin et al., 2019) and showing astounding morphological diversity, pselaphines have conquered a plethora of terrestrial habitats. They are most abundant in the upper soil layers (Chandler, 2001;Park, 1942Park, , 1947, but many groups have adapted to highly specialized conditions of deep soil (e.g., all Mayetiini and Imirini, some Bythinoplectini, Trichonychini, Trogastrini, Bythinini and Amauropini; Besuchet, 1980;Coiffait, 1955Coiffait, , 1958Jeannel, 1948), caves (e.g., all Thaumastocephalini, many Batrisitae, and some Metopiasini, Goniaceritae and Tyrini; Asenjo et al., 2017;Carlton, 2012;Hlav ač et al., 2019;Yin, 2020), or the similarly unique and demanding environment of ant colonies (all Clavigeritae, and many species in most tribes; e.g., Jałoszy nski et al., 2020; Luo et al., 2021). Among Pselaphinae, there are also highly interesting, relatively small lineages that include beetles morphologically rather uniform, and yet showing apparently various degrees or stages of adaptation to life in deeper soil layers and caves. ...
... A derived character state of B. saulcyi is the greatly reduced condition of the compound eyes. This is likely an autapomorphy of the genus, even though reduction of light sense organs occurs frequently in Pselaphinae, especially in cave-dwelling species and inquilines (e.g., Asenjo et al., 2017;Hlav ač et al., 2019;Jałoszy nski et al., 2020). ...
Article
The pselaphine Bergrothia saulcyi shows features seemingly linked with life in deep soil layers, such as greatly reduced and non-functional compound eyes, a sensorium of long tactile setae, long appendages, and flightlessness. However, the tiny beetles occur in forest leaf litter, together with a community of beetles with wings and well-developed eyes. We hypothesize that B. saulcyi moves into deep soil under dry conditions, and returns to upper layers when humidity increases again. Despite the evolutionary cost of a reduced dispersal capacity, this life strategy may be more efficient and less hazardous than moving to different habitats using flight and the visual sense in an environment periodically drying out. We also discuss cephalic features with potential phylogenetic relevance. Plesiomorphies of B. saulcyi include the presence of anterior tentorial arms, well-developed labral retractors, and a full set of extrinsic maxillary and premental muscles. Apomorphic cephalic features support cla-des Protopselaphinae + Pselaphinae, and Pselaphinae. A conspicuous derived condition , the clypeo-ocular carina, is a possible synapomorphy of Batrisitae and genera assigned to Goniaceritae. A complex triple set of cephalic glands found in B. saulcyi is similar to a complex identified in the strict myrmecophile Claviger testaceus (Clavigeritae). It is conceivable that glands linked with food uptake in free-living pselaphines were genetically reprogrammed in ancestors of inquilines, to enable them to appease the host ants. We suggest that behavioral studies are necessary to understand the poorly known life habits of B. saulcyi. Additional information is required to explain why a species with irreversibly reduced visual sense and other adaptations typical of endogean or cave-dwelling beetles was only collected from the upper leaf litter layer. K E Y W O R D S 3D reconstruction, micro/anophthalmy, micro-CT, musculature, rove beetles
... Comprising over 10,000 described species (Yin et al., 2019) and showing astounding morphological diversity, pselaphines have conquered a plethora of terrestrial habitats. They are most abundant in the upper soil layers (Chandler, 2001;Park, 1942Park, , 1947, but many groups have adapted to highly specialized conditions of deep soil (e.g., all Mayetiini and Imirini, some Bythinoplectini, Trichonychini, Trogastrini, Bythinini and Amauropini; Besuchet, 1980;Coiffait, 1955Coiffait, , 1958Jeannel, 1948), caves (e.g., all Thaumastocephalini, many Batrisitae, and some Metopiasini, Goniaceritae and Tyrini; Asenjo et al., 2017;Carlton, 2012;Hlav ač et al., 2019;Yin, 2020), or the similarly unique and demanding environment of ant colonies (all Clavigeritae, and many species in most tribes; e.g., Jałoszy nski et al., 2020; Luo et al., 2021). Among Pselaphinae, there are also highly interesting, relatively small lineages that include beetles morphologically rather uniform, and yet showing apparently various degrees or stages of adaptation to life in deeper soil layers and caves. ...
