New Acanthopetalum from Bulgaria 155

International Journal of Myriapodology 2 (2008) 155-170 Sofi a–Moscow

A new of Acanthopetalum Verhoeff , 1900 (Diplopoda: : ) from Bulgaria, with a review of the A. richii (Gray, 1832) group

Pavel Stoev National Museum of Natural History, Tsar Osvoboditel Blvd. 1, 1000 Sofi a, Bulgaria. E-mail: [email protected]

Abstract Th e genus Acanthopetalum (Callipodida: Schizopetalidae) comprises 11 species distributed from the southern Apennines in the west to the Aegean coast of Turkey in the east. A single species, A. carinatum (Brandt, 1840), has been recorded from Bulgaria (Rhodopes Mts.). Recent study of freshly collected material coming from the same area has revealed that A. carinatum is erroneously recorded for the country, and the genus is actually represented in the Rhodopes by a new species, Acanthopetalum rhodopinum sp. n., whose description is provided. It is well distinguished from the other congeners by the presence of two sharp processes on the coxa of male leg-pair 7, and processes m, n and g on the inner branch of femoroid. New illustrations of and male leg-pair 7 of A. albidicolle (Verhoeff , 1900) and A. richii and an identifi cation key for all species of the A. richii (Gray, 1832) group are provided for better distinction of the new species. A. richii is herewith reported for the fi rst time from Turkey; its presence there might be due to human introduction.

Key words , , Acanthopetalum rhodopinum sp. n., superfi cial underground compartment, Rho- dopes Mts., Bulgaria

Introduction Th e northern part of the Mediterranean region – the Iberian, Apennine, and Balkan peninsulas, and Asia Minor – together with southern North America and southeast- ern Asia constitutes one of the three primary centers of diversifi cation of the milli- pede order Callipodida. In this vast and highly diverse area the order is known from 75 (sub-)species, 18 (sub-)genera and 3 families – Callipodidae, and

© Koninklijke Brill NV and Pensoft, 2008 DOI: 10.1163/187525408X395922

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Schizopetalidae (Shear et al. 2003; Stoev et al. 2008). Th e genera Eurygyrus C.L. Koch, 1847 and Acanthopetalum Verhoeff , 1900, with 18 and 11 species, respectively, outnumber all the remaining genera in the region. Since the beginning of the 20th century, Acanthopetalum (Callipodida: Schizopetali- dae) has been treated in numerous taxonomic, systematic and faunistic works (for a complete review see Stoev et al. 2008). Undoubtedly, the most serious contributions are due to Karl W. Verhoeff who, besides establishing the genus, also described several new species mainly from Greece and Turkey (e.g., Verhoeff 1900, 1901, 1903, 1929, 1940, 1941), and Carl Attems who added new species from the same area (e.g., Attems 1902, 1903, 1935). In more recent times, Hoff man & Lohmander (1964) and Hoff man (1973) commented on the status of some of the old species, provided new illustrations of A. koss- wigi (Verhoeff , 1940) and described new subspecies of A. hamatum (Attems, 1903) from Turkey. In a series of publications in the 1960s and 1970s Karl Strasser revised the genus diagnosis, re-described several taxa, proposed new synonyms and other taxonomic alter- nations, and erected the subgenus Patalysium Srasser, 1976 to accommodate the species of the A. carinatum (Brandt, 1840) group (cf. Strasser 1967, 1970a, 1970b, 1971, 1973, 1974, 1976). Recently, Mauriès et al. (1997) described a new species from Albania and revised the subgenus Patalysium Srasser, 1976, proposing the synonymy of all the species known therein under A. carinatum (Brandt, 1840). Ćurčić et al. (2001) resurrected the full species rank of A. minotauri (Attems, 1902), and Mauriès & Karamaouna (1984) reported A. hoplites Strasser, 1973 from the Greek islands Paros and Naxos. Th e A. richii (Gray, 1832) group was defi ned (under the name of A. sicanum Ber- lese, 1883 group) by Verhoeff (1933), who considered it to comprise the following species: A. sicanum, A. verhoeffi i Strasser, 1933, A. argolicum Verhoeff , 1900 and A. albidicolle (Verhoeff , 1900). Strasser (1970b) proposed the synonymy of Lysiopetalum argolicum, A. verhoeffi i, L. argolicum montivagum Verhoeff , 1901 and Acanthopetalum argolicum epiroticum Attems, 1935 under A. sicanum. Jeekel (2000) revived the long forgotten statement of Pocock (1893) that Lysiopetalum anceps Latzel, 1884 and “very possibly” L. sicanum are synonyms of Lysiopetalum richii (Gray, 1832). Th is view was adopted by Enghoff & Kime (2007) in the most contemporary species’ list of Euro- pean millipedes, and by Stoev et al. (2008) in the world catalogue of Callipodida. In Bulgaria a single species of Acanthopetalum has so far been recorded, i.e., A. (Petalysium) carinatum (Brandt, 1840) found in the Rhodopes Mts. (Ceuca 1973). Being generally confi ned to the western part of the Balkan Peninsula, and showing signifi cant diff erences from the other species in the genus in the presence of a large un- ciform process on the gonopod femoroid (on in Strasser 1974, fi g. 43), its occurrence in the country raised some concerns that led Strasser (1975) to express doubts about the accuracy of Ceuca’s identifi cation. Recent study of freshly collected material com- ing from the same area has confi rmed Strasser’s suspicion, revealing that A. carinatum is erroneously recorded from Bulgaria. Th e genus is actually represented in the Rhodopes by a new species, Acanthopetalum (s.str.) rhodopinum sp. n. belonging to the richii spe- cies group. To be consistent with my previous descriptions of schizopetalid millipedes (e.g. Stoev & Enghoff 2003, 2004; Stoev 2007) and for better separation of the new

