ILLYRIONETHES VERHOEFF, 1927

Bedek, Jana, Taiti, Stefano, Bilandžija, Helena, Ristori, Emma & Baratti, Mariella, 2019, Molecular and taxonomic analyses in troglobiotic Alpioniscus (Illyrionethes) species from the Dinaric Karst (Isopoda: Trichoniscidae), Zoological Journal of the Linnean Society 187, pp. 539-584 : 546-567

SUBGENUS ILLYRIONETHES VERHOEFF, 1927

Type species: Illyrionethes strasseri Verhoeff, 1927 by monotypy.

Diagnosis: Unpigmented body, eyes absent. Pleon narrower than pereon. Antennula of three articles, distal article bearing aesthetascs on subapical and/ or apical margin. Antennal flagellum with five to 30 segments. Mandibles with one/two penicils in the right and three penicils in the left; molar process with none/ one penicil in the right and none in the left. Outer branch of maxillula with ten/11 teeth, apically entire and one/two slender stalks; inner branch with three penicils. Maxilla with setose apex. Pereopods with setose dactylar seta. Uropod endopod shorter than exopod. Male pereopod 7 ischium with straight sternal margin. Genital papilla with rounded apical part. Male pleopod 1 endopod elongated, bearing apical seta. Male pleopod 2 endopod longer than exopod; of two articles, with proximal article distinctly longer than distal article, and distal article bearing terminal seta.

Remarks: The above diagnosis is based on all species included in Illyrionethes , with the exception of Alpioniscus escolai Cruz & Dalens, 1989 from Catalonia, which needs a taxonomic revision owing to some morphological differences. The only character that distinguishes subgenus Illyrionethes from subgenus Alpioniscus is the male pleopod 2 endopod with proximal article distinctly longer than distal one.

ALPIONISCUS (ILLYRIONETHES) STRASSERI ( VERHOEFF, 1927)

( FIGS 6–12; TABLE 1)

Illyrionethes strasseri Verhoeff, 1927: 270–274 ; figs 1–7. – 1929 a: 28. – 1929b: 51. – 1931b: 20–21. – 1931c: 234. – 1932: 24. – Frankenberger, 1937: 173–175. – 1938: 34. – 1939: 98–99. – Brian, 1938: 176. – Strouhal, 1938: 272–273; figs 3–5. – 1939 a: 116, 118–119 [partim: nec NW.- Kroatien; Lokve Špilja, Kroatien]. – 1939b: 6, 20. – 1940a: 94. – 1940b: 14. – Wolf, 1938: 78–79. – Frankenberger & Strouhal, 1940: 448–449 [partim: nec NW.- Kroatien]. – Creuwels, 2018. – Natural History Museum, 2018a. – 2018b. – 2018c – Natural History Museum, 2018d.

Alpioniscus strasseri – Kesselýak, 1930: 247; figs 2a, 3b–e. – Vandel, 1947: 264, 267, 271. – Potočnik, 1989: 64. – Schmalfuss, 2003: 14 [partim: nec island Cres]. – Lukić et al., 2009: 16–17. – Lukić et al., 2011: 19–22. – Bedek et al., 2011: 275–278; fig. 16 [partim: nec Čampari; Ovčarica; Piskovica; Rabakova Špilja; Vrtare male; Lokvarka; Medvjeđa Špilja]. – Bedek et al., 2017: 213. – Ozimec et al., 2011: 166 [partim: Jama kod Burići]. – Polak et al., 2012: 10–16 [partim: nec Kubik, Piskovica].

Illyrionethes enriquesi Arcangeli, 1932: 1042–1046 ; pls. 3–9; figs 7–26. – Brian, 1938: 175–176. – Wolf, 1938: 765.

Alpioniscus (Illyrionethes) strasseri – Buturović, 1957: 6–49; fig. 24. – Schmölzer, 1965: 59 [partim: NO-Italien]. – Karaman, 1966: 376 [partim: nec Lokve Špilja; o. Cres (pećina kod Petrićevi)]. – Paoletti, 1978: 4 [partim: nec Isola di Cherso]. – Potočnik, 1979: 66. – 1993: 85–86, 163, 178. – Potočnik & Novak, 1980: 4. – Argano, Baldari & Manicastri, 1982: 132 [partim: nec Isola di Cherso]. – Argano et al., 1995: 14. – Tabacaru, 1996: 15, 40. – Horvatović, 2014: 125, 172–174; figs 7.28.a, 7.33.

Alpioniscus cf. strasseri – Polak et al., 2012: 10–16 [partim: RačiŠka pećina].

? Illyrionethes strasseri – Verhoeff, 1938: 134. – 1940: 111.

? Alpioniscus (Illyrionethes) strasseri – Karaman, 1966: 376 [partim: o. Cres (pećina kod Petrićevi)]. – Paoletti, 1978: 4 [partim: Isola di Cherso]. – Argano et al., 1982 [partim: Isola di Cherso].

? Alpioniscus strasseri – Schmalfuss, 2003: 14 [partim: including island Cres]. – Bedek et al., 2011: 275 [partim: Čampari; Ovčarica; Piskovica; Rabakova Špilja]. – Ozimec et al., 2011: 166 [partim: Ročka Špilja]. – Polak et al., 2012: 11, 16 [partim: Kubik, Piskovica].

? Alpioniscus cf. strasseri – Polak et al., 2012: 10–11 [partim: Polina peč, Jama pod Krogom, Rabakova Špilja].

nec Alpioniscus strasseri – Casale et al., 2004: 310.

nec Illyrionethes strasseri – Royal Belgian Institute of Natural Sciences, 2017.

Material examined (collection details in Supporting Information, Table S1)

