Eupolybothrus cavernicolus Komericki & Stoev, 2013

Stoev, Pavel, Komericki, Ana, Akkari, Nesrine, Liu, Shanlin, Zhou, Xin, Weigand, Alexander M., Hostens, Jeroen, Hunter, Christopher I., Edmunds, Scott C., Porco, David, Zapparoli, Marzio, Georgiev, Teodor, Mietchen, Daniel, Roberts, David, Faulwetter, Sarah, Smith, Vincent & Penev, Lyubomir, 2013, Eupolybothruscavernicolus Komericki & Stoev sp. n. (Chilopoda: Lithobiomorpha: Lithobiidae): the first eukaryotic species description combining transcriptomic, DNA barcoding and micro-CT imaging data, Biodiversity Data Journal 1, pp. 1013-1013 : 1013

Eupolybothrus cavernicolus Komericki & Stoev, 2013 sp. n.

Materials

Type status: Holotype. Occurrence: recordedBy: M. Lukić; individualCount: 1; sex: male; lifeStage: adult; Location: country: Croatia; stateProvince: Knin; locality: NP Krka, village Kistanje, Hydroelectric power plant Miljacka, cave Miljacka II ; verbatimElevation: 115 m; verbatimLatitude: 44°00'01.1''N; verbatimLongitude: 16°00'58.5''E; Event: samplingProtocol: hand collected under clay sediment; eventDate: 9 February 2013; Record Level: institutionCode: CBSS; collectionCode: CHP536 Type status: Paratype. Occurrence: recordedBy: A. Komerički; individualCount: 1; sex: female; lifeStage: adult; Location: country: Croatia; stateProvince: Knin; locality: NP Krka, village Kistanje, Hydroelectric power plant Miljacka, cave Miljacka II ; verbatimElevation: 115 m; verbatimLatitude: 44°00'01.1''N; verbatimLongitude: 16°00'58.5''E; Event: samplingProtocol: hand collected under lump of clay; eventDate: 18 October 2012; Record Level: institutionCode: CBSS; collectionCode: CHP517 Type status: Paratype. Occurrence: recordedBy: M. Lukić; individualCount: 1; sex: male; lifeStage: adult; Location: country: Croatia; stateProvince: Knin; verbatimLocality: NP Krka, village Kistanje, Hydroelectric power plant Miljacka, cave Miljacka II; verbatimElevation: 115 m; verbatimLatitude: 44°00'01.1''N; verbatimLongitude: 16°00'58.5''E; Event: samplingProtocol: hand collected; eventDate: 9 February 2013; Record Level: institutionCode: BGI Type status: Paratype. Occurrence: recordedBy: H. Cvitanović & A. Komerički; individualCount: 1; sex: female; lifeStage: adult; Location: country: Croatia; stateProvince: Knin; verbatimLocality: NP Krka, village Kistanje, Hydroelectric power plant Miljacka, cave Miljacka II; verbatimElevation: 115 m; verbatimLatitude: 44°00'01.1''N; verbatimLongitude: 16°00'58.5''E; Event: samplingProtocol: hand collected; eventDate: 18 October 2012; Record Level: institutionCode: NMNHS; collectionCode: NMNHS-CHILOPODA-1/2013 Type status: Paratype. Occurrence: recordedBy: A. Kirin & A. Komerički; individualCount: 1; sex: female; lifeStage: adult; Location: country: Croatia; stateProvince: Knin; verbatimLocality: NP Krka, village Kistanje, Hydroelectric power plant Miljacka, cave Miljacka II; verbatimElevation: 115 m; verbatimLatitude: 44°00'01.1''N; verbatimLongitude: 16°00'58.5''E; Event: samplingProtocol: hand collected; eventDate: 4 May 2010; Record Level: institutionCode: ZMUC; collectionCode: zmuc00029439 Type status: Paratype. Occurrence: recordedBy: M. Lukić; individualCount: 1; sex: male; lifeStage: adult; Location: country: Croatia; stateProvince: Knin; verbatimLocality: NP Krka, village Kistanje, cave Miljacka IV (= Špilja kod mlina na Miljacki); verbatimElevation: 115 m; verbatimLatitude: 44°00'12.8''N; verbatimLongitude: 16°01'08.8''E; Event: samplingProtocol: hand collected; eventDate: 4 May 2010; Record Level: institutionCode: ZMUC; collectionCode: zmuc00029440 Type status: Paratype. Occurrence: recordedBy: A. Kirin; individualCount: 1; sex: female; lifeStage: subadult; Location: country: Croatia; stateProvince: Knin; verbatimLocality: NP Krka, village Kistanje, Hydroelectric power plant Miljacka, cave Miljacka II; verbatimElevation: 115 m; verbatimLatitude: 44°00'01.1''N; verbatimLongitude: 16°00'58.5''E; Event: samplingProtocol: hand collected; eventDate: 4 May 2010; Record Level: institutionCode: CBSS; collectionCode: CHP420 Type status: Paratype. Occurrence: recordedBy: A. Komerički; individualCount: 1; sex: damaged female; lifeStage: subadult; Location: country: Croatia; stateProvince: Knin; verbatimLocality: NP Krka, village Kistanje, cave Miljacka IV (= Špilja kod mlina na Miljacki); verbatimElevation: 115 m; verbatimLatitude: 44°00'12.8''N; verbatimLongitude: 16°01'08.8''E; Event: samplingProtocol: hand collected; eventDate: 4 May 2010; Record Level: institutionCode: CBSS; collectionCode: CHP416 Type status: Paratype. Occurrence: recordedBy: A. Komerički; individualCount: 1; sex: damaged male; lifeStage: adult; Location: country: Croatia; stateProvince: Knin; verbatimLocality: NP Krka, village Kistanje, Hydroelectric power plant Miljacka, cave Miljacka II; verbatimElevation: 115 m; verbatimLatitude: 44°00'01.1''N; verbatimLongitude: 16°00'58.5''E; Event: samplingProtocol: hand collected; eventDate: 21 October 2012; Record Level: institutionCode: CBSS; collectionCode: CHP552

