Perreau & Pavicevic 2008.pdf

INSTITUTE FOR NATURE CONSERVATION OF SERBIA BELGRADE, 2008 Monograph n'22

Dragan PAV!CEV!C & Michel PERREAU {editors) Advances in the studies of the fauna of the Balkan Peninsula

Papers dedicated to the memory of Guido Nonveiller p. 215-239

THE GENUS HADESIA MULLER, 1911 AND THE PHYLOGENY OF ANTHROHERPONINA (COLEOPTERA, LEIODIDAE, CHOLEVINAE, LEPTODIRINI)

by Michel Perreau (1) & Dragan Pavicevic (2)

(1) Universite Paris 7 case 7139, 2 place Jussieu, 75251 Paris Cedex 05 France, e-mail: michel. perreau @univ-paris-diderot .fr

(2) Institute for Nature Conservation of Serbia, Dr. Ivana Ribara 91, 11070 Novi Beograd, Serbia, e-mail: [email protected]

SUMMARY. The genus Hadesia is revised. Two new species are described. Hadesia vasiceki weiratheri Zariquiey is upgraded to the rank of species. Several new localities are listed. Identification keys are given for the species of the genus Hadesia and for the genera of the subtribe Anthroherponina.

KEY WORDS. Coleoptera, troglobitic species, Bosnia & Herzegovina, Montenegro, Anthroherponina.

1- INTRODUCTION

The representatives of the genus Hadesia Muller, 1911 lives in Herzegovina and Montenegro. Up to now, it contained one species with two subspecies nominotypical Hadesia vasiceki vasiceki Muller, 1911, found only in a single cave: Vjetrenica pecina in Zavala and Hadesia vasiceki weiratheri Zariquiey, 1927, known from the type locality: Vojvode Dakovica pecina in Grahovo (Montenegro) and from jama Bravenik in Grab, near 'frebinje (Herzegovina) (PRETNER, 1974). Recent samplings in jama Bravenik and new localities in Croatia (spilja pod Gromackom Vlakom near Dubrovnik), Herzegovina (peCina Veliko Datlo in Korita) and Montenegro (Krivosije, Duboka jama near Knezlaz, and in Vodena jama in the Dragaljsko polje near Umac) make a revision of this genus necessary. The result is the increase the number of species of the genus to 4 with the description of two new species: one for the population of Veliko Datlo peCina and the other for the population of Jama Bravenik, and by the upgrade of the subspecies weiratheri Zariquiey to the rank of species.

2- METHODS

The microscopic slides of female genitalia have been obtained after treatment in KOH 0.1 N, coloration with Azoblack and inclusion in Euparal or DMHF. The photonic pictures were made on a microscope ZEISS axiolab with a digital camera SPOT. SEM pictures have been taken on a scanning electron microscope JEOL JSM 840A after metal coating. The phylogenetic analysis has been performed with the software PAUP.

The following abbreviations for collections and institutions are used: HZMZ: Hrvatski prirodoslovni muzej, Zagreb, Croatia; MZBS Museo de Zoologia, Barcelona, Spain; CACT: Achille Casale, Torino, Italy; CDCK: David Ceplik, Kosice, Slovakia; CDPV:

216 Advances in the studies of the fauna of the Balkan Peninsula

Dragan Pavicevic, Belgrade, Serbia; CGDY: Gejza Dunay, Kralovce, Slovakia; CJLK: Jan Lakota, Ruzomberok, Slovakia; CMPR: Michel Perreau, Paris, France; CPMG: Pier Mauro Giachino, Torino, Italy; CRLJ: Roman Lohaj, Kosice, Slovakia. Where necessary, the sign '/' separates different labels of a specimen.

3 - DESCRIPTION OR REDESCRIPTION OF THE TAXA

Genus Hadesia Muller Hadesia l'vliiller, 1911: 175, type species Hadesia Va.Sicceki Miiller, 1911 (monotypy)

DESCRIPTION. Length 6.7 to 7.6 mm. General shape of the body elongated, with narrow head and pronotum, and wide elytra, eyeless, wingless and weakly pigmented.

Head approximately 1. 7 times as long as wide (fig. 7). Surface strongly microreticulated, with some strong points on the occipital part only, with no occipital carina. Insertion of the antennae located before the first quarter of the length of the head. Second antennomere very short compared to the other antennomeres, much shorter than the first and the third ones. Labrum trapezoid with long setae on the dorsal side (fig. 7), and with a median notch on the anterior edge (fig. 13). Maxillae short with brushes of setae and sensillae both on the galea and the lacinia which is reduced (figs. 11 & 12). lVIaxillary palps with long first and second palpomeres, and a very short terminal palpomere (fig. 11). Labium with two rows of long setae on the ventral side and a high triangular longitudinal structure on the dorsal side (fig. 10). Labial palps extremely short, the second palpomere transverse (fig. 10). Mandibles angulate in dorsal view, the apex flat and vertical with 7 small teeth (figs. 8 & 9).

Thorax: pronotum narrow, slightly wider than the head, the widest part in the middle, 1.5 to 1.56 times as long as wide according to the species and the specimen, without a marginal lateral carina, but with a thin marginal anterior and posterior carina. Dorsal surface smooth, with no setae, with very tiny punctures and a superficial microreticulation, the macroscopic aspect shiny. Shoulder with a lateral deep groove (fig. 23). Scutellum wide and long, widely covered by the pronotum.

Elytra around two times as long as wide. Elytral sculpture rough with randomly disposed short (probably hydrophobic) setae, the macroscopic aspect matt and silky (fig. 28). Preapical region of the epipleuron more or less interrupted according to the sexes and the species. Humeral region of the epipleuron more or less thickened according to the species (figs. 24 & 25).

Abdomen: mesosterna! carina interrupted, the mesocoxal cavities widely confluent (fig. 26). First abdominal ventrite with a carina roughly parallel to the anterior edge (figs. 26 & 27), delimiting a smooth surface with a weaker microreticulation and a shiny aspect, between the anterior edge and the carina (fig. 27). Such a smooth plate exists also along the anterior margin of the other visible ventrites (fig. 26) but the carina occurs only on the first one.