Northern Istria population : ITALY. Venezia Giulia, Trieste: Basovizza, Caverna presso Basovizza (cave RCCFVG 689) ( CBSSC IT2386; IT2473; IT3870); Longera, Grotta nel Bosco dei Pini (cave RCCFVG 16) ( CBSSC IT2396; MZUF 1); San Dorlingo della Valle, Val Rosandra, Grotta del Guano (cave RCCFVG 527) ( MZUF 2). SLOVENIA. SeŽana, Štorje, Belinca jama (cave ECC 950) ( CBSSC IT995, IT1648, IT1649, IT1650); Istria, Račice, Starod, RačiŠka pečina (cave ECC 942) ( CBSSC IT2464); Divača, Škocjan, Škocjanske jame (cave ECC 735) ( CBSSC IT2813). CROATIA. Istria, Ćićarija Mts.: Lupoglav, Brgudac, BoruŠnjak, BoruŠnjak 1 (cave CCC HR01280) ( CBSSC IT1735); Lupoglav, Brgudac, BoruŠnjak, BoruŠnjak 3 (cave CCC HR01291) ( CBSSC IT1755); Lupoglav, Brest pod Učkom, Vojni poligon iznad Bresta, Crekvica (cave CCC HR01531) ( CBSSC IT2138, IT2139); Lupoglav, Brest pod Učkom, Vojni poligon iznad Bresta, Crekvica 1 (cave CCC HR01527) ( CBSSC IT682, IT683, IT684, IT686, IT687); Lupoglav, Semići, Podbrus, Jama pod kriŽ (cave CCC HR01761) ( CBSSC IT1761, IT2105, IT2107, IT2108); Opatija, Veprinac, Mahenac, Jama SD 2 (cave CCC HR00925) ( CBSSC IT1762); Lupoglav, Brgudac, Bončića kuća, Pećina kod planinarske kuće Pavlovac (cave) ( CBSSC IT1732, IT1733); Opatija, Veprinac, Puhari, Pećina kod sela Puhari (cave CCC HR00437) ( CBSSC IT1713, IT1717, IT1719); Opatija, Veprinac, BrloŽnik, Pećina pod BrloŽnikom (cave CCC HR00429) ( CBSSC IT1759); Lupoglav, Brest pod Učkom, Vela peć (cave CCC HR00614) ( CBSSC IT698, IT2127). Istria, Učka Mt.: Brest, Vojni poligon iznad Bresta, Barinova jama (cave CCC HR01262) ( CBSSC IT1131); Lovran, Potoki, MoŠćenička draga, DruŠka peć (cave) ( CBSSC IT1744); Boljun, Pikulići, Vela draga, Jama iznad Vranjske drage (cave CCC HR01296) ( CBSSC IT2117, IT2119); Veprinac, Menderi, Jama kod potoka Banine 3 (cave CCC HR00430) ( CBSSC IT721, IT1132); Mala Učka, Jama mrtvih golubova (cave CCC HR00927) ( CBSSC IT706, IT708, CNHM 3/1680); Poklon, Dobreč, Jama na BalaguŠtini (cave CCC HR01534) ( CBSSC IT1140); Vela Učka, Molinarska draga, Mala peć (cave) ( CBSSC IT727); Brest, Vojni poligon iznad Bresta, OŠtirova jama (cave CCC HR01261) ( CBSSC IT1125); Mala Učka, Podmaj, Petrc, Petrčeva jama (cave CCC HR00392) ( CBSSC IT1135, IT1136); Lovran, Lovranska Draga, Veliki Knezgrad, Proljetna jama (cave CCC HR00601) ( CBSSC IT1747, IT1749); Boljun, Vranja, Vela draga, Pupićina pećina (cave CCC HR01318) ( CBSSC IT2130, IT2131); Lovran, Medveja, Rudnik u Medveji (mine) ( CBSSC IT1721, IT1722, IT1723); Mala Učka, Podmaj, Rupa II (cave) ( CBSSC IT1142, IT1144); Poklon, Topol, Topolska peć (cave) ( CBSSC IT718, IT1138). Croatian Littoral, Kostrena: Bakar, RoŽmanići, Humčine, Bunetova jama (cave CCC HR00443) ( CBSSC IT783, IT784, IT785, IT1851, IT1852, IT1853); Perilo, Mali Štumbur (cave) ( CBSSC IT1864, IT1865); Bakar, Paveki, Špilja u uvali Predpeć (cave CCC HR00442) ( CBSSC IT779, IT781, IT782, IT1122, IT1123, IT1854, IT1855, IT1856, IT1857, CNHM 3/1679); Bakar, Paveki, Špilja u uvali SveŽanj (cave CCC HR01336) ( CBSSC IT1120, IT1121, IT1860, IT1863, IT1862); RoŽmanići, Humčine, Tončakova jama (cave) ( CBSSC IT1867); Urinj, Urinjska Špilja (cave CCC HR01110) ( CBSSC IT1841, IT1844, IT1846, IT1850, IT1849, IT1848).

Southern Istria population : CROATIA. Istria: Poreč, Nova Vas, Baredine (cave CCC HR00312) ( CBSSC IT1044, IT1046); Labin, Vozilići, Čepić tunel (Tunnel) ( CBSSC IT3551); Žminj, FeŠtini, Festinsko kraljevstvo (cave) ( CBSSC IT1045); Pula, Muntić, Ikinica (cave) ( CNHM 10/1413, CBSSC IT3529); Poreč, Žbandaj, Ivanka (cave) ( CBSSC IT3548); Kanfanar, Burići, Jama kod Burići (cave CCC HR00754) ( CBSSC IT3555, IT2229, IT2230, IT2440, IT2454, IT2455); Novigrad, Punta Karigador, Jama kod komune (cave) ( CBSSC IT3547); Labin, SnaŠići, Jama na SnaŠići (cave) ( CBSSC IT3549); Labin, RaŠa, Karlota (mine) ( CBSSC IT3553); Pula, Marčana, Ljubićeva pećina (cave) ( CBSSC IT3530); Poreč, Tar, Štancija Špin, Markova jama kod Tara (cave CCC HR00981) ( CBSSC IT3540, IT3541, IT3542, IT3543, IT2231, IT2232, IT2233, IT2446, IT2447, IT2450); Umag, Brtonigla, Mramornica (cave) ( SMNS 5326, CBSSC IT3534, IT3531); Poreč, Tar, Vrh, Pincinova jama (cave CCC HR00313) ( CBSSC IT3544, IT2434, IT2441); Svetvinčenat, Šikuti, PliŠkovićeva jama (cave) ( CBSSC IT3537, IT3538); Poreč, Nova Vas, PodŠeŠ (cave) ( CBSSC IT3546); Pula, Muntić, Staja (cave) ( CBSSC IT3528); Pula, Premantura, Špilja na Gradini kod Premanture (cave) ( CBSSC IT3512, IT3516, IT3517, IT3518, IT3520, IT3692, IT3775, IT3776, IT3778, IT3780, IT3782, IT3784, IT3786, IT3788, IT3790, IT3791, IT3793, IT3795, IT3798, IT3799, IT3801).

Redescription: Maximum length: ♂, 5.8 mm; ♀, 8.0 mm. Pereon ( Fig. 6A) with almost parallel sides. Back smooth, with some triangular scale-setae ( Fig. 6C). Some gland pores on lateral margins of pleonites 4 and 5, telson and uropodal protopods ( Fig. 6F). Cephalon ( Fig. 6D, E) with suprantennal line bent downwards, rounded middle part; antennal lobes quadrangular. Posterior margin of pereonite 1 convex, of pereonites 2 and 3 straight, and of pereonites 4–7 progressively more concave ( Fig. 6A, B). Pleonites 3–5 with small posterior points visible in dorsal view ( Fig. 6A, F). Distal part of telson with concave sides and broadly rounded apex ( Fig. 6F). Antennula ( Fig. 6G) distal article flattened and bearing six to eight aesthetascs on apical margin. Antenna ( Fig. 6H) smooth; flagellum of eight to 11 articles, with one row of aesthetascs on two to five different articles, always present on second and third. Mandibles ( Fig. 7A, B) with one penicil in right; molar process with one penicil in right. Maxillula ( Fig. 7C) outer branch with 5 + 6 teeth and two slender stalks; inner branch with outer and middle penicils subequal, inner distinctly longer. Maxilla ( Fig. 7D) apex bilobate, lobes subequal in width. Maxilliped ( Fig. 7E) endite narrow, with large segmented apical penicil; palp distally with three rounded lobes, basal article with two compound small setae; basis with rounded outer lobe protruding posteriorly and covered with long setae on margin. Pereopods ( Fig. 8A) similar in shape, with dactylar seta long and bifid. Uropod ( Fig. 6F) with protopod slightly grooved on outer margin; endopod inserted proximally.