Description

Description of holotype

Body length:approx. 30 mm (measured from anterior margin of cephalic plate to posterior margin of telson); leg 15 - 22.6 mm long, or 75% length of body.

Color: uniformly yellow-brownish to chestnut, margins of cephalic plate slightly darker than inner parts (Fig. 1).

Head: cephalic plate broader than long (4.0 x 3.6 mm, respectively), as wide as T1 (Fig. 2a); surface smooth, with several minute scattered pits, setae generally absent, except for a few emerging from the marginal ridge (above ocelli) and on the median sulcus. Cephalic median sulcus contributing to biconvex anterior margin, marginal ridge with a median thickening; posterior margin straight or slightly concave; transverse suture situated at about 1/3rd of anterior edge; posterior limbs of transverse suture visible, connecting basal antennal article with anterior part of ocellar area. Ocelli: 1+14 blackish, irregular in shape, in 3-4 rows, outermost first seriate ocellus largest, ocelli of the middle two rows medium-sized, those of inferior row smallest (Fig. 2b). Tömösváry’s organ: moderately large (as large as a medium ocellus), oval, situated on subtriangular sclerotisation below the inferiormost row of seriate ocelli (Fig. 2b). Clypeus: with a cluster of 25-30 setae situated on the apex and near the lateral margin (Fig. 3a). Antennae: right antenna composed of 71 articles, left antenna damaged after 61st article; slightly surpassing posterior margin of T11 (right) or T9 (left) when folded backwards, basal 2 articles enlarged, less pilose; posterior 30 articles visibly longer than broad, ultimate article approx. 1.3 times longer than penultimate one (Fig. 3b). Forcipules: coxosternite subpentagonal (Fig. 4a), shoulders almost absent (steep), lateral margins straight; anterior margin set off as a rim by furrow; coxosternal teeth 8+8, median diastema well-developed, V-shaped, steep and narrow, porodont arising from a pit below the dental rim, situated lateral to the lateralmost tooth; base of porodont thinner then adjacent tooth, coxosternite sparsely setose anteriorly; setae moderately large, irregularly dispersed (Fig. 4b). Forcipular trochanteroprefemur, femur and tibia and proximal part of forcipular tarsungulum with several setae. Distal part of forcipular tarsungulum about 3 times longer than proximal part (Fig. 4a).

