Haasea Verhoeff, 1895

Antić, Dragan & Akkari, Nesrine, 2020, Haasea Verhoeff, 1895 - a genus of tumultuous history and chaotic recordsredefinition, revision of taxonomy and geographic distributions, with descriptions of two new species from Austria and Serbia (Diplopoda, Chordeumatida, Haaseidae), Zootaxa 4798 (1), pp. 1-77 : 7-11

Genus Haasea Verhoeff, 1895

Xiphogona Cook, 1895: 3 (junior objective synonym)

Xiphigona Cook, 1895 —misspelled by Hoffman (1980: 130) (junior objective synonym)

Orobainosoma Verhoeff, 1897: 135 (junior objective synonym)

Rhopalogona Silvestri, 1898: 666 (junior objective synonym)

Deuterohaasea Verhoeff, 1898: 226 (junior objective synonym)

Orobainosoma (Brachybainosoma) Verhoeff, 1899: 128 (junior subjective synonym)

Heterohaasea Verhoeff, 1900: 362 (junior objective synonym)

Orobainosoma (Histrosoma) Strasser, 1973a: 419 (junior subjective synonym)

Type species: Craspedosoma flavescens Latzel, 1884 accepted as Haasea flavescens ( Latzel, 1884) [by subsequent designation of Cook, 1895 in Cook & Collins (1895: 3)]

Diagnosis: A genus of Haaseidae with leg-pair 10 in males reduced in size compared to other walking legs (vs. robust in Hylebainosoma and Xylophageuma , an exception is H. nontronensis also with a reduced leg-pair 10 but this species differs from Haasea by numerous other gonopod and vulvar structures). Medial sternal process of the anterior gonopodal block bilobed distally (vs. not bilobed in Hylebainosoma and Xylophageuma ). Angiocoxites sigmoid in lateral or mesal view (vs. mainly straight in Hylebainosoma and Xylophageuma , an exception is H. gulickai Tajovský, Mock & Papáč, 2014b with sigmoid angiocoxites, but this species differs from the other haaseids by the absence of ommatidia). Additional gonopodal processes mainly strongly curved posteriad in lateral or mesal view (vs. mainly straight in Hylebainosoma and Xylophageuma ). Vulval bursae mesally completely merged.

Short description: Body in adults with 28 or 30 segments (including telson); 6.5–12 mm long.

COLORATION ( Fig. 1 ). Variable, from poorly pigmented to light or darker brown. Most of species with yellowish-brown body.

HEAD ( Fig. 2 A–F). Setose, frontal side flattened in males ( Fig. 2A ), slightly convex in females. Labrum with three medial teeth ( Fig. 2B ). Antennae elongated, slender, with dorsomedial and ventromedial lines of small papillae-like outgrowths ( Fig. 2 D–F). Lateral to antennal sockets a group of similar papillae-like outgrowths present ( Fig. 2C ). Ommatidia brownish to black arranged in triangle, number from 9–16 ( Fig. 2A ).

BODY ( Figs 1 , 2 G–K). With lateral humps. Macrochaetae long and trichoid, present on large knobs. Posterior metazonal margins with small, triangular denticles. Collum and telson not modified.

LEG- PAIRS 1 AND 2 ( Fig. 3E ). In both sexes with tarsal combs; prefemora with several long and robust setae; femora and postfemora each with a group of several long and robust setae.

MALE SEXUAL CHARACTERS ( Fig. 3 E–G). Leg-pair 2 with genital openings on coxae ( Fig. 3E ). Leg-pairs 3–7 enlarged. Distal half of tarsi 3–7 with ventral papillae ( Fig. 3F ). Leg-pair 10 significantly smaller than the other walking legs; with coxal sacks and with a robust, blunt coxal process ( Fig. 3G ). Leg-pair 11 with coxal sacks, no other modifications.

