Travuniidae Absolon & Kratochvíl, 1932 version 2.5

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1 Travuniidae Absolon & Kratochvíl, 1932 version 2.5 Adriano B. Kury Departamento de Invertebrados, Museu Nacional/UFRJ Quinta da Boa Vista, São Cristóvão, , Rio de Janeiro - RJ BRAZIL 1. Introduction The Travuniidae have been regarded as a relictual group, represented by a few species of tiny delicate laniatorids of the Northern Temperate areas, mainly in Southeastern Europe, with one species from lava tubes in Western USA and another cavedwelling in Japan. Most of the, 18 known species have been found only in caves. Because of the unique special claw structure called the peltonychium, the Travuniidae have been recognized relatively early, while the related family Cladonychiidae had many species included in the groundless subfamily Tricommatinae of Roewer. Adequate descriptions lack for most species and taxonomy will still suffer changes Subtaxa included. There are two subfamilies not used by any author, Peltonychiinae (distitarsi I with 2 joints) and Travuniinae (distitarsi I with 3 joints), 9 nominal genera and 18 nominal species. Status of genera and species is highly unstable. Kratochviliola is much probably a synonym of Peltonychia (Rambla, 1973). A great deal of synonymies is to be expected. Included genera, in parentheses number of valid species, hy = hypogean species, ep = epigean: Abasola Strand, 1928 (3, hy) BOSNIA & HERZEGOVINA, CROATIA, SERBIA & MONTENEGRO. Arbasus Roewer, 1935 (1, hy) FRANCE. Buemarinoa Roewer, 1956 (1, hy) ITALY (Sardinia). Dinaria Hadži, 1933 (1, hy) BOSNIA & HERZEGOVINA. Kratochviliola Roewer, 1935 (1, hy) FRANCE. Peltonychia Roewer, 1935 (7, hy/ep) FRANCE, ITALY, SPAIN, SWITZERLAND. Speleonychia Briggs, 1974 (1, hy) Western USA. Travunia Absolon, 1920 (2, hy) BOSNIA & HERZEGOVINA, CROATIA. Yuria Suzuki, 1964 (1, hy/ep) JAPAN.

2 1.2. Systematic historical background This family includes the worst taxonomic mêlée of the laniatorid story in 20th century. First travuniiid ever described was Scotolemon leprieurii Lucas, 1860 in Phalangodidae. Followed by Phalangodes claviger Simon, 1879 and Scotolemon piochardi Simon, Then in the beginning if the 20th century: Phalangodes caecus Simon, 1911, Absolonia troglodytes Roewer, 1915 and Scotolemon anophthalmum Absolon, All six species were in Phalangodidae. Absolon (1920) created the genus Travunia as a replacement for Absolonia, preoccupied and made many mistakes and confusion with nomenclatorial rules. Travunia is the Latin name of the city Trebinje in Herzegovina. Hadži (1933) also stated that his new species Travunia vjetrenicae (Hadži, 1932) should constitute the new genus Dinaria, but inconsistently continued treating it as a member of Travunia. Roewer (1935) reconciled this paradox considering that Dinaria has been proposed as a subgenus of Travunia. The family Travuniidae was for the first time recognized by Hadži (see Hadži s claims in Hadži, 1933), but he started to share his knowledge with Kratochvíl, who acted quickly and had it published first (Absolon & Kratochvíl, 1932a). They used the name Peltaeonychidae, although at that time there was only the generic name Travunia included with two species. So, family name was not based on the name of the type genus. Aware of that, the authors correctly changed family name to Travuniidae a few months later (Absolon & Kratochvíl, 1932b). The genus Peltonychia was to be created by Roewer only three years later. Hadži (1933) considered Travuniidae to be a subfamily of Triaenonychidae, in the same way he would later consider his Cladonychiinae (1935), being thus the first author to recognize the Travunioidea in Martens sense (Martens, 1980). Roewer (1935) reviewed the European Laniatores and placed all of the above cited Phalangodidae in Travuniidae, described many new genera and species. The family was divided into two subfamilies Peltonychiinae and Travuniiinae (only with genus Travunia) by Kratochvíl (1958), who tried to mimic the unsuccessful Roewerian dichotomy Phalangodinae versus Tricommatinae, but subsequent authors did not adopt them. The three genera described later (one each decade 50s, 60s, 70s) have not been assigned to any of the subfamilies. Martens (1978) cast doubt on the validity of many genera, but formally proposed only a few generic and specific synonymies. For about 30 years no new species of Travuniidae has been discovered Natural history. The travuniids are delicate laniatorids which are nowhere of outstanding importance. Some species have been regarded as troglobites or troglophiles (Marcellino, 1982), while others are also found outside caves. Suzuki (1975) reported Japanese travuniids collected