Male: Pereopod 1 ( Fig. 8A) carpus bearing six to seven setae. Pereopod 1–4 with carpus and merus bearing numerous short scales on rostral surface. Pereopod 1–6 ( Fig. 9) merus with sternal margin from straight (pereopod 1) progressively up to slightly concave (pereopod 6) and with sternal margin bearing no lobe (pereopod 1) progressively up to small lobe (pereopod 6). Pereopod 7 ( Fig. 8B, C) merus with slightly concave sternal margin and large hook-shaped lobe in proximal part directed ventrally, bearing one seta; carpus with straight tergal margin and large rounded hump proximally, ratio of carpus length to top hump position 3.73 ± 0.53 (N = 10), and ratio of carpus length to ending hump position 2.09 ± 0.22 (N = 10). Pleopod 1 ( Fig. 10A) exopod with posterior apex narrowly rounded, ratio of length to posterior apex width 4.76 ± 0.38 (N = 10), deeply concave outer margin, ratio of length to concavity turning point position 1.73 ± 0.07 (N = 10), slightly concave inner margin; endopod narrow with almost parallel sides, apical seta plumose. Pleopod 2 ( Fig. 10C) exopod triangular with concave outer margin; endopod distinctly longer than exopod, distal article narrower than proximal; terminal seta strong and bifid. Pleopod 3–5 exopods as in Fig. 10D–F.

Type locality: In the original description, Verhoeff (1927) clearly stated that the described specimens were collected in the cave ‘ Höhle am Mt. Spaccato in Istrien, eine Stunde von Triest entfernt’ (presently known as Grotta ad W di Basovizza , Trieste , Italy; Friuli Venezia Giulia, RCCFVG 68) and that the specimens from the cave ‘ Höhle Belinca bei Storje’ (presently known as Belinca jama, Štorje, SeŽana, Slovenia; ECC. 950) were not included in the description. Therefore, the type locality encompasses only the cave Grotta ad W di Basovizza and not the cave Belinca jama, as previously reported by Bedek et al. (2011) .

Remarks: The Grotta ad W di Basovizza was not available for collecting topotypic specimens owing to the Golf Club Trieste facilities. Therefore, in the species redescription we used specimens collected in two closest possible caves: the 2.5 km distant Caverna presso Basovizza, Basovizza, Trieste, Italy (RCCFVG 689) and the 1 km distant Grotta nel Bosco dei Pini, Basovizza, Trieste , Italy (RCCFVG 16).

The populations of A. strasseri from southern Istria show some small differences ( Figs 8D, 10B, 11, 12; Table 1). Arcangeli (1932) described I. enriquesi from the Špilja na Gradini kod Premanture cave near Pula (caverna presso Promontore, dintorni di Pola) in Istria, which Strouhal (1939a) considered to be a junior synonym of A. strasseri . Taxonomic analyses confirm the differences in the males, i.e. the pereopod 7 merus with a small hump-shaped lobe on sternal margin in the proximal part; ratio of carpus length to carpus top hump position 4.42 ± 0.97 (N = 7); and ratio of pleopod 1 exopod length to concavity turning point position 1.90 ± 0.09 (N = 7). The population is geographically separated from A. strasseri s.s. and shows some molecular differences. More sampling should be conducted in regions between their distribution areas, followed by taxonomical and molecular analyses in order to validate the taxonomic status of the population.

Alpioniscus strasseri differs from the other species of the strasseri- lineage in the male characters ( Fig. 12; Table 1). It differs from A. christiani , A. balthasari , A. hirci sp. nov. and A. iapodicus in the male pereopod 7 with a larger merus hook-shaped lobe. It is distinguishable in the ratio of the pereopod 7 carpus length to carpus top hump position from A. hirci sp. nov. (1.66 ± 0.09) and A. velebiticus sp. nov. (2.08 ± 0.15); in the ratio of the male pereopod 7 carpus length to ending tergal hump position from A. velebiticus sp. nov. (1.16 ± 0.06), A. iapodicus s.s. (1.26 ± 0.03) and A. hirci sp. nov. (1.27 ± 0.03); in the position of the concavity turning point of the outer margin of the male pleopod 1 exopod from A. velebiticus sp. nov. (2.21 ± 0.14); in the ratio of the male pleopod 1 exopod length to posterior apex width from A. hirci sp. nov. (3.57 ± 0.19).

Alpioniscus strasseri is considered to be an amphibious species, because it was also observed in water ( Verhoeff, 1927). The citations with ‘?’ are doubtful, and the material from those localities needs to be re-examined with both molecular and morphological analyses.

Distribution: Italian karst of Trieste, the southern part of the Slovenian Littoral, Croatian Istria and Kostrena surroundings ( Figs 4, 11).

ALPIONISCUS (ILLYRIONETHES) CHRISTIANI POTOČNIK, 1983

( FIGS 12–14 TABLE 1)

Alpioniscus (Illyrionethes) christiani Potočnik, 1983: 389–395 ; figs 1–7. – 1993: 84, 178. – Tabacaru, 1996: 34–40. – Horvatović, 2014: 126, 173, 174, 178 [partim: island Krk]; figs 7.28.c, 7.33.

Alpioniscus christiani – Potočnik, 1989: 64. – Gottstein et al., 2001: 2257. – Gottstein Matočec et al., 2002: 48. – Schmalfuss, 2003: 13. – Tvrtković et al., 2004: 76. – Bedek et al., 2006: 15, 92. – Bedek et al., 2011: 241, 268; fig. 11. – JalŽić et al., 2010: 20. – JalŽić et al., 2013: 13, 30, 43–45; fig. on p. 44.

Alpioniscus strasseri – Bedek et al., 2011: 277 [partim: Vrtare male].

? Alpioniscus christiani – Horvatović, 2014: [partim:

BiH: Hercegovina, Mosor, Bubanj ].

Material examined (collection details in Supporting Information, Table S1)

CROATIA. Croatian Littoral: Krk Is., OmiŠalj, ČiŽići, Rudine, Biserujka (cave) ( CBSSC IT2629, IT1657,

IT1652, IT1653, IT2092, IT2093, IT2409, IT2410, IT3654, IT2478); Crikvenica, Dramalj, Vrtare male (cave) ( MZUF 9880, CBSSC IT325, IT3571, IT3573, IT3574, IT3577, IT3579, IT3580, IT3581).