... A derived character state of B. saulcyi is the greatly reduced condition of the compound eyes. This is likely an autapomorphy of the genus, even though reduction of light sense organs occurs frequently in Pselaphinae, especially in cave-dwelling species and inquilines (e.g., Asenjo et al., 2017;Hlav ač et al., 2019;Jałoszy nski et al., 2020). ...
Article
Full-text available
The pselaphine Bergrothia saulcyi shows features seemingly linked with life in deep soil layers, such as greatly reduced and non‐functional compound eyes, a sensorium of long tactile setae, long appendages, and flightlessness. However, the tiny beetles occur in forest leaf litter, together with a community of beetles with wings and well‐developed eyes. We hypothesize that B. saulcyi moves into deep soil under dry conditions, and returns to upper layers when humidity increases again. Despite the evolutionary cost of a reduced dispersal capacity, this life strategy may be more efficient and less hazardous than moving to different habitats using flight and the visual sense in an environment periodically drying out. We also discuss cephalic features with potential phylogenetic relevance. Plesiomorphies of B. saulcyi include the presence of anterior tentorial arms, well‐developed labral retractors, and a full set of extrinsic maxillary and premental muscles. Apomorphic cephalic features support clades Protopselaphinae + Pselaphinae, and Pselaphinae. A conspicuous derived condition, the clypeo‐ocular carina, is a possible synapomorphy of Batrisitae and genera assigned to Goniaceritae. A complex triple set of cephalic glands found in B. saulcyi is similar to a complex identified in the strict myrmecophile Claviger testaceus (Clavigeritae). It is conceivable that glands linked with food uptake in free‐living pselaphines were genetically re‐programmed in ancestors of inquilines, to enable them to appease the host ants. We suggest that behavioral studies are necessary to understand the poorly known life habits of B. saulcyi. Additional information is required to explain why a species with irreversibly reduced visual sense and other adaptations typical of endogean or cave‐dwelling beetles was only collected from the upper leaf litter layer. This article is protected by copyright. All rights reserved.
... Only 12 species of Pselaphinae are recorded from the Central Dinarides, compared to 27 species from the Northern and 37 species from the Southern Dinarides (Hlaváč pers. database, see also Hlaváč et al., 2017;Hlaváč et al., 2019). As the Mt Matokit finding place of P. apfelbecki is very near the type locality of P. alenkirini n. sp., this record should be considered carefully and additional material from Mt Matokit should be collected to confirm its identity. ...
Article
Full-text available
Two new species of the subterranean and anophthalmous genus Paramaurops Jeannel, 1948 are described and illustrated: P. alenkirini sp. n. and P. struyvei sp. n. from Croatia. The distribution of all Croatian records of the genus is given in the map.
... Despite the fact that the cave beetle fauna of this region has been systematically studied for almost two centuries, new taxa are discovered and described almost every year Perreau and Pavićević 2008;Quéinnec 2008;Lakota et al. 2010;Lohaj and Lakota 2010;Casale et al. 2012;Hlaváč et al. 2019;Delić et al. 2020;etc.). ...
Article
Full-text available
Balkan's Dinaric karst is a continuous source of specialized, cave adapted fauna-so called troglobionts. Much of this exceptionally rich biodiversity is attributed to beetles (Coleoptera) from the ground beetle tribe Trechi-ni (family Carabidae), with hundreds of species with different degrees of morphological modifications for life in subterranean habitats-so called troglomorphism. In this contribution, we present a discovery of one new genus and species of subterranean Trechine beetle, Orcusiella prokletijensis gen. et sp. nov. from a high-altitude pit situated on Prokletije Mountains, Montenegro. This new taxon is described, illustrated and compared with closely related taxa. An identification key for all so far known genera of Dinaric subterranean Trechini, as well as data on the distribution and ecology of these remarkable species are provided and discussed.
... Despite the fact that the cave beetle fauna of this region has been systematically studied for almost two centuries, new taxa are discovered and described almost every year Perreau and Pavićević 2008;Quéinnec 2008;Lakota et al. 2010;Lohaj and Lakota 2010;Casale et al. 2012;Hlaváč et al. 2019;Delić et al. 2020;etc.). ...
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