Downloaded from Brill.com09/28/2021 02:42:07PM via free access New Acanthopetalum from Bulgaria 157 species, I also provide illustrations of the gonopods of A. albidicolle, based on topotypic specimens from Corfu Island, and A. richii, based on topotypic specimens from Malta and additional material from Turkey and Greece.

Material and methods Part of the material of the new species was collected by traps specially designed for catching invertebrates inhabiting the deeper soil layers and the crevices of maternal rock (the so called mesovoid shallow substratum (MSS) or superfi cial underground compart- ment). Traps were made from PVC pipe with diameter of 8 cm and length of 80 cm and were set vertically in holes dug in a limestone spot overgrown with bushes and Pinus nigra. One hundred and eight holes with diameter 8 mm were drilled at 10 cm distance from the pipe’s lower end. A plastic cup, tied to a polythene rope, and fi lled with a solu- tion of ethyleneglycol with a few drops of formalin was put at the bottom of the pipe. Traps were covered by solid plastic covers in order to avoid penetration of superfi cial fauna and infi ltration of water during heavy rains. All specimens are preserved in 70% ethanol. Drawings were made with the aid of a camera lucida mounted on a Wild micro- scope, type 181300. Close up photos were taken under an Olympus SZH 10 research microscope with an Olympus U-PMTVC Q-Color 3 camera, and were processed using the program Adobe Photoshop CS2. A complete chronological list of citations related to the species of the A. richii group can be found in Stoev et al. (2008).

Abbreviations: NHMW - Naturhistorisches Museum Wien; ZMB - Museum für Naturkunde, Ber- lin; SMF - Senckenberg Museum Frankfurt; NMNHS - National Museum of Natural History, Sofi a; HT - Holotype; PT - Paratype/s; ST - Syntype/s

Taxonomic part Family Schizopetalidae Verhoeff , 1909 Genus Acanthopetalum Verhoeff , 1900 Subgenus Acanthopetalum Verhoeff , 1900 Th e Acanthopetalum richii group

Diagnosis: Moderately large callipodidans (ca. 30-60 mm), composed of 44-47 pleu- rotergites (PTs), coxa of 7th male leg-pair modifi ed; gonopods in situ extending well beyond pleurotergal margin; tibiotarsus reduced, process a never extending to the level of the solenomere. Currently valid species: A. richii, A. albidicolle, A. subpatens Mauriès, Golovatch & Stoev, 1997, and A. rhodopinum sp. n. Th e status of A. albidicolle aetolicum (Verhoeff , 1903) is uncertain.

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Distribution: Th e species of the richii group occur in the Italian provinces Venezia and Puglia and the islands Sicily and Lipari; Malta, incl. island of Gozo; ; , incl. island of Cres; Mainland Greece and Aegean coast of Turkey (A. richii); Greek island of Corfu and Epirus, southern Albania (A. albidicolle); western Albania (A. subpatens) and southern Bulgaria (A. rhodopinum) (Fig. 21). Remark: For complete synonymic lists of the species referred to the group see Stoev et al. (2008).

