AnimaliaORDOFAMILIADerkarabetianShahanStarrettJamesTsurusakiNobuoUbickDarrellCastilloStephanieHedinMarshalA stable phylogenomic classification of Travunioidea (Arachnida, Opiliones, Laniatores) based on sequence capture of ultraconserved elementsZookeys2852018201876013610.3897/zookeys.760.24937 TRAVUNIIDAE Absolon & Kratochvíl, 1932Type genus.

Travunia Absolon, 1920.

Type species.

Travunia troglodytes (Roewer, 1915).

Diagnosis.

It is difficult to diagnose the Travuniidae as all taxa have yet to be examined for all relevant characters. For all species in which male genitalia have been examined, the glans is widened and flattened with lateral extensions, tooth-like in Trojanella and wing-like in Travunia and Dinaria. The Travuniidae as defined here are restricted to the European Dinaric Karst and are highly troglomorphic, completely blind with a highly reduced ocularium (Figure 1F). The penis of Travunia and Dinaria is undifferentiated, while that of cladonychiids shows a clear division between glans and shaft. Trojanella shows some similarities in glans structure to Holoscotolemon and Peltonychia (e.g., divided glans and shaft), but the penis musculature is restricted to the apical portion of the shaft and glans in Trojanella, while the musculature of the European cladonychiids are restricted to the basal portion of the shaft.

Included genera and species.

Travunia. The genus Travunia includes four described species that are all highly troglomorphic and restricted to caves in the southern Dinaric Karst region of Europe: T. borisi (Hadži, 1973) from Bosnia and Herzegovina, T. hofferi (Šilhavý, 1937) from Montenegro, T. jandai Kratochvíl, 1937 from Croatia, and T. troglodytes (Roewer, 1915) from Croatia and Bosnia and Herzegovina.

Trojanella (Figure 1F). This monotypic genus is represented by T. serbica Karaman, 2005, a highly troglomorphic species restricted to a single cave on Stara Planina Mountain in Serbia.

Dinaria. A monotypic genus represented by the highly troglomorphic species D. vjetrenicae (Hadži, 1932) known only from Vjetrenica Cave in southern Bosnia and Herzegovina.

Remarks.

It is not surprising that Trojanella is included in the most early-diverging travunioid lineage given Karaman’s (2005) statement that this species is a “unique and isolated phylogenetic line in the superfamily”. Karaman’s decision to leave Trojanella unplaced in Travunioidea was made to highlight, and is a consequence of, the commonly used morphological characters that have hindered a reliable taxonomy within this group. It is unclear how many species of Travunia actually exist. Novak (2004) questioned the validity of Travunia, Dinaria, and Abasola at the generic level, and Abasola was later synonymized with Travunia (Kury and Mendes 2007). Novak (2004, 2005) argues that Travunia may be oversplit and includes only 2–3 species of questionable status. The status of Travunia and Dinaria as distinct genera has been questioned based on similarity in male genitalic morphology (Novak 2004); together these might represent a single lineage.

Photographs of live travunioid harvestmen. A Theromaster brunneus B Erebomaster sp. C Cryptomaster leviathan D Holoscotolemon lessiniense E Peltonychia leprieurii F Trojanella serbica G Briggsus sp. H Isolachus spinosus I Speleonychia sengeri J Yuria pulcra K Paranonychus brunneus L Sclerobunus nondimorphicus M Metanippononychus sp. N Zuma acuta O Kainonychus akamai. All photos by MH, except D, E (courtesy of and copyright A. Schönhofer), and F (courtesy of and copyright I. Karaman).

KaramanIM (2005) Trojanella serbica gen. n., sp. n., a remarkable new troglobitic travunioid (Opiliones, Laniatores, Travunioidea). Revue Suisse de Zoologie 112: 439456.https://doi.org/10.5962/bhl.part.80308 NovakT (2004) An overview of harvestmen (Arachnida: Opiliones) in Croatia. Natura Croatica 13: 231296. https://hrcak.srce.hr/10554 KuryABMendesAC (2007) Taxonomic status of the European genera of Travuniidae (Arachnida, Opiliones, Laniatores). Munis Entomology & Zoology 2: 114. NovakT (2005) An overview of harvestmen (Arachnida: Opiliones) in Bosnia and Herzegovina. Natura Croatica 14: 301350. https://hrcak.srce.hr/1726