Lasiochernes pavlekae Hlebec & Harvey, 2024

Hlebec, Dora, Harms, Danilo, Kučinić, Mladen & Harvey, Mark S., 2024, Integrative taxonomy of the pseudoscorpion family Chernetidae (Pseudoscorpiones: Cheliferoidea): evidence for new range-restricted species in the Dinaric Karst, Zoological Journal of the Linnean Society (Zool. J. Linn. Soc.) 200 (3), pp. 644-669 : 659-665

Lasiochernes pavlekae Hlebec & Harvey , sp. nov.

Zoobank registration: http://zoobank.org/ NomenclaturalActs/ B1738410-6E08-448A-8BD0- F5CFF08AABD9 ( Figs 10–14 )

Type material: Holotype ♂ ( CBSSI608 ), CROATIA: Dubrovnik-Neretva County: Pelješac, Orebić, Čagjina jama cave, 42°58ʹ37.6″N, 17°09ʹ18.7″E, 107 m a.s.l. (cave entrance), 3 November 2016, F. Belak . Paratypes 1♂ ( CBSS292 A) , 4♀♀ ( CBSSI292 B, CBSSI292 C, CBSSI292 D, CBSS32 ), same data except 14 November 2018, F.Šarc ; 2♀♀ ( CBSSI293 , CBSS43 ), same data except 14 November 2018, P. Visković .

Additional material examined: 1♂ ( CNHM470 ), 42°50ʹ48.8″N, 17°38ʹ12.1″E, 415 m a.s.l. (cave entrance): Pelješac , Ston , Vukasi, Bijelo jezero, 29 March2019, B. Jalžić ; 1♂ ( CBSS312 A), 42°84ʹ72.7″N, 17°63ʹ64.7″E, 410 m a.s.l. (cave entrance): Pelješac , Ston, Crno jezero, 29 March 2019, A. Kirin; 2♂♂, 1♀ ( CBSS37 , CBSSI346 , WAMT160123 ), 42°58ʹ45.8″N, 17°17ʹ45.6″E, 275 m a.s.l. (cave entrance): Pelješac , Orebić , Oskorušno, Jama više vodospreme, 23 April 2019, V. Sudar ; 1♀ ( CBSS344 N), same data except 11 October 2022, B. Jalžić; 1♀ ( CBSS54 ), same data except 23 April 2019, L. Ružanović ; 1♂ ( CNHM657 ), 42°55ʹ08.4″N, 17°23ʹ12.8″E, 250 m a.s.l. (cave entrance): Pelješac , Trstenik , Čućin Vrh, Prnčeva špilja, 10 November 2020, B. Jalžić; 1♂, 1♀ ( CNHM780 , CNHM781 ), same data except 11 October 2022 ; 3♂♂ ( CBSS502 A, CBSS502 B, WAMT160122 ), same data except 10 November 2020 , M. Pavlek; 1♂ ( CBSSI319 ), 42°97ʹ85.4″N, 17°12ʹ90.9″E, 53 m a.s.l. (cave entrance): Pelješac , Orebić , Kučište, Šimunkovića špilja, 29 April 2019 , M. Jagić; 1♂ ( CBSS64 ), 42°52ʹ28.9″N, 17°36ʹ59.4″E, 434 m