Appendages: femora enlarged in the basal parts. Protibias without a longitudinal and lateral row of little spines and without an external apical spur. Mesotibias and meta tibias without an apical ring of spines of equal length. Em podium of tarsi with two setae. Claws widely dilated (fig. 29). MALE.

EJytra: preapical region of the epipleuron with a weak sinuosity. Abdomen: metasternum without lateral hollows as are in females. Appendages: all tarsi five segmented and not dilated.

M. Perreau & D. Pavicevic: The genus Hadesia Muller 217

Aedeagus: median lobe long, strongly curved, thick with a prepical widening (in dorsal view) and an apical expansion in lateral view (figs. 30 to 33). The thickness of the prepical part of the aedeagus is variable according to the species, varying from a strong hump (H. vasiceki Muller), to an hardly perceptible sinuosity (H. lakotai n. sp, H. weiratheri Zariquiey). The length of the apical expansion varies also according to the species. Internal sac with a long and slim stylus (figs. 34 to 37). Parameres thin, slightly shorter than the median lobe, with four setae, two short subapical ones and and two apical one which are variable in size. The size of the two apical setae varies within populations and cannot be used as a specific discriminant.

Genital segments: urite IX reduced to a ring, as most of Leptodirini.

FEMALE.

Elytra: preapical region of the epipleuron with a tooth followed towards the apex of the elytra by an indentation. The size of the tooth and the depth of the indentation are variable according to the species (figs. 14 to 16).

Abdomen: first abdominal visible ventrite with two deep lateral hollows (fig. 26).

Appendages: protarsi four segmented, meso and metatarsi five segmented. All tarsi not dilated.

Genital segments: ventrite VIII without expansion on the anterior edge. Urite IX with reduced sclerified parts, without appendicular parts (gonocoxites and gonosubcoxites, cf. Deuve, 2001 for terminology), with the epipleurites reduced to small circular lateral plates and the mediotergite reduced to a semicircular arc (fig. 21). Spermatheca long and curved, with the apex differently shaped according to the species (figs. 17 to 20).

Four species are presently recorded in the genus.

1. Hadesia vasiceki Muller Hadesia Va.Siceki Miiller, 1911: 175.

loc. typ.: Hohle bei Zavala (Herzegovina) [= pecina Vjetrenica] Hadesia Va.Siceki J. Miiller, 1911: Jeanne!, 1924: 426. Hadesia Va.Siceki Miill.: Winkler, 1925: 144. Hadesia Vasiceki Miiller: Fagniez, 1927: 23. Hadesia Va.Siceki J. Miiller, 1911: Remy, 1940: 2. (Biology) Hadesia vasiceki J. Miiller: Gueorguiev, 1990: 269.

DIAGNOSIS. Habitus in fig. 3; in addition to the characters of the genus: humeral region of the epipleuron thickened (figs. 23, 24). Female preapical indentation of the elytral epipleuron deep (fig. 14). Spermatheca regularly curved, rounded at the apex (fig. 17). Aedeagus with a strong dorsal preapical hump and a long apical expansion (fig. 30).

DISTRIBUTION. Bosnia-Herzegovina: Vjetrenica pecina in Zavala, in the Popovo polje. Croatia: spilja pod Gromackom Vlakom in Orasac not far from Dubrovnik (2 females seen in HZMZ).

DISCUSSION. Up to now H. vasiceki was only known from the type locality: Vjetrenica pecina. We have examined two females from spilja pod Gromackom Vlakom in Croatia (B. Jalzic, in HZMZ) which fit the main morphological diagnostic characters of this species.

2. Hadesia lakotai n. sp.

TYPE MATERIAL: holotype d', Bosnia and Herzegovina, peCina Velika Datlo in Korita, 21.06.2003, S. Ognjenovic leg. (CMPR). Paratypes: 1 d' and 2 ~' same data as holotype, S. Ognjenovic and J. Lakota leg. (CDPV, CJLK, CMPR); 1 ~' same cave, 13.09.2003 (CJLK); 1 d', same cave, 24.07.2005 (CJLK). All specimens have been found dead but in fairly good conditions.


Figure 1. Hadesia asamo n. sp. in jama Bravenik (photo: 1\!I. Perreau).

M. Perreau & D. Pavicevic: The genus Hadesia Miiller 219

Figure 2. The entrance of jama Bravenik (photo: M. Perreau).

ADDITIONAL MATERIAL: 1 <;J (debris) 23.06.2003 (CJLK); 2 <;J (debris) 24.07.2005 (CJLK); 1 d (debris) 11.08.2007 (C!viPR); two pairs of elytra (CDCK), all specimens in the same cave.

DIAGNOSIS. Habitus fig. 4; In addition to the characters of the genus: size slightly less than the other species (generally less than 7 mm whereas more than 7 mm for the other species). However this character is difficult to measure accurately as the known specimen are pieces of dead ones put together. Humeral region epipleuron thickened as in H. vasiceki Muller (fig. 24). Female preapical indentation of the elytral epipleuron deep and the preapical tooth projected below the edge of the epipleuron (fig. 15). Female with a spermatheca regularly curved and rounded the apex (fig. 18), as H. vasiceki IVIuller. Aedeagus long. strongly curved, with the basal and the apical regions making an acute angle, with a weak preapical hump and a long apical expansion (fig. 31).

DISTRIBUTION. Known only from the type locality.

ETYMOLOGY. The species is dedicated to Jan Lakota, who has collected some of the few known specimen.

3. Hadesia asamo n. sp. Hndesia vasiceki wciratheri Zariquiey: Pretner, 1974: 12. (pars) liadesia vasiceki weiratheri Zariquiey: Pretner. 1977: 114. fh;desia vasiceki weiratlwri Zariquicy: Gueorguiev, 1990: 270. (pars) 1-ladcsia orjensis: Gueorguiev, 1990: 270. (nomen nudum)

TYPE LOCALITY: Bosnia and Herzegovina: Jama Bravenik in Grab near Zupci (Zubacko polje), (nomen fictum vVeiratheri: peCina Vodenica)

TYPE MATERIAL: holotype d': Bosnia and Herzegovina: Jama Bravenik in Grab near Zupci (Zuhacko polje) 05/06/2004 (Cl\,IPR). Paratypes: 1 <? same data (CMPR); 1 d'and 1 <;J, same


Figures 3 to 6. Genus Hadesia: habitus of males. 3: H. vasiceki Muller. 4: H. lakotai n. sp. 5: H. asamo n. sp. 6. H. weiratheri Zariquiey. The scale bar is 1 mm.