Acanthopetalum albidicolle (Verhoeff , 1900) Figs 1-3, 15.

Lysiopetalum (Acanthopetalum) albidicolle Verhoeff , 1900, Zool. Jahrb. Abt. Syst. Oekol. Geogr. Tiere, 13: 55, Taf. IX, fi gs 38-39. ST: ♂, ♀, juv. (NHMW), ♂, ♀, juv., microscope slides (ZMB) from Corfu, Greece. Lysiopetalum albidicolle aetolicum Verhoeff , 1903, Arch. Naturgesch., 69 (1): 147, fi gs 1-2 (see also p. 151). ST: 2 ♂♂, 2 ♀♀ (NHMW), 2 ♂♂, 3 ♀♀, 4 juv. (ZMB) from Stoliko in Aetolien, Greece (see Stagl & Stoev 2005 for the origin of the types). Acanthopetalum albidicolle and Acanthopetalum albidicolle aetolicum: Stoev et al., 2008, Zootaxa, 1706: 19.

Material examined: adult male, “Lysiopetalum albidicola [sic!], Corfu, Verhoeff , 1900” (SMF, No. 1635).

e ps f

e n d fo tt k sg j sb 123p n k a k

p

Figures 1-3. A. albidicolle, gonopods: 1 – posterior view, 2 – mesal view, 3 – lateral view. Abbreviations: a – femoroidal process “a”, cx – coxa, d – thickening of solenomere branch, e – tibiotarsal process “e”, f – tibiotarsal apex, fo – tibiotarsal fold, j – spine on solenomere branch, k – process “k” of solenomere branch, n – process “n” of solenomere branch, p – coxal process, ps – parasolenomere, sb – solenomere branch, sg – seminal groove, tt – tibiotarsus. Scale line: 1 mm.

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Remarks: Th is species is well distinguished from the other species in the richii group by having the male 7th coxa heavily expanded ventrad, devoid of processes (Fig. 15), the gonocoxa having a subtriangular process, and the femoroid bearing a long process k which is crossing in situ with the opposite process (Fig. 1). Th e species can be easily rec- ognized also by its white collum contrasting with the generally darker brown-yellowish body. A. albidicolle has hitherto been recorded only from the Greek island of Corfu (type locality) and southern Albania (three records from a cave near Himarë, and from Llogora and Ducali). Th e subspecies aetolicum is known from the region of Epirus in Greece, but its status should be confi rmed by examination of the type specimens or further material from northwestern Greece. Th e poorly known Lysiopetalum erberi L. Koch, 1867, also described from Corfu (L. Koch 1867), is considered by Verhoeff (1900) as a possible senior synonym of A. albidicolle (see also Stoev et al. 2008).

Acanthopetalum richii (Gray, 1832) Figs 4-9, 14.

Craspedosoma Richii Gray, 1832, In: Griffi th and Pidgeon, Th e Insecta arranged by Baron Cuvier, pl. 135, fi g. 4. Type (sex and whereabouts unknown) from Malta. Acanthopetalum richii: Stoev et al., 2008, Zootaxa, 1706: 21.

Material examined: adult ♂ and ♀, “Acanthopetalum richii, H. Enghoff det.”, Malta, Rdum-il-Bies, 10.3.1983, P.S. Schembri leg. (NMNHS, ex coll. Natural History Mu- seum of Denmark); adult ♂, “Lysiopetalum argolicum, K. Verhoeff det. 1900.” Argos, Greece (SMF No. 1639); 2 ♂♂, ♀, “Makri, Asieminor” [presently Fethiye city in west- ern Turkey], (ZMB). Remarks: In spite of being the oldest species in the genus (and one of the oldest in the whole order), the true identity of A. richii remains uncertain. Th is is mainly due to the fact that there are no contemporary illustrations of the gonopods based on topo- typic material from Malta. All taxa that have hitherto been synonymized under richii, namely anceps, verhoeffi i, argolicum, argolicum montivagum, argolicum epiroticum show certain diff erences in the shape of gonopod tibiotarsus (cf. Strasser 1970b). To fi ll this gap and to show the morphological variability between diff erent populations, here I provide illustrations of male gonopods and leg-pair 7 based on topotypic specimens from Malta and others from Greece (identifi ed by Verhoeff as A. argolicum) and Turkey. Th ere are no signifi cant diff erences in the shape of male leg 7 (Fig. 14) between the examined specimens. However, the tibiotarsus in Greek specimens has three terminal processes subequal in length (Fig. 5, f, f2, f3) and a much longer spear-shaped process e (Fig. 5). Th e Maltese and Turkish specimens have only two processes – a larger one (f) and a smaller subterminal one (f2, Figs 7, 9). However, the shape of process e in the former is triangular, while in the later it is serrated (Figs 7, 9). In all other respects the gonopods of the three quite distantly located populations resemble each other. Th e specimens from Fethiye show gonopod shape similar to that of A. sicanum, A. verhoeffi i