Redescription: Maximum length: ♂, 5.3 mm; ♀, 8.0 mm. Pereon with almost parallel sides ( Fig. 13A). Back smooth, with some triangular scale-setae. Some gland pores on lateral margins of pleonites 4 and 5 and uropodal protopods ( Fig. 13C). Cephalon ( Fig. 13B) with suprantennal line bent downwards, rounded middle part; antennal lobes broadly rounded. Posterior margin of pereonite 1 convex, of pereonites 2 and 3 straight, and of pereonites 4–7 progressively more concave. Pleonites 3–5 with small posterior points visible in dorsal view ( Fig. 13C). Distal part of telson with concave sides and broadly rounded apex ( Fig. 13C). Antennula ( Fig. 13D) distal article flattened and bearing four to eight aesthetascs on apical margin. Antenna ( Fig. 13E) smooth; flagellum of seven to nine articles, with one row of aesthetascs on two to four articles, always present on second and third. Mouthparts as in A. strasseri . Pereopods ( Fig. 14A) similar in shape, with dactylar seta long and bifid. Uropod ( Fig. 13C) with protopod slightly grooved on outer margin; endopod inserted proximally.

Male: Pereopod 1 ( Fig. 14A) carpus bearing five to six setae. Pereopod 1–2 carpus and merus bearing numerous short scales on rostral surface. Pereopod 1–6 merus with sternal margin from straight (pereopod 1) progressively up to slightly concave (pereopod 6) and with a small lobe in proximal part increasing in size from pereopod 2 to pereopod 6. Pereopod 7 ( Fig. 14B, C) merus with slightly concave sternal margin and small hump-shaped lobe with three to five teeth in proximal part bearing one seta; carpus with straight sternal margin and large rounded dorsal hump in proximal part; ratio of carpus length to top hump position 2.93 ± 0.27 (N = 5), and ratio of carpus length to ending hump position 1.68 ± 0.17 (N = 5). Pleopod 1 ( Fig. 14D) exopod with pointed or narrowly rounded posterior apex, ratio of length to posterior apex width 4.67 ± 0.32 (N = 5), deeply concave outer margin, ratio of length to concavity turning point position 2.01 ± 0.07 (N = 5), slightly concave inner margin; endopod narrow, slightly enlarged at base, apical seta plumose. Pleopod 2 ( Fig. 14E) exopod triangular with concave outer margin; endopod distinctly longer than exopod, articles subequal in width, terminal seta strong.

Type locality: Höhle »Vitezičeva pećina« bei Rudine, OmiŠalj, Insel Krk ( Potočnik, 1983) [presently known as Biserujka (cave), Rudine, ČiŽići, Island of Krk, Croatia].

Remarks: Alpioniscus christiani is the smallest species in the strasseri- lineage. It differs from all other species of the lineage ( Fig. 12; Table 1) in having the lobe of the proximal part of the male pereopod 7 merus with teeth and the posterior apex of the male pleopod 1 exopod pointed instead of rounded. It differs in the ratio of the male pereopod 7 carpus length to carpus top hump position from A. hirci sp. nov. (1.66 ± 0.09) and A. velebiticus sp. nov. (2.08 ± 0.15); in the ratio of the male pereopod 7 carpus length to ending tergal hump from A. velebiticus sp. nov. (1.16 ± 0.06), A. iapodicus s.s. (1.26 ± 0.03) and A. hirci sp. nov. (1.27 ± 0.03); and in the ratio of the male pleopod 1 exopod length to posterior apex width from A. hirci sp. nov. (3.57 ± 0.19).

The citation with ‘?’ is doubtful, and the material from this locality needs to be re-examined with both molecular and morphological analyses.

Distribution: Croatian island of Krk and Dramalj surroundings ( Fig. 4).

ALPIONISCUS (ILLYRIONETHES) BALTHASARI ( FRANKENBERGER, 1937)

( FIGS 12, 15–19; TABLE 1)

Titanethes albus – Girometta, 1913: 13–16. – 1914: 3–8 [nec Špilja na Bardarovici (Vis Island); jama Propod (Hvar Island)]. – Langhoffer, 1915a: 19–20 [nec Ledenica kod Lokava; Mlađenova pećina; VraŽić- pećina; Ledenica; Pustinja; Vrlovka; VraŽić- pećina; pećina Jezero; Baračeva pećina II; Zobenica-pećina; Šupljara].

Illyrionethes balthasari Frankenberger, 1937: 175– 176 , figs 1–6. – 1938: 26. – 1939: 99. – Frankenberger & Strouhal, 1940: 449. – Strouhal, 1938: 273–274. – 1939b: 116; 119–122, figs 1, 2. – 1939c: 7, 22, 25, 28, 31. – 1939d: 17–18. – 1940a: 94. – Flasarová, 1967: 203–205, figs 12–14.

Alpioniscus (Illyrionethes) balthasari – Vandel, 1946: 155. – Buturović, 1955: 131–133. – 1957: 7–49; fig. 35. – Schmölzer, 1965: 59; fig. 221. – Karaman, 1966: 6–7. – Potočnik, 1993: 83, 178. – Tabacaru, 1996: 35. – Horvatović, 2014: 125, 173–175, 223; figs 7.28.b, 7.33.

Alpioniscus balthasari – Vandel, 1947: 271. – Potočnik, 1989: 64. – Erhard, 1997: 3. – Schmalfuss, 2003: 13. – Bedek et al., 2006: 34, 92, 127. – Bedek, 2007: 411–420. – Bedek et al., 2011: 237–240, 259–266; fig. 9 [nec Manita peć]. – 2017: 213 [nec eastern part of Mt. Velebit]. – JalŽić et al., 2007b: 498 [nec Velika Paklenica]. – 2010: 20, 104–105. – 2013: 17, 30. – Ozimec et al., 2012: 181, 217. – Stoev et al., 2013: 20. – Senckenberg, 2018.

Alpioniscus sp. – Hlaváč et al., 2019: 92, 97, 98.

? Alpioniscus (Illyrionethes) balthasari – Horvatović, 2014: 125 [partim: Milića Špilja].

? Alpioniscus balthasari – Bedek et al., 2011: 260– 266 [partim: Čude; Rastovača; Velika pećina; Kusa nad Manastirskim lukama; Kad jaganjci utihnu; Ivina jama; Pisanica; BuŠljeta; Jama kod Jatare 2; Jatara; Vaganka; Gornja Cerovačka Špilja].