M. Perreau & D. Pavicevic: The genus Hadesia Muller

Figures 7 to 13. Genus Hadesia, head (H. asamo n. sp.) and mouthparts (H. vasiceki Muller). 7: head dorsal view (scale bar= 0.5 mm). 8: left mandibule ventral view. 9: right mandibule dorsal view. 10: labium and hypopharinx dorsal side. 11: right maxilla and maxillary palp dorsal side. 12: right maxilla and maxillary palp, ventral side, with details of the apical palpomere and sensillae of the galea. 13: labrum and epipharinx ventral side. (figs. 8 to 13 from Casale & al., 2000.)

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Figures 14 & 15. Urite IX (genital segment) <j>. 14: Remyella sp. (the membraneous spermatheca is not visible). 15: Hadesia asamo n. sp. ( ep: epipleurites; tg: tergite; gs: gonosubcoxite; gc: gonocoxite; gt: gonostyles; sp: spermatheca). Scale bar is 0.2 mm.

JVI. Perreau & D. Pavicevic: The genus Hadesia Muller

Figures 16 to 22. Genus Hadesia female, apex of elytra (16 to 18, scale bar = 0.5 mm) and spermathecae (19 to 22, scale bar= 0.1 mm). 16 & 19: H. vasiceki Miiller. 17 & 20: H. lakotai n. sp. 18 & 21: H. asamo n. sp. 22: H. wciratheri Zariquiey (from Vodena jama).

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Figures 23 to 25. Humeral region of the right elytron. 23: H. vasiceki Miiller (g = lateral groove of the shoulder) (scale bar = 100 J-Im). 24: the detail of fig. 23, (hm = thickened humeral region of the epipleuron). 25. The same detail in H. asamo n. sp.

pit, 1910612003 (CMPR); 3d', same data as holotype (CGDY); 6 d' and 5 <j?, same pit, R. Mlejnek lgt. 18.06.2003 (CGDY, CJLK); 7 d' and 8 <j?, same pit, 05.06.2004, leg. S. Ognjenovic (CDPV); 1 d' and 1 <j?, same pit, 03.03.2006 leg. Miroslav Djokic (CDCK); 2 <j?, same pit, 03.03.2006 leg. Miroslav Djokic (CRLJ); 6 d' and 8 <j?, H6 Vodenica, Orjen Gau, without date n78-1666 to 78-1679 MZB (MZBS); 1 d' H6 Vodenica, Orjen Gau, without date (CMPR); 4 d' and 3 <j?, H6 Vodenica, Orjen Gau, without date (one male labelled as H. orjensis type) (CACT, CPMG). The 22 last paratypes with labels handwritten by Weirather.

DIAGNOSIS. Habitus fig. 5, in vivo fig. 1. In addition to the generic characters: humeral region of the epipleuron flat (fig. 25). Preapical indentation of the elytral epipleuron moderately deep, at least the apical tooth weak (and often also the preapical one) (fig. 16). Spermatheca regularly curved, with a preapical constriction and a sinuosity, narrowed at the apex (fig. 19). Aedeagus with a low preapical hump and a short apical expansion (fig. 32). DISTRIBUTION. Bosnia & Herzegovina, found only in the type locality: Jama Bravenik in Grab near Trebinje, (nomen fictum Weiratheri: peCina Vodenica). The entrance of the cave is shown in fig. 2. Details of this pit, including a bibliography and a topography are given in the report of expeditions (KuRTOVIC & al., 2008: 502). ETYMOLOGY. Asamo is the roman empire name of the town Trebinje.

4. Hadesia weiratheri Zariquiey stat. n. Hadesia vasiceki subsp. Weiratheri Zariquiey, 1927: 163. (holotype d' in MZBS)

loc. typ.: Hiihle 9, Dobra P. [= Vojvode Dakovica pecina], Lisac Gau Hadesia vasiceki weiratheri Zariquiey: Pretner, 1974: 12. (faunistic) (pars) Hadesia vasiceki weiratheri Zariquiey: Gueorguiev, 1990: 264.

TYPE MATERIAL: holotype (MZBS) with the following labels: typus [printed, black on red background] I H9 Dobra P. Lisac Gau zwischen Trebinje und NikSic, Siidwest Mont. [hand written] I Hadesia weiratheri [manuscript] Dr. Zariquey det. [printed] I 78 917 MZB [printed]

Jv1. Perreau & D. Pavicevic: The genus Hadesia Miiller

Figures 26 to 29. H. vasiceki Muller, abdomen, elytra and onychium. 26: ventral view of the abdomen (m = confluent mesocoxae, h = lateral hollow, c = adominal carina, sp = smooth plates) (scale bar= 1mm). 27: abdominal carina, detail of fig. 25. 28: elytralmicrosculpturc (scale bar = 0.05 mm). 29: onychium of the protarsus. Scale bar= 0.1 111111.

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STUDIED MATERIAL: holotype d': Montenegro, Vojvode Dakovica peCina in Grahovo (Grahovsko polje); 1 <jl, same locality, 15.04.2006 (CDCK); 1 d' (dead), same locality, 22.08.2007, leg. M. Plcca.S (CMPR); 1 <jl: Montenegro, Dragaljsko poljc, Umac env., 615 111 a.s.l., Vodna [Vodena] jama pit, 28.7.2004, lgt. R. Mlejnek (CGDY); 1 d': Montenegro, Krivosije, village of Knezlaz, Duboka jama, 600 111., 13.07.1997 leg. Dejan Vuckovic (CDPV)

DIAGNOSIS. Habitus fig. 6; in addition to the generic characters: humeral region of the epipleuron flat (fig. 25). Preapical indentation of the elytral epipleuron moderately deep as in H. asamo (fig. 16) but with some variability. Both characters similar as for H. asamo n. sp. Spermatheca regularly curved with the apical region not narrowed (fig. 20). Aedeagus without preapical hump and with a short apical expansion (fig. 33).