a.s.l. (cave entrance): Pelješac , Ston , Sparagovići, Vranja jama, 16 November 2018 , P. Visković; 1♂, 2♀♀ ( CNHM628 , CNHM625 , CNHM627 ), 42°40ʹ37.9″N, 18°08ʹ10.3″E, 103 m a.s.l. (cave entrance): Dubrovnik , Rožat , Vilina špilja-Izvor Omble cave, 29 June 2020 , B. Jalžić; 1♂ ( CBSSI533 ), 42°73ʹ46.2″N, 16°91ʹ06.9″E, 100 m a.s.l. (cave entrance): Lastovo , Radaš do, Rača, 10 September 2018 , M. Lukić; 1♂, 1♀ ( CNHM779 , CNHM778 ), 42°45ʹ29.2″N, 17°29ʹ16.8″E, 190 m a.s.l. (cave entrance): Mljet , Babino Polje , Blato, Dočina špilja, 23 February 2021 , B. Jalžić; 1♂ ( CBSSI423 ), 42°73ʹ17.8″N, 17°54ʹ24.1″E, 128 m a.s.l. (cave entrance): Mljet , Babino Polje, Jama pod Sv. Spasom, 19 June 2021, A. Ćukušić; 1♂, 1♀ ( CBSSI424 A, CBSSI424 B), 42°76ʹ44.4″N, 17°47ʹ10.7″E, 55 m a.s.l. (cave entrance): Mljet, Babino Polje, Blato, Mala spila, 16 October 2021, M. Pavlek; 1♀ ( CNHM513 ), same data except 28 September 2019, B. Jalžić; 1♂ ( CBSSI458 ), 42°76ʹ13.3″N, 17°43ʹ08.0″E, 265 m a.s.l. (cave entrance): Mljet , Babino Polje , Goveđari, Male ponte, 16 June 2021 , A. Ćukušić; 1♀ ( CBSSI595 A), same data except 14 February 2021 , T. Rožman ; 1♀ ( CBSS436 I), 42°45ʹ30.2″N, 17°26ʹ10.0″E, 201 m a.s.l. (cave entrance): Mljet , Babino Polje , Goveđari , Špilja kod Nerezinog dola, 16 October 2021 , N. Kuharić; 1♂ ( CBSSI453 ), 42°76ʹ71.1″N, 17°45ʹ36.1″E, 102 m a.s.l. (cave entrance): Mljet, Babino Polje, Blato, Špilja na Strmici, 17 June 2021, H. Cvitanović; 1♀ ( CBSSI99 A) same data except 20 October 2015, M. Pavlek; 1♀ ( CBSSI430 ), 42°76ʹ46.8″N, 17°47ʹ20.4″E, 79 m a.s.l. (cave entrance): Mljet , Babino Polje , Blato, Velika špilja, 16 June 2021 , M. Pavlek; 1♀ ( CBSS433 ) same data except 06 October 2021 , I. Čupić; 1♀ ( CBSSI534 A), 42°75ʹ25.5″N, 17°55ʹ11.6″E, 210 m a.s.l. (cave entrance): Mljet , Babino Polje, Movrica, 15 June 2021 , M. Pavlek; 1♂, 1♀ ( CBSSI91 , CBSSI188 ), 43°77ʹ42.6″N, 15°34ʹ97.1″E, 15 m a.s.l. (cave entrance): Šibenik-Knin County, Kornati , Gustac Islet , Vjetruša, 18 April 2018 , T. Čuković and A. Ćukušić .