Material examined (collection details in Supporting Information, Table S1)

Cetina–Krka population : CROATIA. Dalmatia, Vrlika: Civljane, KotluŠa, KotluŠa (cave) ( CBSSC IT1599, IT1602, IT1603, IT1601, IT1600, IT1654, IT1655, IT4028); GlavaŠ, Klačina draga, Babića pećina (cave) ( CBSSC IT4041); Kijevo, Golubinjka kod ValidŽića (cave CCC HR00595) ( CBSSC IT4040); Civljane, Tutići, Gornja Tutićeva Špilja (cave) ( CBSSC IT1616); Civljane, Četnici, Gospodska Špilja (cave) ( SMNS 5365, 5348; CBSSC IT1605, IT1607, IT1604, IT1606, IT1608, IT1609, IT4031, IT4032, IT4030, IT4029); Dinara Mt., GlavaŠ, Pekasova draga, Pećine (cave) ( CBSSC IT1618); Vukovići, Civljane, Rudelića Špilja (cave) ( CBSSC IT1610, IT1611, IT1612); Koljane, Spilja ispod manastira Dragović (cave CCC HR00604) ( CBSSC IT4038, IT4039); Kijevo, Velika Ćulumova pećina (cave CCC HR01591) ( SMNS 5356; CBSSC IT1656). Dalmatia, Krka surroundings: Kistanje, Lalići, Lalića stanovi, Drenovača (cave) ( CBSSC IT2100, IT278, IT280, IT281, IT2099, IT2101, IT2102, IT2103, IT2873); Skradin, Golubinka (cave) ( CBSSC

exopod length to apex width. The edge of each box marks the 25th and 75th percentiles; the whiskers show the largest and smallest values; yellow line indicates the median; and thick blue lines indicate the mean and SD.

IT2221); Benkovac, Bjelina, Golubinka kod Bjeline (cave CCC HR01468) ( CBSSC IT3955); Kistanje, Grulovići, Golubnjača u Grulovićima (cave CCC HR00250) ( CBSSC IT2857, IT2858, IT2859); Oklaj, Ljubotić, Jagodnjak (cave) ( CBSSC IT2871, IT2872); DrniŠ, LaŠkovica, RoŠki slap, Jama na Bibnovcu (cave CCC HR01364) ( CBSSC IT2839, IT2833, IT2838); DrniŠ, LaŠkovica, RoŠki slap, Jama na Selinama (cave CCC HR01290) ( CBSSC IT2875); DrniŠ, Ključ, kanjon Čikole, Torak, Jama nasuprot Torka (cave CCC HR01264) ( CBSSC IT2222, IT3986); Oklaj, Puljanje, Jazinka (cave) ( CBSSC IT2204, IT2205, IT2868, IT2869); DrniŠ, Ključ, Kaverna u kanjonu Čikole (cave) ( CBSSC IT2206, IT2207); Dalmatia, Vrlika, Jare, Crvene grede, Kranjica (cave) ( CBSSC IT1617); Oklaj, Ljubotić, Makeljina jama (cave) ( CBSSC IT2876, IT3978); Skradin, Martina jama (cave CCC HR01367) ( CBSSC IT305); Oklaj, HE Miljacka, Miljacka II (cave) ( CBSSC IT3412, IT298, IT300, IT301, IT3065, IT2156, IT2157, IT2159, IT2162, IT2163, IT2820, IT3982); Oklaj, HE Miljacka, Miljacka I-V (cave) ( CBSSC IT288, IT289, IT2212, IT2213, IT2224); Oklaj, Bogatići, RoŠki slap, Oziđana pećina (cave CCC HR01356) ( CBSSC IT2210, IT2211, IT3984, IT3985); Kistanje, Prčova pećina (cave CCC HR01306) ( CBSSC IT284); Kistanje, Burze, Sedrena Špilja iza mlina (cave CCC HR01269) ( CBSSC IT3140, IT2192, IT2194, IT2195, IT2847); Kistanje, Burze, Sedrena Špilja kod stola (cave CCC HR00244) ( CBSSC IT2201, IT2202, IT2203, IT2853); Kistanje, Burze, Sedrena Špilja s jamskim ulazom (cave CCC HR00245) ( CBSSC IT2196, IT2197, IT2198, IT2199, IT2200); Konjevrate, Gornji Krnjići, Punčka draga, Stara jametina (cave CCC HR01349) ( CBSSC IT303, IT304, IT2177, IT2178, IT2179, IT2180, IT2181, IT2182, IT2829, IT2830, IT3990, IT3991); Konjevrate, Škarići, Škarin Samograd (cave CCC HR00246) ( CBSSC IT286, IT3010, IT302, IT287, IT2885, IT2886); Skradin, Gornje Grahovo, Visovac, Špilja ispod Vukovića poda (cave) ( CBSSC IT2878, IT2884); Skradin, Špilja iznad Skradinskog mosta (cave CCC HR01550) ( CBSSC IT2225); Oklaj, HE Miljacka, Špilja kod mlina na Miljacki (cave) ( CBSSC IT293, IT3061, IT294, IT2214, IT2216, IT2228); Skradin, HE Jaruga, Špilja u Skradinskom buku (cave CCC HR01232) ( CBSSC IT2208, IT2209); DrniŠ, Ključ, kanjon Čikole, Štrikinica (cave CCC HR01191) ( CBSSC IT2219, IT2220); Kistanje, Lalići, Lalića ograde, Tavnica (cave CCC HR01579) ( CBSSC IT274, IT275, IT277); DrniŠ, Ključ, Kanjon Čikole, Topla pećina (cave CCC HR01217) ( CBSSC IT290, IT291, IT292, IT2184, IT2186, IT2189, IT3987, IT3988); Kistanje, Lalići, Lalića ograde, TriŠića jama (cave) ( CBSSC IT282, IT283, IT315, IT2874, IT3981, IT3979, IT3980); Kistanje, Burze, Trogrla (cave CCC HR00248) ( CBSSC IT2217, IT2218).