DISTRIBUTION. Montenegro: Orjen Mt. from the Grahovsko polje to the Krivosije. The holotype collected in 1926, one female collected in april 2006, and fragments of a male


collected in 2007 are the only known specimens from the type locality: Vojvode Dakovica pecina (nomen fictum Weiratheri: Dobra pecina) in the Grahovsko polje (PRETNER, 1974: WEIRATHER, 2006). Despite there being no sampling date written on the label of the holotype, nor in the description of Zariquiey, we know from the diaries of vVeirather that he collected the specimen during one of the three visits he made to Vojvode Dakovicain 1926, on the lOth, 11th and 13th of .July (\iVEIRATHER. 2006). Details of this cave, including a topography and bibliography are given in the report of expedition (KuRTOVIC & al.. 2008: 530). One female was collected in Vodcna jama which is the main sinkhole (ponor) of the Dragaljsko polje (TISSERAND & GODARD, 1970). One male has been collected in Duboka jama, in Kameno 1\Iore near Knezlaz in Krivosije. The five specimens are presumed to belong to the same species. The morphological characters are compatible with this hypothesis, but further sampling is necessary to get data on the geographical variability of the species.

4 - PHYLOGENY OF THE GENUS HADESIA MULLER

A morphological phylogenetic analysis of the species of Hadesia can be carried out with the following characters: morphology of humeral region of the elytra, shape of the female apical elytral notch, hump and apical tooth of the aedeagus. We considered Remyella and Leptodirus as outgroups. For an optimality criterion based on parsimony, two other characters: the apex of the spermathcca and the angle between the base and the apex of the median lobe are non informative. Indeed, they arc characters of the single H. asamo and H. lakotai respectively. However they are informative with an optimality criterion based on distances.

The parsimony analysis gives a well supported monophyletic group for H. vasiceki and H. lakotai (bootstrap 100 %). However, the possible sister group H. asamo-H. weiratheri is weakly supported (bootstrap 50%) because their shared characters are mainly plesiomorphic: humeral region of elytra fiat, short apical expansion of the aed~agus, and weak indentation of the female apex of elytra. This result seems paradoxical because both H. asamo and H. weiratheri are extremely similar and geographically localised in the same massif: Orjen (fig. 40). As a matter of fact, an analysis based on distances as optimality criterion, supports both groups H. vasiceki- H. lakotai and H. asamo-H. weiratheri as sister groups at the same level of bootstrap. In the present situation, the distances give a result more coherent with the biogeography than the parsimony, but molecular analysis would help to clarify the relationships of these four species.

Here we provide an identification key of the species:

1 - Humeral region of the cpipleuron thickened (fig. 24). Aedeagus with a longer apical expansion in lateral view (figs. 30 & 31). Preapical part of the female epipleuron with a strong tooth followed towards the apex by a deep indentation of the dytra (figs. 16, 17). Spermatheca with a rounded apex, without a preapical constriction (figs. 19, 20) . 2

- Humeral region of the epipleuron fiat (fig. 25). Aedeagus with a short apical expansion in lateral view (figs. 32 & 33). Preapical part of the female epipleuron with a strong tooth and a weak indentation (fig. 18) (with some variability) . 3

2 - SizE' generally more than 7 mm. Basal and apical parts of the aedeagus making an obtuse angle, median lobe with a high preapical hump in lateral view (fig. 30) vasiceki Miiller

- Size less than 7 mm. Basal and apical part of the aedeagus making an acute angle, median lobe without a preapical hump in lateral view (fig. 31) lakotai n. sp.

3 - Median lobe of the aedE'agus thicker and shorter in dorsal view (fig. 19). Spermatheca with preapical constriction and sinuosity, and a sharp apex (fig. 21) ... asamo n. sp.


Figures 30 to 37. Genus Hadesia, aedeagi, lateral view (30 to 33) and dorsal view (34 to 37). 30 & 34: H. vasiceki Miiller. 31 & 35: H. lakotai n. sp. 32 & 36: H. asamo n. sp. 33 & 37: H. weiratheri Zariquiey (from Duboka jama, Krivosije). Scale bar = 1 mm.


-Median lobe of the aedeagus more slender and in dorsal view (fig. 37). Spermatheca regularly curved with a rounded apex (fig. 22) . . . . . weiratheri Zariquiey

5- EVOLUTION AND PHYLOGENY OF THE SUBTRIBE ANTHROHERPONINA

Several morphological characters of whole or part of the subtribe Anthroherponina are of evolutionnary or biogical importance. We will emphasise some of them in this section.

5- 1 - The mouthparts

The mouthparts are strongly modified compared to most of the other species of Leptodirini. Similar morphologies have been observed to a more or less advanced degree of modifications in the genera Cansiliella Paoletti, 1972, Radziella Casale & Jalzic, 1988, Tartariella Nonveiller & Pavicevic 1999, Croatodirus Casale, Giachino & Jalzic, 2000 and Velebitodromus Casale, Giachino & Jalzic, 2004, Kircheria Giachino & Vailati, 2006. This special morphology has been illustrated by Jeanne! (1924) and more recently detailed scanning electron microscope illustrations and comparisons with other species have been published by Casale & al. (2000) and Moldovan & al. (2004). Some of these pictures are reproduced on figs. 8 to 13 with the courtesy of Casale. The most spectacular features are: 1 - the presence of very long and dense setae on labrum and labium (figs. 7 & 13), 2 - a rotation of the apex of the mandibles from an horizontal to a vertical position (figs.