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f e f2 f3 d fo t ps j n k k 4 a 5

p

f e f2 fo sg j j d t k

a k

p

6 7 p

f e f2 fo j d t j

a k k 8 9 p

p

Figures 4-9. A. richii: 4-5, Argos, Greece, SMF; 6-7, Rdum-il-Bies, Malta, NMNHS; 8-9, Fethiye, Tur- key, ZMB: mesal (fi gs 4, 6, 8) and lateral (fi gs 5, 7, 9) views of gonopod, respectively. Abbreviations: a – femoroidal process “a”, e – tibiotarsal process “e”, d – thickening of solenomere branch, f – tibiotarsal apex, f2 – process “f2” of tibiotarsus, f3 – process “f3” of tibiotarsus, fo – tibiotarsal fold, j – spine on solenomere branch, k – process “k” of solenomere branch, n – process “n” of solenomere branch, p – coxal process, ps – parasolenomere, sg – seminal groove, t – process “t” of solenomere branch. Scale lines: 1 mm.

Downloaded from Brill.com09/28/2021 02:42:07PM via free access New Acanthopetalum from Bulgaria 161 and A. argolicum epiroticum (cf. Strasser 1933, fi gs 1-2; Verhoeff 1939, fi gs 5-6; Attems 1935, fi gs 9-10), and it is very likely that the species has been imported in Turkey by human activities, most likely with a cargo from the Apennines. Th is formally represents a new country record.

Acanthopetalum rhodopinum sp. n. Figs 10-13, 16-20.

Lysiopetalum carinatum: Ceuca 1973, Bull. Inst. Zool. Mus. Sofi a 38: 242, fi g. 1, misidentifi ca- tion! See also Strasser (1975). Acanthopetalum sp. n.: Stoev et al. 2008, Zootaxa 1706: 40. Previous records: Rhodopes: Narechenski bani (Ceuca 1973).

Material examined: Holotype: adult ♂ with 44 or 45 PTs + telson; Bulgaria, Western Rhodopes Mts., Karadzhov Kamak Protected Area, N 41°49.367, E 24°57.015, 1410 m alt., under stones, 22.4.2006, B. Petrov leg. (NMNHS); – Paratypes: subadult ♂ with undeveloped gonopods, 44 PT + telson, same locality, date and collector (NMNHS); 2 adult ♂♂, one ♂ with 45 PT + telson, 1 subadult ♂ with undeveloped gonopods, 44 PT + telson, 1 juvenile with 38 PT + telson, Western Rhodopes Mts., Laki, approx. 1100 m from the crossroad to village of Borovo in the direction of Belitsa; left side of the road, small rocky valley overgrown with bushes and Pinus nigra, underground trap, N41°50.340, E24°51.564, 666 m alt., 17.7.2006-1.4.2007, B. Petrov leg. (NMNHS, one adult ♂ in SMF); one subadult ♂, 44 PT + telson, same locality and collector, 1.4- 25.11.2007 (NMNHS). Diagnosis. Diff ers from the other species in the richii group by the presence of two sharp processes on coxa of male leg-pair 7, by the reduced and twisted gonocoxal proc- ess p, which crosses the opposite gonopod’s coxal process, and by the specifi c shape of processes g, n, m, k and h on gonopods. Etymology. From mountain Rhodopes, the type locality. Description. Maximum length ca. 60 mm, maximum width ca. 3.4 mm, 44-45 PT + telson. In cross-section PT higher than broad (ratio: 1.6 x 1.4). PT 4 with a small ante- rioventral hook. Body colour: generally dark brown; a yellow, submoniliform mid-dorsal stripe from second PT to body end. Prozonites light brown-grayish; metazonites with posterior dark brown band. Pleurotergal crests darker than remaining part of pleuroter- gite. Ozopores encircled by a lighter, yellowish spot. Antennae long, extending slightly beyond the posterior margin of PT 7 when folded backward (Fig. 13). All antennomeres dark brown. Frontal part of head in males strongly concave, without setae. Long, sparse setae only on labrum. Epicranium uniformly dark brown, frontal concavity marbled with lighter yellowish spots; a lighter yellowish stripe above labrum. Ocellaria subtriangular, transparent, lying on a black background, composed of 41 ocelli in 7 rows. Tömösváry’s organ bigger than an ocellus, situated closer to anterior side of ocellaria than to antennal base. Legs brown. Hypoproct tripartite, median sclerite largest, subtrapezoidal, bearing