Biokovo–Mosor population : CROATIA. Dalmatia, Biokovo Mt.: Podgora, ŠoŠići, Štedovac, Baba (cave) ( CBSSC IT2540); Zagvozd, Gornje RaŠćane, Bebeci, Baba pećina (cave) ( CBSSC IT3813); Zavodi, Ravna vlaŠka, Benova jama (cave) ( CBSSC IT851); Župa, Lozovci, BoŠak (cave) ( CBSSC IT213, IT466); Makarska, Veliko Brdo, Bubnjavača (cave CCC HR00361) ( CBSSC IT495, IT3171, IT3173, IT2304, IT2480); Zagvozd, SvaguŠa, Čelar (a cave unknown at present) ( SMNS 5273); Bartulovići, Grljak, Ćulića jama (cave) ( CBSSC IT85, IT86); Gornja Brela, Bartulovići, Drinova, Drinova II (cave) ( CNHM 3/1417, CBSSC IT66, IT949, IT2264, IT3824); Zagvozd, Kaoci, Duboka trepetljika (cave) ( CBSSC IT92); Zagvozd, GačeŠe, Gaćina jama (cave) ( CNHM 5/1677); Gornja Brela, Bartulovići, Grljak, Gornja jama na Grljaku (cave) ( CBSSC IT215); Župa, Gradska spila (cave CCC HR00362) ( CNHM 425; CBSSC IT2799); Sv. Jure, Lipi Dočić, Jama iznad Lipog dočića (cave) ( CBSSC IT67); Gornja Brela, Nevistine stine, Jama iznad Prozorčića (cave) ( CBSSC IT83, IT884, IT82, IT246, IT472); Kozica, Saranač, Jama iznad Saranača (cave CCC HR00798) ( CBSSC IT99, IT3187, IT3149); Zagvozd, SvaguŠa, Jama kod SvaguŠe (a cave unknown at present) ( SMNS 5353); Tučepi, Ravna vlaŠka, Jama kod Tučepske vilenjače (cave CCC HR00802) ( CBSSC IT847); Prisika, Jama na grebenu (cave) ( CBSSC IT69, IT70, IT868); Topići, Brezdan, Jama na Brezdanu (cave) ( CBSSC IT260, IT223); Kozica, Saranač, Jama na Docima (cave) ( CBSSC IT89, IT90, IT918, IT895, IT3154); Zagvozd, Kačka glavica, Kaoci, Jama na Kačkoj glavici 3 (cave) ( CBSSC IT94); Zagvozd, Dedići, Jama na MrŽdolcu (cave CCC HR01853) ( CBSSC IT443); Sv. Jure, Jama na putu za Staru Školu (cave) ( CNHM 3/1416); Sv. Jure, Jama pod Sv. Jurom (cave) ( CBSSC IT849, IT850, IT3815); Makarska, VoŠac, Jama pod VoŠcem (cave) ( CBSSC IT3144); Zagvozd, ČikeŠi, Jama u ČikeŠima (cave) ( CBSSC IT490); Zagvozd, Kaoci, Jama u trepetljikama (cave) ( CBSSC IT945); Podgora, Vrata Biokova, Supin, Jama za Supinom (cave CCC HR00959) ( CBSSC IT867, IT241, IT459, IT2535); Nevistine stine, Jamina (cave) ( CBSSC IT886, IT764); Kozica, Jujnovići, Jujnovića Špilja (cave CCC HR00363) ( CNHM 4/1418, ( CBSSC IT865, IT483, IT484, IT3189, IT2544, IT3195); Makarska, Velo Brdo, Krjava 1 (cave) ( CBSSC IT42); Makarska, Velo Brdo, Krjava 2 (cave) ( CBSSC IT43, IT862); BaŠka Voda, Bast, Kukor (cave) ( CBSSC IT11, IT41, IT830, IT832, IT229, IT225, IT228, IT508, IT501, IT503, IT750, IT751, IT3183, IT3191, IT3148, IT2270, IT2301, IT2539, IT3823); Čedor, Kuna (cave) ( CBSSC IT856, IT858); Sv. Jure, Ledenica kod Stare Škole (cave) ( CNHM 1/1419); Župa, Mala jama (cave) ( CBSSC IT97); Zagvozd, Spiljice, Kaoci, Mala jama u stijenama (cave) ( CNHM 4/1676); Župa, MatijaŠeva peć (cave CCC HR01820) ( CNHM 4/1600, CBSSC IT435, IT3184); Župa, Lozovci, MravinjuŠa (cave) ( CBSSC IT207); Zagvozd, SvaguŠa, Sridnja gora, MuŽina Špilja (cave) ( CBSSC IT854, IT855); Topići, Pasji manastir (cave) ( CBSSC IT245); Gornja Brela, Nevistine stine, PloŠnica (cave CCC HR00055) ( CNHM 5/1601); Podgora, Podrum (cave CCC HR00255) ( CBSSC IT860, IT416, IT417); Dedići, Rastovac, Potkameničina jama (cave CCC HR00261) ( CBSSC IT756); Gornja Brela, RaŽanj, Pozjata (cave) ( CBSSC IT88); Makarska, Lokve, Pretnerova jama (cave) ( CBSSC IT3804; MZUF 9879); Zagvozd, Prodani, PruŽina (cave) ( CBSSC IT864, IT87, IT897, IT3806); Zagvozd, Kaoci, Pukotina kod doca StraŽbenica (cave) ( CNHM 7/1414; CBSSC IT899); Zagvozd, Rastovac, Raputinova Špilja (cave) ( CBSSC IT444, IT447, IT450); Gornja Brela, Drinova, Spasiteljica (cave) ( CBSSC IT925, IT915); Kozica, Jujnovići, Spila, Spila 1 (cave) ( CBSSC IT905, IT3817, IT3819); Kozica, Jujnovići, Spila, Spila 2 (cave) ( CNHM 3/1678; CBSSC IT84, IT907, IT206, IT256, IT3186, IT2315, IT3816); Župa, Roglići, Stonjska peć donja (cave CCC HR00264) ( CBSSC IT96, IT3803); Župa, Roglići, Stonjska peć gornja (cave) ( CBSSC IT95, IT2803); Gornja Brela, Dubci, Svetica (cave) ( CBSSC IT487); Zagvozd, KatuŠići, Špilja bei KatuŠići (a cave unknown at present) ( SMNS 5346); Zagvozd, Kaoci, Špilja do ObiljeŽene (cave) ( CBSSC IT943); Zagvozd, Dedići, Sridnja gora, Špilja na MrŽdolcu (cave) ( CBSSC IT441, IT442); Zagvozd, Brnasi, Špilja u Gaju (cave) ( CBSSC IT841, IT840); Tučepi, Srida sela, Ravna vlaŠka, Tučepska vilenjača (cave) ( CBSSC IT757, IT758, IT759, IT760, IT845, IT846, IT2633, IT2267, IT4086, IT4089, IT4096, IT4108; MZUF 9877, 9878); Zagvozd, Dedići, Bukovci, Velika Špilja (cave) ( CBSSC IT481, IT482); Zagvozd, Kaoci, Vodena trepetljika (cave) ( CBSSC IT93). Dalmatia, Mosor Mt.: Dugopolje, BaleŠina jama (cave) ( CBSSC IT4059, IT4060); Balići, Dugopolje, Balića Špilja (cave CCC HR00478) ( SMNS 5354, CBSSC IT2630, IT3046); Gornje Sitno, Jabukovac, Javor draga, Dana Špilja (cave) ( CBSSC IT4062); Kotelnice, Tukići, Bradarića staje, DrinovčuŠa (cave) ( CBSSC IT2959, IT3072, IT3074, IT3117, IT3126); Dugopolje, KriŽani, Ðuderina jama (cave) ( CBSSC IT3085); Gornja Poljica, Trnbusi, Jama u Dočiću (cave) ( CBSSC IT4067); Trilj, Ugljane, Jama u Krčevini (cave) ( SMNS 5352); Kotelnice, Tukići, Bradarića staje, Kravska jama (cave) ( CBSSC IT4045, IT4047, IT4048); Trilj, Ugljane, Majica pećina (a cave unknown at present) ( SMNS 5332); Kute, Čevrljin klen, Dugopolje, Maklutača (cave) ( CBSSC IT3124, IT3077, IT4050, IT4051); Žrnovnica, Miličevića pećina (cave) ( CBSSC IT4056, IT4057); Kolić, Dugopolje, Peć u Čulinovim raljevinama (cave CCC HR01164) ( CBSSC IT2991, IT3019, IT3137, IT3076); Dugopolje, Slipica (cave) ( SMNS 5327); Ljubljan, SnijeŽnica pod Ljubljanom (cave) ( CBSSC IT2973); Podgrade, Šćadin, Svinjarača (a cave unknown at present) ( CBSSC IT4066); Gornje Poljice, Trnbusi, Špilja Jame (cave) ( CBSSC IT3071); Gornji Dolac, Gornja Poljica, Trojama (cave CCC HR00875) ( CBSSC IT3090, IT3100, IT3108, IT2966, IT3050, IT4064); Gornje Sitno, Velika Gajina (cave) ( CBSSC IT4061); Kotlenice, Dugopolje, Vranjača (cave) ( SMNS 5320, 5043; CBSSC IT2631, IT2986, IT3051, IT3045, IT3012); Palin umac, Žmirino gumno, Dugopolje, Žmirina jama (cave) ( CBSSC IT4052, IT4054). Dalmatia, Omiška Dinara Mt.: Brela, Gornja Brela, Dovanj, Čolnjača (cave) ( CBSSC IT98, IT226); Zadvarje, Slime, Jama na podu Slime (cave) ( CBSSC IT3007); Zakučac, Naklice, Mandića Špilja (cave) ( CBSSC IT3040); Kučiće, Nikolina Špilja (cave) ( CBSSC IT2996). Dalmatia, Vrgorac: Šibenik, Kozica, Antunovići, Mala Špilja (cave) ( CNHM 4/1415; CBSSC IT3152); Stilja, Martina jama (cave) ( CBSSC IT306); Šibenik, Kozica, Antunovići, Velika Špilja (cave) ( CBSSC IT3192); Jurilji, Breskula u staroj paski (a cave unknown at present) ( SMNS 5105).