8 & 9), 3 - very short third maxillary palpomeres. The second palpomere is at least 2.5 times

as long as the third, versus less than 1.8 time in other Leptodirini. All genera of Leptodirini in which the characters 1 and 2 have been observed live in a

similar biotope: the flow of running water leaking on the stalagmitic walls, sometimes called the 'cave hygropetric', which will be extensively discussed in the section devoted~to the biology. However, the character 3 is observed only in the hygropetricolous Anthroherponina (table I). This difference is very important on a phylogenetic ground as characters 1 and 2 are likely homoplasic as adaptative to the hygropetricolous conditions (they have been discarded from the phylogenetic analysis), while character 3 is possibly synapomorphic. Comparisons of the ratio of the length of the second palpomere to the length of the third one, between several genera of Leptodirini are given in table I, especially the comparison between hygropetricolous species and phylogenetically related non hygropetricolous ones.

The morphologies including characters 1 and 2 are generally referred to by authors as 'filtering mouthparts' (JEANNEL, 1924; CASALE & aJ., 2000; MOLDOVAN & al. 2004; CASALE & al., 2004) especially because of the numerous long setae. Our observations that the animal seems to brush the surface of stalagmitic walls on which it runs, by alternative moving of the head from right to left and left to right, suggest that it is eating deposits (algae?) off the surface of the wall. Then the so called 'filtering mouthparts' could be called 'brushing mouthparts' as well. Actually, no experimental protocol has ever been set out to verify these hypothesis, and the feeding pattern of the species of Hadesia (and of the other genera with similar mouthparts) is still unknown.

5- 2 - The female urite IX

The reduced shape of the IXth urite and especially the lack of the IXth appendages have been explicitely noticed in the genera Croatodirus and Velebitodromus by Casale

M. Perreau & D. Pavicevic: The genus Hadesia MUller

Figure 38. Right: lateral view of Leptodirus lwchenwarti Schmidt, head horizontal. Left: lateral view of Anthroherpon primitivum Absolon, head vertical, with elongation of the forehead (from Jeannel, 1924, modified).

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& al. (2000; 2004) and in Kircheria by Giachino & Vailati (2006). It occurs also in Anthroherpon, Leptomeson, and Paranthrophilon. Only one female Nauticiella stygivaga Moravec & Mlejnek, 2002 has been found and recently described (MLEJNEK & MORAVEC, 2003). Nothing is said about the evolution of the appendicular parts of the urite IX. The tergum IX alone is drawn and shows a reduction similar to that of the Hadesia, so that it is highly probable that a similar lack of appendicular parts occurs also in Nauticiella. In the genus Remyella however, the urite IX is fully developed (GIACHINO & ETONTI, 1995 and fig. 15).

5- 3 - The tarsal claws

Most genera of Anthroherponina have widened and apically blunt claws (fig. 29) which are very different from the thin and acute claws of most of Leiodidae.

Recently, Giachino & Vailati (2006) made a very interesting observation. Three specimens have been found of their new genus Kircheria, two are mature and one immature'. The two mature specimens have short and blunt claws, the immature one shows long and acute claws. This suggests that the blunt apical conformation results from the attrition of the claws in conditions which are presently unknown. The claws are still significantly wider than the usual conformation, but this observation decreases once more the phylogenetic importance of this character.

5- 4 - The antennae

Antennal insertions are located in the basal first quarter of the length of the head in Anthroherponina (fig. 7) (and also in Spelaobatina) and on the middle of the head in the other Leptodirini.

A functional interpretation of this character could be linked to the extreme dilatation of the abdomen correlated to the position of the insertion of the pro thorax on the abdomen. The figure 38 shows the lateral view of two genera each with an extremely dilated abdomen: Leptodirus and Anthroherpon. The difference is that the insertion of the prothorax is in a low position (close to the ground) for Leptodirus and in an upper position for Anthroherpon. In the latter case, the necessary position of the mouthparts, close to the ground to collect food, implies an elongation of the forehead.

The relative length of the two first antennomeres also clearly isolates the Anthroherponina. The second one is much shorter than the first one, compared to the other genera of Leptodirini in which it is at least nearly as long as the first one (table I).


Maxillary palpomeres Antennomeres length H ygropetricolous length ratio 2/1 ratio 2/1 ratio 3/1 Y=yes, N=no

Anthroherponina Hadesia vasiceki 2.65 0.35 1.92 y Kircheria beroni 2.55 0.67 1.88 y Nauticiella stygivaga 2.65 0.62 1.55 y Croatodirus bozicevici 2.60 0.65 1.25 y Velebitodromus smidai 2.60 0.60 1.00 y Leptomeson Jeonhardi 1.20 0.80 1.98 N Anthroherpon Jatipenne 1.60 0.65 1.84 N Anthroherpon harbichi 1.25 0.68 2.05 N Paranthrophilon speJaebatoides 1.40 0.55 1.72 N Leptodirina Remyella scaplwides 1.75 0.90 1.58 N Razdiella styx 1.75 1.25 1.38 y Leptodirus hochenwarthi 1.70 3.87 2.67 N Pholeuina Pholeuon angusticolle 1.50 1.43 0.90 N Spelaeobatina Spelaeobates kraussi 1.42 1.16 1.30 N Bathysciina Cansiliella servadeii 1.55 0.91 1.39 y Lessiniella trevisioJi 1.40 1.09 1.19 N Aphaotus jureceki 1.35 1.03 1.05 N Tartariella dunnitorensis 1.60 1.37 1.63 y Leonhardella angulicollis 1.70 1.22 1.05 N

Table I. Comparison of the relative lengths of the second to the third maxillary palpomeres, of the second to the first antennomeres and of the third to the first antennomeres in Anthroherponina and in the hygropetricolous genera of Leptodirini with some phylogenetically related non hygropetricolous species.

5-5- Phylogeny

Jeannel (1924) used the combination of the position of the antennal insertions and the enlarged and apically blunt claws to characterize the tribe "Anthroherponini" composed of "phyletic line of Anthroherpon" (today: subtribe Anthroherponina) and "phyletic line of Spelaeoba.tes" (today: subtribe Spelaeobatina). As a matter of fact, at this time, no other genus than those of Anthroherponini (sensu Jeannel) have a similar morphology of claws and antennal insertions.

However several other genera were subsequently discovered with similar claws in other subtribes of Leptodirini: Ca.nsiliella., Razdiella ...