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Table 1. Partial chaetotaxy in A. rhodopinum

Anterior setae Posterior setae Collum a, b, c, d, e + a, b, c, d, e 2nd PT a, b, c, d, e + a, b, c, d, e 3rd PT a, b, c, d, e + a, b, c, d, e 4th PT a, b, c, d, e + a, b, c, d, e 5th PT a, d + a, d, b, c, e + b, c, e 6th PT a, e + a, eb, c, d, f + b, c, d, f 7th PT a, b, c, d, e, f + a, b, c, d, e, f a pair of macrosetae situated in the middle. Lateral sclerites smaller, subtriangular, with one seta each. Paraprocts subdivided into smaller dorsal and a bigger ventral plates, each bearing a pair of long setae. Spinnerets long, each ending with a seta their length.

s

h ps f e r fo g tt sb

n j g n k 10 11 12 p k m a sg k p

13

Figures 10-13. A. rhodopinum: 10-12, gonopods: 10 – posterior view, 11 – mesal view, 12 – lateral view; 13 – left antenna, lateral view, setation not shown. Abbreviations: a – femoroidal process “a”, e – tibiotarsal process “e”, f – tibiotarsal apex, fo – tibiotarsal fold, g – tibiotarsal process “g”, h – tibiotarsal tooth, j – tooth of solenomere branch, k – process “k” of solenomere branch, m – spear-like process “m”, n – process “n” of solenomere branch, p – coxal process, ps – parasolenomere, r – ridge of solenomere branch, s – so- lenomere, sb – solenomere branch, sg – seminal groove, tt – tibiotarsus. Scale lines: 1 mm.

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Crests on anterior 1-5 PTs poorly developed, more pronounced on middle and poste- rior PTs; 18 crests between ozopores on 15th PT. Ozopores visible from PT 6 to 5th PT from caudal end, situated between crests 9 and 10 on midbody PTs. Chaetotaxy: as in table 1. Male fi rst and second leg-pairs markedly shorter, third slightly shorter than subse- quent legs. Tarsus of leg-pairs 1-3 single; bipartite from 4th to ultimate pair. Pads con- fi ned to the distal part of tarsi on leg-pairs 4-11. All legs densely covered with long fi ne setae. Coxae of leg-pairs 5 and 6 with ventromedian processes, pointing anteriad (Fig. 17); similar process but less pronounced on leg-pair 4; coxa of leg-pair 7 with one acumi- nate ventromedian and one ventrolateral processes (Figs 16, 18). Coxa of leg-pair 9 and following legs unmodifi ed. Prefemora of leg-pairs 4-11 swollen, more so on leg-pair 7. Male gonopods (Figs 10-12, 19-20): in situ extending well beyond pleurotergal mar- gin; sternum reduced, gonocoxae connected through a chitinized sternal lamina. Coxae with a long, sigmoid, apically pointed coxal process (p) arising from the anteromedian side of coxa; p subequal in length to prefemoroid, directed ventrad, crossing opposite coxal process in situ. Femoroid composed of smaller inner (solenomere branch, sb) and larger outer branch (tibiotarsus, tt), the latter partly entwining the inner process from the anterior side. A long, but apically blunt process (a) arising basally from median side of femoroid, curving slightly anteriad and then posteriad, ending in close proximity to process g of tibiotarsus. Tibiotarsus with pointed apex (f) and serrated median margin,

14 15

17

16

Figures 14-17. 14-16 – male leg-pair 7, anterior view, setation not shown: 14 – A. richii (Greece), 15 – A. albidicolle, 16 – A. rhodopinum; 17 – male leg-pair 6 of A. rhodopinum, anterior view, setation not shown. Scale lines: 1 mm.