Redescription: Maximum length: ♂, 6.5 mm; ♀, 7.8 mm. Pereon with almost parallel sides. Back smooth, with ridges near posterior margins of cephalon and pereonites 1–4 ( Fig. 15A, B) (absent in populations from the cave Stara jametina) with some triangular scale-setae. Gland pores on lateral margins of pleonites 4 and 5, telson and uropodal protopods ( Fig. 15E). Cephalon ( Fig. 15D) with suprantennal line bent downwards, rounded middle part; antennal lobes quadrangular with central depression. Posterior margin of pereonite 1 convex, of pereonites 2 and 3 straight, and of pereonites 4–7 progressively more concave. Pleonites 3–5 with small posterior points visible in dorsal view ( Fig. 15A, E). Distal part of telson with concave sides and broadly rounded apex ( Fig. 15E). Antennula ( Fig. 16A) distal article flattened and bearing four to seven aesthetascs on apical margin. Antenna ( Fig. 16C) smooth; flagellum of eight to 13 articles, with one row of aesthetascs on two to seven articles, always present on second and third. Mouthparts as in A. strasseri . Pereopods ( Fig. 16D) similar in shape, with dactylar seta large and bifid. Uropod ( Fig. 15E) with protopod slightly grooved on outer margin; endopod inserted proximally.

Male: Pereopod 1 ( Fig. 16D) carpus bearing five to seven setae. Pereopod 1–3 with carpus and merus bearing numerous short scales on rostral surface. Pereopod 1–6 merus with sternal margin from straight (pereopod 1) progressively up to slightly concave (pereopod 6) and from none (pereopod 1) progressively up to small lobe (pereopod 6) in proximal part. Pereopod 7 ( Fig. 17A–C) merus with slightly concave sternal margin and small hook-shaped lobe in proximal part bearing one seta; carpus with straight sternal margin and large more or less rounded dorsal hump in proximal part, ratio of carpus length to top hump position 2.81 ± 0.33 (N = 25), and ratio of carpus length to ending hump position 1.45 ± 0.12 (N = 24). Pleopod 1 ( Fig. 18A) exopod with narrowly rounded posterior apex, ratio of length to posterior apex width 4.75 ± 0.36 (N = 25), deeply concave outer margin, ratio of length to concavity turning point position 2.23 ± 0.18 (N = 25), straight to slightly concave inner margin; endopod narrow with almost parallel sides, apical seta plumose. Pleopod 2 ( Fig. 18C) exopod triangular with narrowly rounded posterior apex, concave outer margin; endopod articles subequal in width; terminal seta strong and bifid.

Type locality: ‘eine Höhle in der Umgebung der Stadt Knin; es ist dies jene Höhle, die in der Nähe des Dorfes KotluŠa am westlichen Fussse des Dinara-Gebirges; »Höhle mit Wasser«’ ( Frankenberger, 1937) [presently known as KotluŠa (cave), KotluŠa, Civljane, Vrlika, Croatia].

Remarks: The Biokovo –Mosor population of A. balthasari shows some differences ( Figs 12, 15C, 16B, 17D, 18B, D, 19; Table 1), as follows: the body is considerably larger (♂, 8.2 mm; ♀, 10.7 mm), without ridges near posterior margins of cephalon and pereonites; the distal article of the antennula bears seven to 14 aesthetascs on the apical margin; the male pleopod 1 exopod has a broadly rounded posterior apex, ratio of length to posterior apex width 3.97 ± 0.28 (N = 33), ratio of length to concavity turning point position 1.70 ± 0.10 (N = 33), slightly concave inner margin; endopod slightly enlarged at base; the male pleopod 2 exopod has a broadly rounded posterior apex and the endopod is distinctly longer than the exopod, with the distal article narrower than the proximal. The population is geographically separated from A. balthasari s.s., and also shows some molecular differences. More sampling should be conducted in regions between their distribution areas, with morphological and molecular analyses in order to validate the taxonomic status of that population.

Alpioniscus balthasari differs from other species of the strasseri- lineage ( Fig. 12; Table 1) in the presence of ridges on the posterior part of cephalon and pereonites 1–4 (except the population from Stara jametina). Males differ in the ratio of the pereopod 7 carpus length to carpus top hump position from A. hirci sp. nov. (1.66 ± 0.09) and A. velebiticus sp. nov. (2.08 ± 0.15); in pereopod 7 carpus length to ending hump position from A. velebiticus sp. nov. (1.16 ± 0.06); in the ratio of the pleopod 1 exopod length to turning point position from A. strasseri s.s. (1.73 ± 0.07); and in the ratio of the pleopod 1 length to apex width from A. hirci sp. nov. (3.57 ± 0.19) and A. iapodicus (Paklenica) (3.68 ± 0.22).

This species is amphibious, because it was also observed in water.

The citations with ‘?’ are doubtful, and the material from those localities needs to be re-examined with both molecular and morphological analyses.

Distribution: Croatian river Cetina source surroundings, river Krka surroundings, Biokovo Mt., Mosor Mt., OmiŠka Dinara Mt. and their surroundings ( Figs 4, 19); Bosnia and Herzegovina, Tomislavgrad surroundings.

ALPIONISCUS (ILLYRIONETHES) IAPODICUS BEDEK, HORVATOVIĆ & KARAMAN, 2017

( FIGS 12, 20, 21; TABLE 1)

Titanethes ( albus ?) dahli – BoŽičević, 1965: 438 [partim: Manita peć].