Tergites: T1 wider than long, subtrapeziform, wider anteriorly, posterior margin straight or slightly emarginated, marginal ridge with a small median thickening; TT3 and 5 more elongated than T1, posterior margin slightly emarginated medially, posterior angles rounded; posterior angles of T4 rounded; posterior margin of T8 slightly emarginated medially, angles rounded; TT6 and 7 with posterior angles abruptly rounded (Fig. 5a); TT9, 11, 13 with well-developed posterior triangular projections (Fig. 5b); posterior margin of TT10, 12, 14 slightly emarginated, posterior-most part densely setose; intermediate tergite subpentagonal, posterior margin deeply emarginated, lateral edges bent upwards, covered with setae; middle part of posterior third of tergite densely covered with setae; laterally, on both sides of the central setose area there are two specific seta-free regions (Fig. 6a, sfa). All tergites smooth, setae present only on their lateral margins.

Legs: leg 15 longest; leg 14 approx. 25% longer than legs 1-12, leg 13 only slightly longer than legs 1-12; pretarsus of legs 1-14 with expanded fundus, larger posterior accessory claw (approx. 1/3rd of fundus) and a slightly thinner and shorter anterior accessory claw (= spine, sensu Bonato et al. 2010) (Fig. 6b); pectinal (seriate) setae missing on tarsi 1 and 2 of leg 15, present in one short row on tarsus 2 of leg 14, and in one row on tarsus 1 and two rows on tarsus 2 of legs 1-13 (Fig. 7a); pretarsus of leg 15 without accessory spines (Fig. 7b). Length of podomeres of leg 15: coxa 1.5 mm, prefemur 3.7 mm, femur 4.0 mm, tibia 5.2 mm, tarsus 1 5.0 mm, tarsus 2 3.0 mm, pretarsus 0.25 mm. Prefemur of leg 15 with a large apically rounded proximal knob (Fig. 8) protruding mediad, latter slightly bent dorsad and bearing a peculiar cluster of long setae on tip (Fig. 9a); posterior edge with well defined circular protuberance at mid-distance between spines a and p dorsally, covered with long setae (Fig. 9b), rest of prefemur covered with sparse setae. Dorsal spine p on prefemur (but also in other podomeres and other legs) with characteristic bi- and tripartite tip (Fig. 10a). Legs 1-14 without particular modifications. Coxal pores: generally round, arranged in 4-5 irregular rows, pores of inner rows largest, size decreasing outwards; pores separated from each other by a distance more than, or equal to their own diameter; number of pores on leg-pair 12: L-36/R-33, 13 L-41/R-44, 14 L-52/R-49, 15: L-39/R-34 (Fig. 10b).

Sternites: all sternites smooth, subtrapeziform, with few sparse setae, mainly at lateral margins; posterior margins straight.

Genitalia: posterior margin of male first genital sternite deeply concave, up to half of its length, posterior margin densely covered with long setae, the rest of sternite sparsely covered with shorter setae; gonopod small, hidden behind the edge of first genital sternite, with 4-5 short setae (Fig. 11).

Plectrotaxy: as in Table 1.

Description of male paratypes

All characters like in the holotype, except the following: length of leg 15: prefemur 2.5 mm; femur 3.5 mm; tibia 4 mm; tarsus 1 3.7 mm; tarsus 2 2.5 mm; pretarsus 0.3 mm; ocelli: 1+12-1+13; antennae composed of 68-70 articles; coxosternal teeth: 6+7.

Description of adult female paratype

Body length: approx. 31 mm; leg 15 approx. 20-21 mm, or 68% length of body. Color: uniformly yellow-brownish to chestnut, head and T1 darker, legs yellowish, margins of tergites darker; distal parts of tarsungulum, coxosternal teeth and pretarsi of all legs dark brown to blackish.

Head: cephalic plate broader than long (3.9 x 3.5 mm, respectively), as wide as anterior part of T1; surface smooth, with several pits scattered throughout the head and giving rise to trichoid setae. Cephalic median sulcus contributing to biconvex anterior margin, marginal ridge with a median thickening; posterior margin slightly concave; transverse suture situated at about 1/3 of anterior edge; posterior limbs of transverse suture visible, connecting basal antennal article with anterior part of ocellar area. Ocelli: 18 blackish, subequal in size, in 3-4 rows. Tömösváry’s organ: moderately large (as large as or slightly larger than a medium ocellus), oval, situated slightly above the cephalic edge below the inferiormost row of ocelli. Clypeus: with a cluster of about 25 trichoid setae situated on the apex. Antennae: approx. 22 mm long, composed of 67 articles, reaching the middle of T10 when folded backwards, basal 2 articles enlarged, less setose; posterior 30 articles visibly longer than broad, ultimate article approx. 1.3 times longer than penultimate one. Forcipules: coxosternite subpentagonal, shoulders almost absent, lateral margins straight; anterior margin set off as a rim by furrow; coxosternal teeth 7+7, median diastema well-developed, V-shaped, subparallel and narrow, porodont arising from a pit below the dental rim, situated lateral to the lateralmost tooth; base of porodont thinner then adjacent tooth, coxosternite sparsely setose anteriorly; setae moderately large, irregularly dispersed. Medial side of forcipular trochanteroprefemur, femur and tibia and proximal part of forcipular tarsungulum setose. Distal part of forcipular tarsungulum about 3 times longer than proximal part.