Downloaded from Brill.com09/28/2021 02:42:07PM via free access 164 P. Stoev / International Journal of Myriapodology 2 (2008) 155-170 ending basally with a long, subfalcate process (g) directed medioposteriad; apical part of tibiotarsus strongly incised to form blunt process (e) at the point where the tibiotarsal fold (fo) starts; a tiny pointed tooth (j) on the inner surface of solenomere branch close to the junction with tibiotarsus; a similar one (h) on the inner surface of tibiotarsus, just below the fold. Solenomere branch with two large black processes at base (k and n), arising from its posteriomedial side: the larger process (k) distally falcate, pointing dorsomediad, with a spear-like process (m) on its inner surface, pointing medioventrad; the smaller process (n) apically blunt, pointing dorsolaterad. Solenomere (s) slightly larger than parasolenomere (ps). Solenomere branch enlarged just below the division point of solenomere and parasolenomere forming a short and rounded ridge (r). Female: unknown. Remarks: A. rhodopinum was hand-collected under stones and also caught in traps at 60-70 cm depth in the soil in limestone areas. Th ough encountered at considerable depth it lacks modifi cations for subterranean life, like e.g. compact body, shortened legs and antennae, faded coloration and eye reduction. Th e Rhodopes region is center of origin of the genus Balkanopetalum Verhoeff , 1926, closely related to Acanthopetalum, with 4 species known from the area (Stoev & Enghoff 2003; Stoev et al. 2008). Unlike the species of Balkanopetalum, which generally live in caves, A. rhodopinum is hitherto registered to inhabit habitats outside caves and most likely is an euedaphibiont.

Acanthopetalum subpatens Mauriès, Golovatch & Stoev, 1997

Acanthopetalum subpatens Mauriès, Golovatch & Stoev, 1997, Zoosystema 19 (2-3): 264, fi g. 4 A-F. Male HT (NMNHS), ♂ and ♀ PT (MNHN) from a cave on the road Permet- Leskoviku, Leskoviku District, Albania. Acanthopetalum subpatens: Stoev et al. 2008, Zootaxa 1706: 22.

Material examined: 2 paratype ♂♂, Albania, Leskoviku District, cave on the road Per- met-Leskoviku, 5 km before Leskoviku, 900 m, 12 May 1995, P. Stoev & B. Petrov leg. (NMNHS). Remarks: Th is species is known only from its type locality in southern Albania, close to the Greek border. By having a single median process on male 7th coxa and coxal process broadened at base and abruptly narrowing in the middle, it is most closely re- lated to A. richii. However, the tibiotarsus in subpatens has a serrated median margin, while in richii it is smooth.

Discussion Th e subgenus Acanthopetalum comprises 10 species (15 subspecies) distributed from the southern Apennines in the west to the Aegean coast of Turkey in the east (Stoev et

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18

19 20

Figures 18-20. A. rhodopinum: 18 – coxa, trochanter and prefemur of male leg-pair 7, anterior view; 19 – gonopod, mesal view; 20 – gonopod, posterolateral view. Without scale.

Downloaded from Brill.com09/28/2021 02:42:07PM via free access 166 P. Stoev / International Journal of Myriapodology 2 (2008) 155-170 al. 2008). On the basis of gonopod shape three groups of species can be distinguished: 1) the furculigerum group characterized by the presence of well-developed uncinate process on the outer surface of tibiotarsus (here A. furculigerum (Verhoeff , 1901) with 4 subspecies, A. minotauri (Attems, 1902) and A. hamatum (Attems, 1903)); 2) the blanci group having the femoroidal process a extending to the level of solenomere (here A. cycladicum (Verhoeff , 1901), A. blanci (Brolemann, 1932) and A. mendeli- cum Strasser, 1974); 3) the richii group showing a comparatively reduced tibiotarsus, and process a never extending to the level of the solenomere. Acanthopetalum hoplites Strasser, 1973 from the Greek islands Paros, Antiparos and Naxos, has a strongly reduced tibiotarsus and uncinate process, and the expanded basal part of solenomere branch bearing 5 processes; it most likely constitutes an independent phyletic line closely related to the furculigerum group. According to Strasser (1976) the species of the furculigerum group are distributed mainly along the western coast of Turkey, on the Dodecanese Island of Kos, Crete and southern part of Peloponnese, while the blanci group is restricted to the Macedonia and Attica provinces of Greece, and the Cycladic island Syros. With the exception of some introductions, the richii group shows a typical trans-Adriatic distribution. Noteworthy are the absence of the group in the central part of the Balkans and its presence only in the southernmost part of the Italian Mainland (Fig. 21). Being generally troglo- and petrophilous, the species of A. richii group are usually found in the aphotic zone of caves, deeply underground in limestone areas, or under stony debris. A. richii is undoubtedly the most euryecious of all. Besides from caves,