Diagnosis: Lasiochernes pavlekae sp. nov. differs from other Lasiochernes species by the presence of dense setation patterns on all pedipalpal segments (♂, ♀), slightly more setose in males ( Figs 10A, B , 11A, B ), and a normal posterolateral margin at coxa IV ( Fig. 10C, D ). Pedipalpal setation is absent in Lasiochernes jalzici sp. nov. ( Figs. 7 , 8 ) and pronounced basally on the prolateral face of the male femur in Lasiochernes marinae sp. nov. ( Figs 4 , 5 ).

Paratype: CBSS32 (DNA voucher CROBD583 ; GenBank ON 841917) served as the reference sequence, because the amplification of the COI gene fragment from the holotype was unsuccessful.

Other sequenced material: CBSSI 312A (DNA voucher CBSSI 312A; GenBank OQ 330000); CBSSI 319 (DNA voucher CBSSI 319; GenBank OQ 330001); CBSSI 595A (DNA voucher CBSSI 595A; GenBank OQ 330002); CBSSI 533 (DNA voucher CBSSI 533; GenBank OQ 330003); CBSSI 534A (DNA voucher CBSSI 534A; GenBank OQ 330004); CBSSI 91 (DNA voucher CBSSI 91; GenBank OQ 330005); CNHM 470 (DNA voucher CROBD 269; GenBank ON842238); CNHM 657 (DNA voucher CROBD 767; GenBank ON842072); CBSS 37 (DNA voucher CROBD 588; GenBank ON842232); CBSS 436I (DNA voucher CROBD 1194; GenBank ON841949); WAMT 160123 (DNA voucher CROBD 599; GenBank CROBD 599; GenBank ON842212); CBSS 54 (DNA voucher CROBD 607; GenBank ON842045); CBSS 64 (DNA voucher CROBD 617; GenBank ON842194); CNHM 627 (DNA voucher CROBD 670; GenBank ON842271); CNHM 628 (DNA voucher CROBD 671; GenBank ON842142); and CNHM 779 (DNA voucher CROBD 1015; GenBank ON842083). See the Supporting Information, Appendix S6 for a list of diagnostic nucleotides in the barcode fragment of the COI gene. Uncorrected pairwise genetic distances (p -distances) between Lasiochernes pavlekae sp. nov. and other Lasiochernes species are listed in the Supporting Information, Appendix S9. Positions refer to the alignment available in the Supporting Information, Appendix S10.

Description (adults)

Colour: Pedipalps, carapace and coxae deep red-brown; tergites and sternites brown, darker in female; legs yellow-brown ( Fig.10 ).

Pedipalp ( Figs 11 , 13A ): All pedipalpal surfaces granulate; all segments setose, all setae long ( Figs 10A, B , 12A, B ); patella with several dorsal lyrifissures ( Fig. 11A, B ); all segments robust, more robust in male; trochanter 1.71 (♂), 1.79 (♀), femur 2.18 (♂), 2.73 (♀), patella 2.13 (♂), 2.12 (♀), chela (with pedicel) 2.82 (♂), 3.00 (♀), chela (without pedicel) 2.66 (♂), 2.76 (♀), hand (without pedicel) 1.22 (♂), 1.39 (♀) × longer than broad, movable finger (without pedicel) 0.90 (♂), 0.97 (♀) × longer than hand. Fixed chelal finger with eight trichobothria, movable chelal finger with four trichobothria ( Fig. 11A, B ): eb and esb situated basally, est situated midway between esb and et, ib and ist situated subbasally, isb situated slightly posterior to est and closer to ist than to it, and it situated subdistally; t situated subdistally, st situated closer to t than to sb, and sb situated much closer to b than to st ( Fig. 11C, D ). Venom apparatus only present in movable chelal finger, venom duct long, terminating in nodus ramosus basal to t. Pseudotactile setae absent. Chelal fingers not gaping; teeth slightly retrorse, juxtadentate; fixed finger with 44 (♂), 42 (♀) teeth, plus 4 (♂), 7 (♀) retrolateral accessory teeth and 5 (♂, ♀) distal prolateral accessory teeth; movable finger with 45 (♂), 43 (♀) teeth, plus 9 (♂), 7 (♀) retrolateral accessory teeth and 2 (♂), 6 (♀) distal prolateral accessory teeth; sense-spots arranged: fixed chelal finger with 12 (♂), 15 (♀) retrolateral from which 13 arranged at the base of the finger, 12 (♂), 9 (♀) prolateral, and movable chelal finger with 2 (♂), 4 (♀) retrolateral and 0 (♂, ♀) prolateral.

Chelicera: With five setae on hand and one subdistal seta (gs) on movable finger; setae es, bs and sbs strongly dentate, ls and is long and acuminate ( Figs 12A, B , 13C ); galea well-developed, long and thin with three distal and three medial rami (♂, ♀) ( Fig. 12D, E ); with two dorsal lyrifissures and one ventral lyrifissure; rallum composed of three blades, all three dentate on anterior face ( Fig. 12C ); serrula exterior with 19 (♂, ♀) blades; lamina exterior present.

Carapace: Granulate and sub-rectangular; 1.13 (♂) and 1.07 (♀) × longer than broad; eyes and eye-spot absent; with two transverse furrows, posterior furrow situated slightly closer to posterior margin than to anterior furrow, posterior furrow with small posteromedian furrow ( Fig. 10A, B ); with 106 (♂), 102 (♀) setae including 7 (♂, ♀) setae near anterior margin and 13 (♂), 11 (♀) setae near posterior margin;

Coxal region: Maxillae rugose, coxae smooth; manducatory process pointed, with three apical acuminate setae, with one small sub-oral seta on medial edge, and 46 (♂), 49 (♀) additional setae. Chaetotaxy of coxae I–IV: ♂: 28: 38: 41: 65; ♀: 34: 29: 50: 69.