Tergites: T1 wider than long, subtrapeziform wider anteriorly, posterior margin slightly concave; TT3 and 5 more elongated than T1, posterior margin slightly concave medially, posterior angles rounded; T2 almost entirely covered by T1, only posteriormost part surpassing the margin of T1; posterior margin of TT4 and 6 straight, posterior angles abruptly rounded; T7 rectangular, posterior margin straight, posterior angles abruptly rounded; T8 approx. 1.4 times longer than T7, posterior margin of T8 slightly concave medially, angles abruptly rounded; TT9, 11, 13 with a well-developed posterior triangular projections; TT10 and 12 subequal in size, approx. 1.2 times longer than T8, posterior margin slightly emarginated; posterior margin of T14 slightly emarginated, surface smooth, posterior-most part covered with just a few trichoid setae (much more setose in male, see Fig. 6a); intermediate tergite subpentagonal, posterior margin deeply emarginated, surface smooth, lateral edges bent upwards, a few trichoid setae emerging from the posterior and lateral edges; areas covered with spines and setae, as well as the specific setose free areas present in male (Fig. 6a, sfa) absent.

Legs: leg 15 longest, leg 14 latter approx. 25% longer than legs 1-12, leg 13 only slightly longer than legs 1-12; pretarsus of legs 1-14 with a more expanded fundus, larger posterior accessory claw (approx. 1/3rd of fundus) and a slightly thinner and shorter anterior accessory claw (= spine, sensu Bonato et al. 2010); pectinal (seriate) setae missing on tarsi 1 and 2 of leg 15, present in one short row on tarsus 2 of leg 14, and in one row on tarsus 1 and two rows on tarsus 2 of legs 1-13; pretarsus of leg 15 without accessory spines. Leg 15 slender and elongate, without particular modifications. Bifurcated spines present irregularly on most podomeres. Coxal pores: generally round, forming 4-5 irregular rows, pores of inner rows largest, size decreasing outwards; pores separated from each other mostly by a distance more than or equal to their own diameter.

Sternites: subtrapeziform in shape, anterior part wider; lateral sides straight in all but ultimate sternite, where they are slightly convex; sternite surface smooth, shining, covered with a few sparse setae, mainly at lateral margins.

Female gonopods: densely setose, with 2+2 long and pointed spurs slightly bent outwards, and a single blunt claw; outer spur 1.4-1.5 times longer than the inner one, approx. 4 times longer than broad at base; 3-4 dorsal setae on article 1; 12 on article 2.

Plectrotaxy: as in Table 2.

Description of other female paratypes

Length: 19-22 mm; ocelli: 1+10-1+11; antennae composed of 65-68 articles; coxosternal teeth: 7+7. Tergites: TT8, 10 and 11 slightly emarginated; posterior margin of TT2, 4, 6, 7 straight. Legs: seriate setae missing on the tarsi 1 and 2 of leg 15, present in one short row only on posterior part of tarsus 2. Female gonopods: with 2+2 elongated sharply pointed spurs slightly bent outwards and a single blunt claw; 3-4 dorsal setae on article 1; 8 on article 2. Sometimes, a small, pointed spine occurs posteriorly in the middle of the first genital segment; so far, it has been detected only in two adult females [ Kaczmarek (1973) reported similar spur in Polybothrus ochraceus (Folkmanova, 1936) (= E. transsylvanicus , cf. Stoev 2002)].