ANTERIOR GONOPODS ( Fig. 3 A–C) Sternum (s) wide, with medial sternal process (msp, see Figs 25D and 34C ) bilobed distally and anterior sternal lobe (asl) with several long setae, on both lateral sides. Long setae (1–3) also present on antero-paramedial and lateral parts of s. Angiocoxites (a) erected, sigmoid in lateral or mesal view, characterized by the presence of a deep, posterior and longitudinal furrow (f, see Figs 9B , 14C , 18C, E or 34D), extending from the base to the distal part, dividing superficially posterior portions of the angiocoxite into the mesal and lateral sides. Angiocoxites supplied with a few additional processes or extensions on both mesal and lateral sides. Mesal process (mp) always present, more or less well-developed, with or without additional teeth. Apart from mp, two more structures are always present on mesal side, viz., papillated cone (pc) and “ Opuntia -like” membranous lobe (ml) which might be a colpocoxite. Both structures more or less identical in all species. Laterally, two processes present, viz., anterior lateral process (alp) in only a few species and posterior lateral process (plp) in all species. The distal half of a bends anteriad. The most distal part (dp) complex, composed of structures with “hairs” or fimbriaelike outgrowths. In some species, this part has a mesodistal lobe (mdl) sometimes with fimbriae. Posteriorly, dp has a seminal opening (so) surrounded by minute fimbriae-like outgrowths. Below so, posterior lamella pl (sometimes also pl1, pl2) present, simple or bilobed. Inside a, (?) seminal canal (s c) is visible, starts from the basal half and opens distally in so. The most basal part of sc enlarged, in form of seminal chamber (sch) and ends with a cone structure (cs) covered with cilia or sensilla.

POSTERIOR GONOPODS ( Fig. 3D ). With reduced sternum (s, see Fig. 26J ). Telopodites (t) well-developed, robust, with two podomeres, covered by long setae. Coxal processes (cp) present mesally, with two or three branches (ab, mb, pb).

LEG- PAIR 3 IN FEMALES ( Fig. 3 H–J). With lateral sternal lobes (sl).

VULVAE ( Fig. 3H ). Operculum (o, see Figs 35G and 37A ) membranous, with serrated edge and with a row of long setae. Bursae (b) merged mesally with bulbous structure, long setae and triangular denticles present on mesal and lateral parts. Postgenital plate (pgp) present, with lateral lobes and medial lobe. For vulvae details see Kurnik (1988: 234, taf I, 246–251).

Figures

FIGURE 1. Living Haasea Verhoeff, 1895. A. H. germanica (Verhoeff, 1901), female from Reiflding, Germany. B. H. ger- manica (Verhoeff, 1901), mating pair from Riedl, Germany. C. H. flavescens (Latzel, 1884), male from Schwaighofen, Austria. D. H. flavescens (Latzel, 1884), mating pair from Gaisberg, Austria. E. H. hungarica (Verhoeff, 1928), male from Fruška gora Mt., Serbia. F. H. vidinensis (Strasser, 1973a), female from the Ponor Radovanske reke Cave, Serbia. Photos credit: A–D. Jörg Spelda, E. Dragiša Savić (photo in lab), F. Nikola Vesović (photo in lab).

FIGURE 2. General habitus of Haasea Verhoeff, 1895. A–K. H. flavescens (Latzel, 1884). A, C–E, H–K. Male from Lower Austria, Rekawinkel, Vienna Woods. B, F. Female from Austria, Vienna, Heuberg, Vienna Woods. G. Male from Austria, Vienna, Heuberg, Vienna Woods. A. Head, lateral view. B. Head, anterior view. C. Papillae-like outgrowths lateral to antennal socket, lateral view. D. Right antenna, lateral view. E. Right antennomere 6, ventrolateral view. F. Antennomeres 4 and 5, ventral view. G. Segment 15, dorsal view. H. Apex of midbody macrochaeta. I. Posterior margin of midbody metazona. J. Midbody segments, dorsolateral view. K. Body end, dorsolateral view. Scale bars: 0.2 mm (A, B), 0.1 mm (C, D, F, G, K), 0.05 mm (J), 0.02 mm (E, I), 0.01 mm (H).