Legs: femur + patella of leg IV 4.87 (♂), 5.27 (♀) × longer than deep; tarsus IV with long tactile seta located medially, TS ratio = 0.49 (♂), 0.51 (♀); arolium shorter than claws, claws simple ( Fig. 12G ).

Abdomen: Tergites I‒II partially divided, other tergites and sternites IV‒XI with broad medial suture line. Tergal chaetotaxy: ♂, 16: 18: 19: 25: 23: 22: 22: 21: 23: 15: 2: 2; ♀, 14: 16: 19: 22: 24: 23: 22: 21: 21: 16: 10: 2; setae lightly dentate. Sternal chaetotaxy: ♂, 46: (4) 28 [4 + 3] (4): (4) 19 (4): 24: 29: 29: 28: 28: 27: 14: 2; ♀, 33: (3) 13 (3): (4) 9 (4): 20: 26: 26: 23: 23: 21: 12: 2. Tergites and sternites without tactile setae. Pleural membrane wrinkled plicate, without setae.

Genitalia: Male with typical chernetid morphology; female with single T-shaped spermatheca, length of each receptaculum slightly shorter than median stalk ( Fig. 12F, H, I ).

Dimensions (mm): Male holotype (CBSSI608), followed by 12 other males in parentheses: body length 3.600 (2.940 –4.015). Pedipalps: trochanter 0.575 / 0.335 (0.490–0.660/0.290– 0.390), femur 0.930 / 0.425 (0.775–1.120/0.335–0.490), patella 0.920 / 0.430 (0.770–1.050/0.380–0.510), chela (with pedicel) 1.520 / 0.540 (1.390–1.680/0.460–0.615), chela (without pedicel) 1.440 (1.290 –1.540), hand (without pedicel) length 0.660 (0.640 –0.800), movable finger length 0.770 (0.660 – 0.815). Carapace 1.090 / 0.960 (0.990–1.200/0.820–1.160). Leg IV (holotype): femur + patella 0.950 / 0.195, tibia 0.770 / 0.160, tarsus 0.495 / 0.105, TS = 0.490.

Dimensions (mm): Female paratype (CBSSI293), followed by 16 other females in parentheses: body length 4.950 (3.500 –5.100). Pedipalps: trochanter 0.625 / 0.350 (0.490–0.690/0.275–0.395), femur 1.010 / 0.370 (0.775–1.150/0.300–0.415), patella 0.890 / 0.420 (0.750–1.040/0.315–0.460), chela (with pedicel) 1.650 / 0.550 (1.330–1.780/0.430–0.600), chela (without pedicel) 1.520 (1.290 – 1.650), hand (without pedicel) length 0.765 (0.605 –0.790), movable finger length 0.790 (0.620 –0.860). Carapace: 1.190 / 1.110 (0.905–1.250/0.920–1.340). Leg IV (paratype): femur + patella 1.080 / 0.205, tibia 0.845 / 0.135, tarsus 0.510 / 0.105, TS = 0.510.

Remarks: Lasiochernes pavlekae has been recorded in seven caves on the Pelješac Peninsula, Vilina špilja-Izvor Omble cave, as well as 10 additional caves on islands (Mljet Island, Lastovo Island, and Gustac Islet in the Kornati archipelago). All of these locations are within a aerial distance of less than 50 km from the coast. Most caves range from 7 to 114 m in total length, with the exception of Špilja Crno jezero cave, which is 239 m long, and Vilina špilja-Izvor Omble cave, which streches 3050 m in length. The entrances of these caves are situated at elevations between 15 and 434 m above sea level. The majority of them harbour colonies of bats, including maternity colonies of the Mediterranean horseshoe bat ( Rhinolophus euryale Blasius, 1853 ), the greater horseshoe bat [ R. ferrumequinum (Schreber, 1774) ], Geoffroy’s bat [ Myotis emarginatus (E. Geoffroy, 1806) ], and the lesser mouse-eared bat ( M. blythii Tomes, 1857 ) ( Pavlinić et al. 2010).