2 2 2

3

2 2 1 4 2 1 1 2 2 2 1 2 2 2 2 2 2

2

Figure 21. Distribution map of the A. richii group: 1 – A. albidicolle, 2 – A. richii, 3 – A. rhodopinum, 4 – A. subpatens.

Downloaded from Brill.com09/28/2021 02:42:07PM via free access New Acanthopetalum from Bulgaria 167 on Malta it has been found also from leaf-litter in woodlands, clay slopes, garrigue, in- side cellars and also from specifi c rocky habitats called Rdum and Widien (Enghoff & Schembri 1989). Th e species is tolerable to anthropogenic infl uence and often occurs in human settlements, sometimes in substantial quantity. Th e fi nd of A. rhodopinum in the mesovoid shallow substratum in Bulgaria is the fi rst record of a callipodidan from this particular environment. Its absence in the well-prospected Rhodopean caves can be explained with the time of colonization of the caves by callipodidans. A. rho- dopinum most likely invaded the Rhodopes later than the ancestors of Balkanopeta- lum, which had penetrated into the caves of this region earlier.

Key for identifi cation of the species of the A. richii group 1 (2) male 7th coxa with two sharp processes (Figs 16, 18); gonocoxal process sig- moid, crossing opposite process in situ; inner (solenomere) branch of femoroid with processes m, n and g; Rhodopes Mts., Bulgaria...... A. rhodopinum 2 (1) male 7th coxa either without or with a single sharp, median process (Figs 14, 15); gonocoxal process straight; processes m, n and g absent ...... 3 3 (4) male 7th coxa heavily expanded ventrad, without sharp process (Fig. 15); gono- coxal process subtriangular (Fig. 2); femoroidal process k very long, pointing dorsad, crossing opposite process in situ (Fig. 1); collum white; Greek island of Corfu, Greek Mainland (subspecies aetolicum) and southern Albania ...... A. albidicolle 4 (3) male 7th coxa less or almost not expanded, always with one sharp median pro- cess (Fig. 14); gonocoxal process broadened at base, abruptly narrowing in the middle and curving anteriad distally (Figs 4, 6, 8); femoroidal process k much smaller, never crossing opposite member; collum usually yellow-brownish ....5 5 (6) tibiotarsus with serrated median margin and a subtriangular process e; femo- roidal process k single, tooth-like; cave near Leskoviku, Albania (see Mauriès et al. 1997, fi g. 4) ...... A. subpatens 6 (5) median margin of tibiotarsus smooth, with 2-3 processes (Figs 5, 7, 9); fe- moroidal process k forked (Fig. 4); Apennine and Balkan peninsulas, Malta, possibly introduced in Turkey ...... A. richii

Acknowledgements I would like to thank Henrik Enghoff (ZMUC, Denmark) and Rowland Shelley (North Carolina State Museum of Natural Sciences, USA) for their numerous linguis- tic corrections and critical reviews on the fi nal draft, and Boyan Petrov (NMNHS) for collecting the material of the new species. Th is paper was made possible with funding from the Bulgarian Ministry of Education and Science, Th e National Science Fund project No. B-1523/05. Part of the work was undertaken also with support from the

Downloaded from Brill.com09/28/2021 02:42:07PM via free access 168 P. Stoev / International Journal of Myriapodology 2 (2008) 155-170

Deutsche Forschungsgemeinschaft in 2007, and the Field Museum of Natural History, Chicago (FMNH, Bass Scholarship) in 2006. I greatly acknowledge the help of Peter Jäger and Petra Sierwald in the course of research visits to the SMF and FMNH.

References

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