Moreover J eannel ( 1931) himself placed the genus Remyella in Anthroherponini in spite of its usual conformation of claws. This placement of Remyella in Anthroherponina was based on the insertion of the antennae on the posterior third of the head ( JEANNEL, 1931). However, Remyella differs from the other genera of Anthroherponina by several important characters, not only the fully developed female urite IX, but also the relative size of the two first antennomeres, and the already mentioned shape of the claws.

Laneyrie (1967) and Gueorguiev (1974; 1976) strictly followed the point of view of Jeannel on this topic.

Later, a first tentative of cladistic analysis of the whole tribe of Leptodirini (CASALE et al., 1991) showed that Spelaeobatina should presumably be more closely related to


Bathysciini than Anthroherponina, breaking the previously assumed monophyly of both Anthroherponina and Spelaeobatina, and decreasing the phylogenetic importance of the position of antennal insertions. Perreau (2000) conserved the results of the work of Casale et al. on these subtribes in a slightly different framework.

Therefore, at the present time, the strongest apomorphy on which a monophyletic group could be established is the reduction of the appendicular parts of the female urite IX. However this would lead to the exclusion of Remyella from Anthroherponina. This exclusion is also supported by two other characters in which the character state of Remyella and the other genera of Anthroherponina are different: tarsal claws and relative length of the second to first antennomere.

To test this hypothesis, we performed a preliminary phylogenetic analysis with the genera of Anthroherponina and with Pholeuon as outgroup, using the 11 following characters:

1 - female urite IX normal ( outgroup, Remyella) versus reduced (other genera) 2- female ventrite VIII with an anterior apophysis (spiculum ventrale) (outgroup,

Remyella, Anthroherpon, Leptomeson, Paranthrophilon) versus without (Hadesia, Kircheria, Nauticiella, Velebitodromus, Croatodirus)

3- spermatheca normally sclerified (outgroup, Hadesia, Nauticiella, Paranthrophilon, Leptomeson, Anthroherpon) versus weakly sclerified (Remyella, Croatodirus, Velebitodromus, Kircheria)

4- aedeagus with internal sclerified structures (Remyella, Pholeuon, Leptomeson, Kircheria, Hadesia) versus inermous (other genera)

5 - abdominal ventrites with glabrous plates (Hadesia, Kircheria) versus without glabrous plates (other genera)

6- female first ventrite with lateral hollows (Hadesia, Kircheria) versus without (other genera)

7- pronotum with a sparse punctuation and erected setae (Hadesia, Anthroherpon, Leptomeson, Paranthrophilon) versus a close punctuation and recumbent setae (Pholeuon, Remyella, Nauticiella, Velebitodromus, Croatodirus, Kircheria)

8 - Third maxillary palpomere short: the second one at least 2.5 times as long as the third one as on figs. 11 & 12 (Hadesia, Kircheria, Croatodirus, Velebitodromus, Nauticiella) versus long: the second one at most 1.8 times as long as the third one ( outgroup, Remyella, Anthroherpon, Leptomeson, Paranthrophilon)

9 - Second antennomere approximately as long as the first one ( outgroup, Remyella), versus much shorter than the first one (other genera)

10- Third antennomere less than 1.3 times as long as the first one ( Croatodirus, Velebitodromus), versus at least 1.5 times as long as the first one (other genera)

11 - Claws narrow ( outgroup, Remyella, Anthroherpon, Leptomeson, Paranthrophilon) versus wide as in fig. 29 ( Hadesia, Kircheria, Croatodirus, Velebitodromus, Nauticiella)

The only mostly parsimonious tree obtained from this analysis is shown in fig. 39 with bootstrap values.

A first well supported group (bootstrap: 94%) is the set of genera of Anthroherponina without Remyella. This confirms the necessary exclusion of Remyella from Anthroherponina. (For this reason, Remyella is set between square brackets in the identification key given in the next section.)


Pholeuon

Remyella

85 I Hadesia

I Kircheiia 72

Nauticiella

94 87 I I

Croatodirus

Velebitodromus

Leptomeson

I Paranthrophilon

I Anthroherpon

Figure 39. The most parsimonious tree with bootstrap values obtained in the phylogenetic analysis of the genera of Anthroherponina with Pholeuon as outgroup.

Two other well supported groups (bootstrap respectively of 85 % and 87 %) are the set Hadesia-Kircheria corresponding to the 'phyletic lineage' of Hadesia (GIACHINO & VAILATI, 2006), and the set Croatodirus- Velebitodromus.

A fourth group is less neatly supported (bootstrap: 72 %) : the set of hygropetricolous genera (Hadesia, Kircheria, Nautriciella, Croatodirus, Velebitodromus). This result deserves more investigation, since the hygropetricolous evolution is likely homoplasic (but the clearly adaptative mouthparts have been excluded from the set of characters used in the analysis). The potential sister group (if so) of the preceding one: the set Anthroh$erpon, Leptomeson, and Parathrophilon is inconsistently supported with a bootstrap of 56 %.

Two other groups are also inconsistently supported with bootstraps around 50 %: the set Nauticiella, Croatodirus, Velebitodromus and the set Anthroherpon-Parathrophilon.

Actually, this preliminary analysis does not give any information about a new placement for Remyella. A phylogenetic analysis of all genera of Leptodirini is outside the scope of this paper, but the following discussion is a preliminary attempt to find the placement of Remyella.

In previous publications, Leptodirina are generally characterised by the following set of characters: male protarsi five segmented; apex of tibias without external spur; protibias with a lateral external row of spines; mesotibias and metatibias with an apical comb of spines, and insertions of the antennae on the middle of the head (JEANNEL, 1924; GuEORGUIEV, 1974; GUEORGUIEV, 1976; PERREAU, 2000). However, as previously noted (PERREAU & PAVICEVIC, 2008), this characterisation can no longer be retained after obvious observations. Indeed, many genera presently included in Leptodirina do not have the protibial lateral external row of spines, for example: Astagobius Reitter, Balcanobius Gueorguiev, Rozajella Curcic & et al., Nonveilleriella Perreau & Pavicevic, Parapropus Ganglbauer, Radziella Casale & Jalzic, Spelaeodromus Reitter (list not exhaustive). Gueorguiev himself, in the description of Balcanobius, clearly noticed this absence (GuEORGUIEV, 1965). So the absence of the protibial lateral external row of spines in Remyella is not an impediment to its placement in Leptodirina. Moreover all


other characters of Leptodirina, except the position of antenna! insertions, are present in Remyella including the absence of the tibial apical external spurs.