Twelve haplotypes were found in the COI gene (p -distances = 0.18–1.09%) and two in 28S (p -distances = 0.36%), but other genes had one haplotype demonstrating that there is low genetic variability within this species despite some variation in the fast-evolving mitochondrial genome.

Etymology: This species is named for the Dr. Martina Pavlek, a Croatian biospeleologist and taxonomic specialist in subterranean spiders.

Figures

Figure 4. Lasiochernes marinae sp. nov.. A, holotype male, habitus (CNHM779), dorsal. B, paratype female, habitus (CNHM491), dorsal. C, holotype male, cephalothorax, ventral. D, paratype female, cephalothorax, ventral. Scale lines = 1 mm (A, B), 0.5 mm (C, D).

Figure 5. Lasiochernes marinae sp. nov.. A, holotype male (CNHM779), right pedipalp, dorsal. B, paratype female (CNHM491), right pedipalp, dorsal. C, holotype male, left chela, retrolateral. D, paratype female, left chela, retrolateral.Scale lines = 0.25 mm.

Figure 7. Lasiochernes jalzici sp. nov.. A, holotype male, habitus (CNHM650), dorsal.B, paratype female, habitus (CNHM778), dorsal. C, holotype male, cephalothorax, ventral. D, paratype female, cephalothorax, ventral. Scale lines = 1 mm (A, B), 0.5 mm (C, D).

Figure 8. Lasiochernes jalzici sp. nov.. A, holotype male (CNHM650), right pedipalp, dorsal.B, paratype female (CNHM778), right pedipalp, dorsal. C, holotype male, left chela, retrolateral. D, paratype female, left chela, retrolateral. Scale lines = 0.25 mm.

Figure 10. Lasiochernes pavlekae sp. nov.. A, holotype male, habitus (CBSSI608), dorsal. B, paratype female, habitus (CBSSI293), dorsal. C, holotype male, cephalothorax, ventral. D, paratype female, cephalothorax, ventral. Scale lines = 1 mm (A, B), 0.5 mm (C, D).

Figure 11. Lasiochernes pavlekae sp.nov.. A, holotype male (CBSSI608), right pedipalp, dorsal.B, paratype female (CBSSI293), right pedipalp, dorsal. C, holotype male, left chela, retrolateral. D, paratype female, left chela, retrolateral. Scale lines = 0.25 mm.

Figure 12. Lasiochernes pavlekae sp. nov.A, holotype male (CBSSI608), left chelicera, dorsal. B, paratype female (CBSSI293), left chelicera, dorsal. C, paratype female, left rallum, lateral. D, holotype male, left galea, dorsal.E, paratype female, left galea, dorsal. F, holotype male, genital opercula (sternites II and III). G, holotype male, left leg IV, retrolateral. H, paratype female, genital opercula (sternites II and III). I, paratype female, spermathecae. Scale lines = 0.25 mm (G), 0.1 mm (A, B, F, H, I), 0.05 mm (C, D, E).

Figure 13. Lasiochernes pavlekae sp.nov., male, scanning electron micrographs.A, left pedipalp, dorsal.B, right chela, dorsal.C, chelicera, dorsal. Scale lines = 1 mm (A), 0.5 mm (B), 150 μm (C).

Figure 14. Genetic and morphological variability of Lasiochernes pavlekae sp. nov.. A, MJ haplotype network for L. pavlekae sp. nov. based on the COI gene. Haplotype circles are colour-coded according to localities and circle size is proportional to overall haplotype frequency. Numbers of mutational steps are given as hatch marks. B, sampling localities (colour coding matches insert in Fig.14A). C, visualization of PCA results performed on morphometric data deriving from the two populations of L. pavlekae sp.nov.. Polygonal shapes indicate the range of variability found in a given population. The points and polygons are coloured according to localities. D, visualization of PCA results performed on morphometric data deriving from male and female specimens of L.pavlekae sp. nov..