Titanethes albus – Langhoffer, 1912: 363 [partim: Medina pećina]. – 1915b: 68 [partim: Medina pećina, Perusić környéke]. – Csiki, 1926: 73 [partim: Medina pećina] – Wolf, 1938: 77 [partim: Medina pećina].

Titanthes dahli – Strouhal, 1939b: 23 [partim: Medina pećina bei PeruŠić].

Alpioniscus sp. – Bregović et al., 2008: 110 [partim: Manita Peć, Velika Paklenica].

Alpioniscus balthasari – JalŽić et al., 2007b: 498 [partim: Velika Paklenica]. – Bedek et al., 2011: 261,

fig. 9 [partim: Manita peć]. – Bedek et al., 2017: 213 [partim: eastern part of Mt. Velebit].

Alpioniscus n. sp. 4 – Horvatović, 2014: 126, 174, 223, 240, figs 7.29, 7.33 [partim: pećina pri MuŠalučkom donjem selu; nekoliko pećina u Lici].

Alpioniscus n. sp. – Cindrić et al., 2017: 51–52.

Alpioniscus (Illyrionethes) iapodicus Bedek et al., 2017: 205–214 , figs 1–7.

Type material re-examined (collection details in Supporting Information, Table S1)

Holotype: ♂, Croatia, Lika, Donji Kosinj, Lipovo polje, Markov ponor (cave CCC HR01480 ) ( CBSSC IT4109 ) . Paratypes: Croatia, Lika : ibid. ( CBSSC IT2479 ; ZZDBE 1142 ; MZUF 9763 ) ; Ličko LeŠće , Pećina (cave) ( CBSSC IT3344 , IT3347 ) ; Donji Kosinj , Lipovo polje, Petranović Draga, Mramorna Špilja (cave CCC HR02057 ) ( CBSSC IT3367 ) ; PeruŠić , Studenci , Milkovići, Budina Špilja (cave CCC HR01195 ) ( CBSSC IT3373 , IT3843 , IT3848 , IT3846 , IT3374 ) ; Donji Kosinj , Mlakva , Javorinske drage, Pećina na Čakovcu (cave CCC HR01706 ) ( CBSSC IT3765 ) ; PeruŠić , Sitvuki, Sitvukova pećina (cave CCC HR01962 ) ( CBSSC IT3383 ) ; PeruŠić , Velika Plana , Rastovac , Prva poŠtena – Jama u Rastovcu (cave CCC HR00953 ) ( CBSSC IT3921 ) ; PeruŠić , Grabovača, Medina pećina (cave) ( CNHM 653 ; CBSSC IT2545 , IT3381 , IT3907 , IT3382 ) ; PeruŠić , Grabovača, AmidŽina pećina (cave) ( CBSSC IT2546 , IT3849 , IT3850 , IT3378 ) ; PazariŠte , Vranovine, Vranovinski ponor (cave CCC HR00490 ) ( SMNS 5276 ) ; Velebit , PazariŠte , Japage, Špilja Japa (cave CCC HR00344 ) ( CBSSC IT544 ) ; Gospić , MuŠaluk, Pećina pri muŠalučkom donjem selu (present unknown cave) ( ZZDBE 0899 ) ; Gospić , Klanac, Grčka pećina (present unknown cave) ( SMNS 5344 ) ;

566 J. BEDEK ET AL.

PazariŠte, Mlakva, Mlakvena greda (present unknown cave) ( SMNS 5295).

Material examined (collection details in Supporting Information, Table S1)

Lika population : Croatia, Velebit Mt. (N): Starigrad kod Senja, Velike Brisnice, Jama II kod Velikih Brisnica (cave) ( CBSSC IT411, IT412, IT580, IT582, IT583, IT3112, IT3427, IT3428); Senj, Donja Klada, Jama na KrŽeljevcu (cave) ( CBSSC IT542); Senj, Velike Brisnice, Jama na Vrančinici (cave) ( CNHM 2/1669; CBSSC IT587); Donja Klada, Špilja u Vukotića dragi (cave) ( CBSSC IT540, IT541, IT593). Croatia, Lika : Ogulin, DreŽnica, DreŽničko polje, Sopača (cave CCC HR01927) ( CBSSC IT3252); Bunić, Zelena pećina (cave) ( CBSSC IT3404).

Paklenica population : Croatia, Velebit Mt. (S): Starigrad Paklenica, Velika Paklenica, Manita peć (cave) (SMNS 5362, 5341; CBSSC IT14, IT15, IT1, IT562, IT563, IT2259, IT2260, IT2261, IT3211, IT2958, IT3208, IT3210, IT3212, IT3223, IT3225, IT3207, IT3221, IT3206, IT3202, IT3200, IT3218, IT3228, IT3229, IT3230, IT3231, IT3232, IT3204).

Additional data to description: Tergite with triangular scale-setae ( Fig. 20A). Antennula distal article bearing four to nine aesthetascs on apical margin. Antennal flagellum of eight to ten articles, with one row of aesthetascs on three to five different articles, always on second and third. Male pereopod 1 carpus bearing five to seven setae. Male pereopod 7, ratio of carpus length to carpus top hump position 2.10 ± 0.36 (N = 7), and ratio of carpus length to ending hump position 1.26 ± 0.03 (N = 7). Male pleopod 1 ratio of exopod length to posterior apex width 3.68 ± 0.52 (N = 7), with deeply concave outer margin, ratio of length to concavity turning point position 1.93 ± 0.10 (N = 7), straight to slightly concave inner margin; endopod apical seta plumose. Male pleopod 2 endopod of two articles ( Fig. 20C).

Type locality: Croatia, Lika, Donji Kosinj, Lipovo polje, Markov ponor (cave CCC HR01480) ( Bedek et al., 2017).

Remarks: The population from Manita peć, Paklenica ( Figs 12, 20B, 21; Table 1) shows some differences from A. iapodicus s.s., i.e. the male pereopod 7 merus with hook-shaped lobe directed ventrally; pereopod 7 ratio of carpus length to carpus top hump position 2.69 ± 0.18 (N = 6); and pereopod 7 ratio of carpus length to ending hump position 1.42 ± 0.09 (N = 6). The population is geographically separated from A. iapodicus s.s., and it also shows some molecular differences. More sampling should be conducted between the type locality and Paklenica area, in order to validate the taxonomic status of this population.

Alpioniscus iapodicus differs from all the other species of the strasseri- lineage in the male characters ( Fig. 12; Table 1). The hook-shaped lobe of the male pereopod 7 merus in the proximal part is directed laterally instead of ventrolaterally as in the A. balthasari Biokovo –Mosor population or ventrally as in all the other species. It differs in the ratio of the male pereopod 7 carpus length to ending tergal hump position from A. strasseri s.s. (2.09 ± 0.22) and A. christiani (1.68 ± 0.17).

Distribution: Croatian Lika and Velebit Mt. slopes ( Figs 4, 21).

ALPIONISCUS (ILLYRIONETHES) HIRCI BEDEK &