These observations support the tentative placement of Remyella in Leptodirina. Within the new recognition of the subtribes, the probability of monophyly of Anthroherponina is greatly increased, but the hypothesis of monophyly of Leptodirina is significantly weakened. Further investigation of the phylogeny of the latter is necessary to clarify its actual status.

5-6 - Identification table of the genera of Anthroherponina

The following identification table of the genera of the subtribe Anthrohetponina (sensu novo) modifies and updates the table given by Moravec & Mlejnek (2002) for the former Anthroherponina.

1 -Female with a fully developped IXth urite (fig. 14). Second antennomere as long as the first one. Claws narrow ....................... : ........ [Remyella Jeannel]

-Female IXth urite reduced, without appendicular parts (fig. 15). Second antennomere much shorter than the first one. Claws wide (fig. 29) (Anthroherponina sensu novo) . 2

2 -Second maxillary palpomere at least 2.5 times as long as the third one (figs. 11 & 12). 'Hygropetricolous' species ........................................... 3

- Second maxillary palpomere at most 1.8 times as long as the third one. Species not 'hygropet~icolous' ................................................. 7

3 - Large size, more than 6 mm. Abdominal ventrites with a transversal glabrous plate along the anterior edge (fig. 26: sp). Female first abdominal ventrite with two deep lateral hollows (fig. 26: h). Internal sac of the aedeagus with a longitudinal stylus (figs. 32, 33, 36, 37) . . . . . . . . . . . . . . . . . . . . . . . . . . . ........................ 4

- Small size, less than 4 mm. Abdominal ventrites without a transversal glabrous plate. Female first abdominal ventrite without deep lateral hollows. Internal sac of the aedeagus inermous ....................................................... 5

4 - Pronotum with a sparse ponctuation and with long erected setae, the macroscopic aspect shiny. Second antennomere approximately 0.35 times as long as the first one. Shoulders with lateral grooves (fig. 24: g). Apex of the epipleuron of elytra interrupted by an indentation, deeper in females (figs. 16 to 18) than in males . ........ Hadesia Muller

- Pronotum with a close ponctuation and short recumbent setae, the macroscopic aspect matt. Second antennomere approximately 0.65 times as long as the first one. Shoulders without lateral grooves. Apex of the epipleuron of elytra continuous ............ .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Kircheria Giachino & Vailati 5 - Pronotum wide, less than 1.3 times as long as wide, with lateral margins. Third

antennomere less than 1.3 times as long as the first one ...................... 6 - Pronotum narrow, at least 1.6 times as long as wide, without lateral margins. Third

antennomere approximately 1.5 times as long as the first one ............... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nauticiella Moravec & Mlejnek

6 - Aedeagus strongly curved in lateral view ...... Croatodirus Casale, Giachino & Jalzic - Aedeagus straight in lateral view . . . . . . . . Velebitodromus Casale, Giachino & Jalzic

7 - Apex of parameres with three setae. Internal sac of the aedeagus without sclerified structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8

- Apex of parameres with four setae. Internal sac of the aedeagus with sclerified structures . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Leptomeson Jeannel

8- Head widely oval, distinctly wider than the pronotum ....... Parantrophilon Noesske - Head cylindrical, at most slightly wider than the pronotum .... Anthroherpon Reitter


6 - GEOGRAPHICAL DISTRIBUTION

The distribution areas are shown on the map of fig. 40. H. vasiceki is located in the Popovo Polje and in Croatia near Dubrovnik, H. lakotai lives in a single cave near Korita, far more north-east than H. vasiceki. H. weiratheri is located in the central part of Orjen and in the eastern Krivosije (the eastern slope of Ledenice) and H. asamo lives in a single pit in the Zubacko polje, on the north-western part of Orjen, not far from Trebinje.

7- BIOLOGY AND HABITAT: "THE CAVE HYGROPETRIC"

The biotope of the genus Hadesia differs in many points from that of most of the troglobitic species of Leptodirini and was described a long time ago (JEANNEL, 1924: REMY, 1940). Adults are always found in the vicinity of strong flows of running water on stalagmitic walls, often inside the film of running water itself, even when it is a few millimeters thick and the flow especially strong. It seems to look for food in this stream of water as discussed above in the description of the mouthparts.

Outside the stream of running water, Hadesia walk as every other species of Coleoptera. However, inside the water flow, the Hadesia turn their back to the water flow (head towards the floor) and the walk is either perpendicular to the axis of the body (like a crab), or climbing up backwards inside the water stream. It is not established whether this position is made mandatory by the feeding process by bringing water to the ventral side of the mouth, or if it is a position which is more stable to hang on to the wall in the flow of water, or even if it corresponds to any other function.

According to this habitat, Hadesia always require huge inputs of water in the cave. But in Jama Bravenik (type locality of H. asamo), based on our own observations since 2003, the abundance of Hadesia is clearly correlated to the rate of flow in the pit. The fluctuations of running water is one parameter which can explain why some populations are so uncommon. In Vojvode Dakovica peCina (type locality of H. weiratheri), 6nly two specimens have been collected since 1926: the holotype in July in 1926, the second one in April 2006. In July 1926, a very high level of precipitation was recorded on Orjen, more than twice the average precipitation for July from 1888-1960, and one of the five wettest Julys during this period (fig. 41, data from RIDANOVIC, 1966). Of course, this precipitation data is recorded in Crkvice, at about ten kilometers away from Grahovo where the records began only after 1950. The precipitation is much less in Grahovo than in Crkvice (fig. 40, and RADOVANOVIC & al., 2008), but the comparison over the period 1950-1960 shows that both series are highly correlated, so probably before as well. In 2006, despite no objective measurements of the precipitation being available (the meteorological stations are no longer active), we observed in summer an exceptionally high flow of water in the cave. The second low passage of the cave (cf. topography page 530) was filled with water and changed to a siphon, so that the deepest part of the cave was not accessible. During the many visits we had made previously to the cave, the conditions of dryness did not allow the sampling of any specimen. It should be noted that the level of precipitation for July is not generally directly correlated with the annual precipitation (fig. 41). For instance, the high precipitation level in July 1926 does not correspond to an annual maximum in 1926. The annual maximum occurred in 1927, the second wettest year during the period 1888-1960, surpassed only in 1937, and hardly equated in 1910 and 1958. However it should be taken into account that the hydrogeological conditions of the Grahovsko Polje were not the same, during all the 20th century, as they are today. The history and hydrogeology