FIGURE 3. General sexualcharacters in Haasea Verhoeff, 1895. A. H. germanica (Verhoeff, 1901), male from Krummnussbaum, Austria, left anterior gonopod, mesal view. B. H. makarovi sp. nov., paratype male (IZB), left anterior gonopod, mesal view. C. H. makarovi sp. nov., paratype male (IZB), left anterior gonopod, base of left anterior gonopod, mesal view. D. H. flavescens (Latzel, 1884), male from Lower Austria, Rekawinkel, right posterior gonopod, mesal view. E. H. makarovi sp. nov., paratype male (NHMW9474), left leg 2, posterior view. F. H. makarovi sp. nov., paratype male (NHMW9474), right tarsus 3, anterior view. G. H. makarovi sp. nov., paratype male (NHMW9474), left leg 10, anterior view. H. H. flavescens (Latzel, 1884), female from Austria, Vienna, Heuberg, Vienna Woods, vulvae, ventral view. J. H. flavescens (Latzel, 1884), female from Austria, Vienna, Heuberg, Vienna Woods, sternum 3, lateral view. Abbreviations: a—angiocoxite; ab—anterior branch; alp—anterior lateral process; asl—anterior sternal lobe; b—bursa; cs—cone structure; cp—coxal process; dp—distal part; mb—medial branch; mdl—mesodistal lobe; ml—“Opuntia-like” membranous lobe; mp—mesal process; pb—posterior branch; pc—papillated cone; pgp—postgenital plate; pl—posterior lamella; plp—posterior lateral process; s—sternum; sc—seminal canal; sch—seminal chamber; sl—sternal lobe; so—seminal opening; t—telopodite. Scale bars: 0.1 mm (A–E, G–J), 0.05 mm (F).

FIGURE 9. Haasea flavescens (Latzel, 1884). A–F. Male from Lower Austria, Rekawinkel. G, H. Male from near Vienna, Austria. A. Left anterior gonopod, mesal view. B. Left anterior gonopod, posterodistal view. C. Left anterior gonopod, lateral view. D. Left anterior gonopod, detail of distal part, mesal view. E. Left anterior gonopod, detail of distal part, distal view. F. Right posterior gonopod, mesal view. G. Right posterior gonopod, mesal view. H. Right posterior gonopod, posterior view. Abbreviations: a—angiocoxite; ab—anterior branch; asl—anterior sternal lobe; cp—coxal process; dp—distal part; f—furrow; mb—medial branch; mdl—mesodistal lobe; ml—“Opuntia-like” membranous lobe; mp—mesal process; pb—posterior branch; pc—papillated cone; pl—posterior lamella; plp—posterior lateral process; so—seminal opening; t—telopodite. Scale bars: 0.1 mm (A–C, F–H), 0.05 mm (E), 0.02 mm (D).

FIGURE 14. Haasea germanica (Verhoeff, 1901), male from Krummnussbaum, Austria. A–E. Left anterior gonopod. F–H. Right posterior gonopod. A. Mesal view. B. Lateral view. C. Distolateral view. D. Detail of distal part, mesal view. E. Detail of distal part, distal view. F, G. Posterior view. H. Mesal view. Abbreviations: a—angiocoxite; ab—anterior branch; asl—anterior sternal lobe; cp—coxal process; dp—distal part; f—furrow; mb—medial branch; mdl—mesodistal lobe; ml—“Opuntia-like” membranous lobe; mp—mesal process; pb—posterior branch; pc—papillated cone; pl—posterior lamella; plp—posterior lateral process; s—sternum; so—seminal opening; t—telopodite. Scale bars: 0.1 mm (A–C, F–H), 0.05 mm (D, E).

FIGURE 18. Haasea inflata (Verhoeff, 1907), males from Posruck, Schlossberg, Austria. A. Right anterior gonopod, lateral view. B. Left anterior gonopod, mesal view. C. Right anterior gonopod, posterodistal view. D. Right anterior gonopod, detail of distal part, lateral view. E. Left anterior gonopod, lateral view. F. Right posterior gonopod, mesal view. G. Right posterior gonopod, mesal view. H. Right posterior gonopod, anterior view. Abbreviations: a—angiocoxite; ab—anterior branch; asl—anterior sternal lobe; cp—coxal process; dp—distal part; f—furrow; mdl—mesodistal lobe; ml—“Opuntia-like” membranous lobe; mp—mesal process; pb—posterior branch; pc—papillated cone; pl1, pl2—posterior lamellae; plp—posterior lateral process; so—seminal opening; sph—spermatophore; t—telopodite. Scale bars: 0.1 mm (A–C, E–H), 0.05 mm (D).