Diagnosis

The species can be readily distinguished from all other congeners by the following set of molecular and morphological characters: interspecific genetic distance in COI from the closest neighbour, E. leostygis : 14.5-15.4%; antennae moderately long (approx. 70% body length), comprised of 67-71 articles; 11-15 ocelli; 6+6-8+8 coxosternal teeth; tergites 9, 11, 13 with posterior triangular projections; intermediate tergite subpentagonal, posterior margin deeply emarginated, middle part of posterior third of tergite densely covered with setae; laterally, on both sides of the central setose area, there are two specific seta-free regions; pretarsus 15 without accessory spines; leg 15 long (approx. 70-75% body length), prefemur of male leg 15 with a large, apically rounded proximal knob protruding mediad, latter slightly bent dorsad and bearing a cluster of long setae on tip; distal part of prefemur with a well-defined circular protuberance covered with setae; posterior margin of male first genital sternite deeply emarginated, nearly as deep as half of the sternite’s length.

Etymology

Cavernicolus means "living in caves or caverns", to emphasise that the species inhabits caves.

Description of the type locality

Eupolybothrus cavernicolus Komerički & Stoev sp. n. is so far known only from the caves Miljacka II and Miljacka IV (= Špilja kod mlina na Miljacki), situated near the village of Kistanje, Krka National Park, Knin District, Croatia (Fig. 12). The two caves are situated close to each other and are formed in Middle Eocene to Early Oligocene conglomerate and marbly limestone. Miljacka II is the longest cave in the Krka National Park, with a large, spacious entrance and a total length of over 2800 m (Fig. 13). Most of the cave passages are under water except for approx. 300 m of main passage. From a hydrogeological point of view, cave Miljacka II contains a periodical spring, while cave Miljacka IV has a permanent water flow. The cave Miljacka IV has two entrances, one dry and one underwater, and a length of approximately 43 m. The land entrance is walled in and with a small door while inside the cave there is a thick drywall separating it in two parts. The climatic conditions in Miljacka II as measured on 4th May 2010 and 8th October 2010 are as follows: Tair = 12.5-13.7°C (Kestrel); RH = 100%; Tsediment = 12.5-13.2°C; Twater = 12.6-13.2°C; in Miljacka IV (measured on 4 May 2010): Tair = 13.1-13.6°C (Kestrel); RH = 100%; Tsediment = 12.5°C; Twater = 12.5°C. In Miljacka II, the specimens were collected in the aphotic zone, approx. 50 m from the cave entrance, in a passage where water never occurs in a periodic flow. In Miljacka IV, they were found closer to the entrance, under stones.

Associated fauna: Gastropoda: Oxychilus cellarius (O.F. Müller, 1774), Hauffenia jadertina Kuščer, 1933, Hadziella sketi Bole, 1961; Araneae : Episinus cavernicola (Kulczynski, 1897), Nesticus eremita Simon, 1879, Tegenaria domestica (Clerck, 1757), Metellina merianae (Scopoli, 1763), Histopona sp.; Pseudoscorpiones : Chthonius tetrachelatus (Preyssler, 1790), Chthonius litoralis Hadži, 1933, Neobisium carsicum Hadži, 1933, Pselaphochernes litoralis Beier, 1956; Opiliones : Nelima troglodytes Roewer, 1910; Acari: Parasitus sp.; Isopoda : Monolistra pretneri Sket, 1964, Sphaeromides virei mediodalmatina Sket, 1964, Alpioniscus balthasari (Frankenberger, 1937), Cyphopleon kratochvili (Frankenberger, 1939); Amphipoda : Niphargus sp.; Decapoda : Troglocaris sp.; Chilopoda: Eupolybothrus tridentinus , Harpolithobius sp., Lithobius sp., Cryptops sp.; Diplopoda: Brachydesmus subterraneus Heller, 1858; Collembola : Troglopedetes pallidus Absolon, 1907, Heteromurus nitidus (Templeton, 1835), Pseudosinella heteromurina (Stach, 1929), Lepidocyrtus sp.; Diplura: Plusiocampa (Stygiocampa) dalmatica Conde, 1959, Japygidae gen. spp.; Coleoptera : Laemostenus cavicola muelleri (Schaum, 1860), Atheta spelaea (Erichson, 1839); Orthoptera : Dolichopoda araneiformis (Burmeister, 1838), Troglophilus ovuliformis Karny, 1907, Gryllomorpha dalmatina Ocskay, 1832; Psocoptera : Psyllipsocus sp.; Lepidoptera : Apopestes spectrum (Esper, 1787); Amphibia: Proteus anguinus Laurenti, 1768; Chiroptera : a colony of bats, Myotis capaccinii (Bonaparte, 1837) ( Marguš et al. 2012).