M. Perreau & D. Pavicevic: The genus Hadesia Muller

Figure 40. Distribution map of the genus Hadesia Muller. 1: pecina Vjetrenica (type locality of H. vasiceki Muller). 2: spilja pod Gromaekom Vlakom (H. vasiceki Muller). 3: pecina Datlo (type locality of H. lakotai n. sp.) 4: jama Bravenik (type locality of H. asamo n. sp.) 5: Orjen and Krivosije (H. weiratheri Zariquiey): peCina Vojvode Dakovica (type locality); Vodena jama and Duboka jama.

235

236

700

600

500

400

300

200

100

0

1885

Advances in the studies of the fauna of the Balkan Peninsula

, _._ Crk~ite ann~i112 , ---il- Crkvice july i -Crkviceaverageofjuly

_ -=7E-Q~ovo ~UI_~ __

1895 1905 1915

1927

·'\

1925 1935 1945 1955 1965

Figure 41. Graphics of the precipitations on Orjen, annual (reported to one month) and for July, in millimeters (stations of Crkvice and Grahovo, data from RIDANOVIC, 1966).

of the Grahovsko Polje and Vojvode Dakovica peCina have been extensively described by Petrovic and Gavrilovic (1986). The drainage of the Grahovsko Polje occurred in two steps. Before 1904, Grahovsko Polje was periodically flooded by water and the Vojvode Dakovica pecina represented an estavelle. Water run from the cave several times a year which corresponded to the inundation of the Grahovsko Polje. From 1904, when the river Grahovska Reka was channelled, the inundations occurred less frequently and were l€lss devastating, and the water run from the cave also less frequently. The second step occurred in 1962 when a dam was built, draining all the water of the river Grahovska reka into an aqueduct and ceasing definitely the flow thorough the river bed. The water run for the last time from the Vojvode Dakovica cave in October 1962. Today, the water conditions in this cave are dramatically different than before 1962 and even more different than before 1904 (but at this time H. weiratheri has not yet been discovered). Consequently, any conclusions based only on meteorological fluctuations are at risk of being irrelevant. H weiratheri was perhaps more common in the Vojvode Dakovica peCina while it represented an estavelle and the records were missing only due to the lack of sampling.

This habitat of strong, fast and thick running water over the surface of stalagmitic walls has been referred to by some authors as "the cave hygropetric" (SKET, 1979; 2004). This attempt to identify a new biotope was based essentially on the fact that several species of different zoological groups are living in similar conditions as the Coleoptera of the genera Cansiliella, Tartariella, Radziella, Velebitodromus; the Collembola: Pseudosinella cabidochei Deharveng & Gouze, P. bessoni Deharveng, Bessoniella procm·a Deharveng & Thiebaut (SKET, 2004), and the observation that coleopterans living in these conditions have convergent morphologies of mouthparts (see the above descriptions).

Little attention was paid to the fact that such conditions occur seasonally or even sporadically. As a matter of fact, in all caves where Hadesia are living (except perhaps Vjetrenica in which these conditions exist nearly permanently), the hygropetric conditions


are never permanent. They are linked to the presence of the stream of water which is highly variable according to the period of the year (jama Bravenik), and often totally absent for many years (Vojvode Dakovica peCina).

Some authors noticed this fact: "Such [water] films are only seldom really permanent ... " (SKET, 2004). However nobody discussed the critical questions which arise from this lack of permanency: where do the Hadesia (and other species living in similar conditions) live when hygropetric conditions are not present in the cave? Are these conditions maintained in other unaccessible parts of the karst in which the Hadesia could live in the same way as observed when possible? Or are the Hadesialiving in other yet unknown special conditions? What are the exact relations between hygropetric conditions and the mouthparts morphologies? This last question being also linked to the problem of the feeding pattern of the hygropetricolous species. These questions remain open, and up to now have not been addressed on a reliable experimental ground. Yet their answers are mandatory to give an actual and relevant characterisation to the 'hygropetric' conditions.

8 - ACKNOWLEDGMENTS

We thank Sinisa Ognjenovic and Jan Lakota, first collectors of the new species H. lakotai, M. Popovic for the drawings; G. Maso and 0. Escola (MZBS) for the facilities to study the Zariquiey collection; A. Casale (Sassari, Italy) for the permission to reproduce the figures 8 to 13; R. Ozimec and B. Jalzic (Zagreb, Croatia) for the loan of two specimen from the spilja pod Gromackom Vlakom; P. M. Giachino (Torino, Italia) for the loan of specimens of Croatodirus, Velebitodromus and Kircheria; G. Dunay (Kralovce, Slovakia) for the loan of the specimen from Vodena jama; D. Ceplik (Slovakia) for the loan of the second known specimen of H. weiratheri from Vojvode Dakovica; R. Mlejnek for the loan of the female of Nauticiella stygivaga; D. Kurtovic and the members of the speleological group ''Zelena brda" of Trebinje (Bosnia-Herzegovina) for their collaboration in the field. We are also indebted to E. Qw§innec (University Paris 6, France) for useful discussions, to I. le Disquet (University Paris 6) for electronic microscopy facilities, and to IUT Paris Jussieu (University Paris 7) for the photonic microscopy facilities. A special thank toM. Schulke (Berlin, Germany) for his help in solving some tricky translations from ancient German.

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Documents

Perreau & Pavicevic 2008.pdf