FIGURE 25. Haasea vidinensis (Strasser, 1973a). A. Allotype female of H. guidononveilleri Makarov, 2008 syn. nov. (IZB), habitus, lateral view. B. Male from Kučaj Mts, Serbia, right anterior gonopod, mesal view. C. Male from Kučaj Mts, Serbia, right anterior gonopod, lateral view. D. Male from Kalafat Mt, Serbia, left anterior gonopod, distomesal view. E. Male from Kalafat Mt, Serbia, right anterior gonopod, detail of distal part, posterodistal view. F. Male from Kalafat Mt, Serbia, right anterior gonopod, detail with lateral processes, lateral view. Abbreviations: a—angiocoxite; alp—anterior lateral process; dp—distal part; f—furrow; mdl—mesodistal lobe; ml—“Opuntia-like” membranous lobe; mp—mesal process; msp—medial sternal process; pc—papillated cone; pl—posterior lamella; plp—posterior lateral process; so—seminal opening. Scale bars: 1 mm (A), 0.1 mm (B–E), 0.01 mm (F).

FIGURE 26. Haasea vidinensis (Strasser, 1973a). A. Lectotype male (NMNHS10807), right anterior gonopod, lateral view. B. Paralectotype male (NMNHS10808), left anterior gonopod, lateral view. C. H. guidononveilleri Makarov, 2008 syn. nov., holotype male (IZB), left anterior gonopod, lateral view. D. Male from Rtanj Mt, Serbia, right anterior gonopod, lateral view. E. Male from Kučaj Mts, Serbia, left anterior gonopod, lateral view. F. Male from Kučaj Mts, Serbia, left anterior gonopod, mesal view. G. Lectotype male (NMNHS10807), right posterior gonopod, anteromesal view. H. Male from Kučaj Mts, Serbia, right posterior gonopod, mesal view. I. H. guidononveilleri Makarov, 2008 syn. nov., holotype male (IZB), posterior gonopods, posterior view. J. Male from Rtanj, Mt, Serbia, posterior gonopods, posteror view. Abbreviations: ab—anterior branch; cp—coxal process; pb—posterior branch; plp—posterior lateral process; s—sternum t—telopodite. Scale bars: 0.1 mm.

FIGURE 34. Haasea makarovi sp. nov., paratype male (NHMW9474). A. Right tarsus 3, anterior view. B. Leg 7, detail with ventral papillae, ventral view. C. Anterior gonopods, anterior view. D. Anterior gonopods, posterodistal view. E. Anterior gonopods, detail of distal part, anterior view. F. Anterior gonopods, detail of distal part, posterodistal view. Abbreviations: a—angiocoxite; alp—anterior lateral process; asl—anterior sternal lobe; f—furrow; mdl—mesodistal lobe; ml—“Opuntia- like” membranous; mp—mesal process; msp—medial sternal process; pc—papillated cone; plp—posterior lateral process; s—sternum; so—seminal opening; Scale bars: 0.1 mm (C, D), 0.05 mm (A, E, F), 0.01 mm (B).

FIGURE 35. Haasea makarovi sp. nov., A. Paratype male (IZB), left anterior gonopod, lateral view. B. Paratype male (IZB), left anterior gonopod, mesal view. C. Paratype male (IZB), left anterior gonopod, detail of distal part, lateral view. D. Paratype male (IZB), left anterior gonopod, detail of distal part, mesal view. E. Paratype male (NHMW9474), anterior gonopods, detail, posterodital view. F. Paratype male (NHMW9474), left posterior gonopod, mesal view. G. Paratype female (IZB), vulvae, ventral view. Abbreviations: a—angiocoxite; ab—anterior branch; alp—anterior lateral process; b—bursa; cp—coxal process; f—furrow; mb—medial branch; mdl—mesodistal lobe; ml—“Opuntia-like” membranous lobe; mp—mesal process; o—operculum; pb—posterior branch; pc—papillated cone; pgp—postgenital plate; plp—posterior lateral process; so—seminal opening; t—telopodite. Scale bars: 0.1 mm (A, B, F, G), 0.05 mm (C, D, E).

FIGURE 37. Haasea makarovi sp. nov., paratype female (IZB).A. Vulvae, ventral view. B. Postgenital plate, posterior view. C. Leg-pair 3, Anterior view. Abbreviations: b—bursa; o—operculum; sl—sternal lobe. Scale bars: 0.1 mm.