3 in caves and surrounding areas by sifting leaf debris in dense forests between 700 and 1000 m. Maybe it is more accurate to speak of "hypogean species" than cavernicolous because many of them live indeed in fissure and crack systems (of the epikarstic habitats) and are therefore rarely to be found in caves. 2. Characterization Size. Small Laniatores, 1-3 mm. Body dorsum/venter. Body convex, mostly rounded posteriorly, only slightly constricted in anterior third. Frontal border of carapace unarmed. Segments of body ill-marked by incomplete grooves, mostly lacking. All areas, tergites and sternites unarmed. Eye mound when present, low, granular, far from the frontal border of carapace. Eyes may be reduced and depigmented. Ninth tergite and lateral free sclerites present in non-european genera. Sternum wedge-shaped. Pedipalps robust and strongly spined, femur dorsally convex, with ventral row of setiferous tubercles and mesal subapical setiferous tubercle. Tibia and tarsus with powerful mesal and ectal setiferous tubercles. Setae inserted subdistally in sockets. Legs I-IV slender and unarmed. Claws III-IV with peltonychium (complex claw formed by central shield and many pairs of lateral branches, sometimes asymmetrical) attached to a stem at distal part of tarsus. Tarsal counts: leg I 3-6 (distitarsi 2-3), leg II 5-6 (distitarsi 3-4) leg III 4, leg IV 4, only Travunia has such high counts as 6(3) in leg I, other genera have 3-5(2). Musculature of penis often reduced to basal portion of truncus. Glans with sclerites fused including the stylus. Ovipositor unsegmented, four-lobed, with only scattered setae. Color in general uniform dark yellow to pale yellow, troglobites are depigmented. No secondary sexual dimorphism. 3. Distribution Europe, Japan and United States. The records for Slovenia are all mistaken (Novak & Gruber, 2000). 4. Relationships Branched claws are present in most Triaenonychidae, but the peltonychium as a unique complex claw would be a potential synapomorphy for the genera of Travuniidae. However, similar structures developed in many apparently unrelated Travunioidea such as Synthetonychiidae (Forster, 1954), some Australian Triaenonychidae of genus Lomanella (Hunt & Hickman, 1993), the Argentinean troglobite Triaenonychidae Picunchenops (Maury, 1988). So, the monophyly of Travuniidae is not corroborated by any unique structure, but

4 even so the presence of peltonychium could work as a synapomorphy (convergently appeared in other Travunioidea). The Travuniidae are most closely related to the Cladonychiidae because of 1) the musculature of penis is restricted to the base and 2) the complex of glans is short with all median and dorsal components fused in a single structure. It is possible that the Travuniidae are paraphyletic in relation to the Cladonychiidae, because some of its genera (at least Peltonychia) appear closer to this family in genital morphology. The replacement of the peltonychium for a cladonychium could be a synapomorphy for the Cladonychiidae in a scenario of a paraphyletic Travuniidae. The presence of additional abdominal sclerites in genera Yuria and Speleonychia is a retention of a plesiomorphic state, shared with the Pentanychidae only in Laniatores. This would add support to a paraphyletic Travunioidea but would need an extra ad hoc hypothesis of independent loss of those sclerites 1) in other Travuniidae plus Cladonychiidae and 2) all other Laniatores. 5. Acknowledgements I wish to thank Dr Tone Novak (University of Maribor, Slovenia) for important critics, comments and suggestions on the draft version. 6. References Absolon, Karel. 1921: O mikrofotografování neprůhledných drobných předmětů. Druhé sdělení. [On microphotographing of opaque small objects. 2nd part]. Časopis moravského musea zemského, Brno, [ ]: Absolon, K. & J. Kratochvíl, 1932a. Peltaeonychidae, une famille nouvelle des Opilionides aveugles des grottes balcaniques [1]. Príroda, Brno, 25(5): Absolon, K. & J. Kratochvíl, 1932b. Peltaeonychidae, une famille nouvelle des Opilionides aveugles des grottes balcaniques [2]. Príroda, Brno, 25(6): Absolon, K. & J. Kratochvíl, 1932c. Zur Kenntnis der höhlenbewohnenden Araneae der illyrischen Karstgebiete. Mitt. Höhlen- u. Karstforschung, 3: Forster, R. R., The New Zealand harvestmen (sub-order Laniatores). Canterbury Mus. Bull., 2:

5 Hunt, G. S. & J.L. Hickman, A revision of the genus Lomanella Pocock and its implications for family level classification in the Travunioidea (Arachnida: Opiliones: Triaenonychidae). Rec. Austr. Mus., 45: Hadži, J., Prilog poznavanju pecinske faune Vjetrenice. (Pseudoscorpionidea: Neobisium (Blothrus) vjetrenicae sp. n., Opilionidea: Travunia vjetrenicae sp. n., Nelima troglodytes Roewer). Glas Srpske kralj. akadem Beograd. CLI. prvi razred, 75: Hadži, J., Beitrag zur Kenntnis der Fauna der Höhle Vjetrenica. Bull. Acad. Sci. Mat. Natur. B. Sci. Natur., Beograd 1: Hadži, J., Ein eigentümlicher neuer Hölen-Opilionid aus Nord-Amerika, Cladonychium corii g.n. sp. n. Biologia Generalis, 11: Kratochvíl, J., Die Höhlenweberknechte Bulgariens (Cyphophthalmi und Laniatores). Práce Brn. zákl. cesk. Akad. Véd., 30(375): Marcellino, I., 1982, Opilioni cavernicoli italiani, Lav. Soc. Ital. Biogeogr., N.S., 7: Martens, J., Genitalmorphologie, System und Phylogenie der Weberknechte (Arachnida: Opiliones). Ent. Germanica, 3(1-2): Martens, J., Weberknechte, Opiliones. Die Tierwelt Deutschlands 64: 464 pp. Martens, J., Versuch eines phylogenetischen Systems der Opiliones. Verh. 8. intern. Arachn. Kongr. Wien: Martens, J., Die Grossgliederung der Opiliones und die Evolution der Ordnung (Arachnida). Actas 10 Congr. Aracnol. Jaca España, 1986, 1: Maury, E.A., Triaenonychidae sudamericanos V. Un nuevo género de opiliones cavernícolas de la Patagonia (Opiliones, Laniatores). Mémoires de Biospéologie 15: Novak, T. & J. Gruber, Remarks on published data on harvestmen (Arachnida: Opiliones) from Slovenia. Annales (Koper) Ser. hist. nat. 10, 2(21): Rambla, M., Contribución al conocimiento de los Opiliones de la fauna ibérica. Estudio de los subórdenes Laniatores y Palpatores (pars). Resumen tesis doctoral. Secret. Publ. Univ. Barcelona: 20 pages. Roewer, C. F., Opiliones. Fünfte Serie, zugleich eine Revision aller bisher bekannten europäischen Laniatores. Arch. Zool. Exp. Gén., 78(1): Suzuki, S., The harvestmen of family Travuniidae from Japan. J. Sci. Hiroshima Univ. B1, 26:

6 Captions for illustrations: Figs 1-2. Travunia anophthalma (Absolon, 1916). Dorsal scute. 1. dorsal view; 2. lateral view (redrawn from Absolon & Kratochvíl, 1932c and Roewer, 1935). Figs 3-4. Examples of adult peltonychial claw of leg IV in lateral view. 3. Travunia anophthalma (Absolon, 1916) distitarsus and claw, (redrawn from Absolon & Kratochvíl, 1932). 4. Yuria pulchra Suzuki, 1964 (redrawn from Suzuki, 1975). Figs 5-9. Examples of adult peltonychial claw in dorsal view. 5. Leg III Abasola borisi Hadži, 1973 (possibly a synonym of Dinaria vjetrenicae). 6. Leg IV. Abasola borisi Hadži, 1973; 7. Dinaria vjetrenicae (Hadži, 1932); 8. Travunia anophthalma (Absolon, 1916); 9. Yuria pulchra Suzuki, Redrawn from several sources (Hadži, 1973; Roewer, 1935; Absolon & Kratochvíl, 1932; Suzuki, 1975). Fig 10. Abasola troglodytes (Roewer, 1915) Sternum, coxae and stigmatic area, ventral view (redrawn from Roewer, 1935). Fig 11. Peltonychia leprieuri (Lucas, 1860) right pedipalp, mesal view (from Martens, 1978). Figs Male genitalia. 12. Dinaria vjetrenicae (Hadži, 1932), penis; 13. Same, distal part ventral view; 14. Same, distal part lateral view; 15. Peltonychia postumicola (Roewer, 1935) (status of species uncertain, it may be a synonymy of P. tenuis or even P. clavigera), penis; 16. Same, distal part ventral view; 17. Same, distal part lateral view;, 18. Yuria pulchra Suzuki, 1964, distal part of penis lateral view;, 19. Same, dorsal view; 20. Same, ventral view (from Martens, 1976; 1978; 1986; Suzuki, 1975). Fig 21. Peltonychia clavigera (Simon, 1879), ovipositor (from Martens, 1978). Fig 22. Yuria pulchra, view of free abdominal sclerites (from Suzuki, 1975).

TROGLOBITIC HARVESTMEN RECENTLY DISCOVERED IN NORT H AMERICAN LAVA TUBES (TRAVUNIIDAE, EREBOMASTRIDAE, TRIAENONYCHIDAE : OPILIONES ) Thomas S.

Briggs, T. S. 1974. Troglobitic harvestmen recently discovered in North American lava tube s (Travuniidae, Erebomastridae, Triaenonychidae : Opiliones). J. ArachnoL 1 :205-214. TROGLOBITIC HARVESTMEN RECENTLY