AnimaliaOpilionesCryptomastridaeDerkarabetianShahanStarrettJamesTsurusakiNobuoUbickDarrellCastilloStephanieHedinMarshalA stable phylogenomic classification of Travunioidea (Arachnida, Opiliones, Laniatores) based on sequence capture of ultraconserved elementsZookeys2852018201876013610.3897/zookeys.760.24937 CRYPTOMASTRIDAE http://zoobank.org/A32A845F-36A7-426B-B3C1-F4E65085F356 Derkarabetian & Hedinfam. n.Type genus.

Cryptomaster Briggs, 1969

Type species.

Cryptomaster leviathan Briggs, 1969

Diagnosis.

The Cryptomastridae can be diagnosed from all other travunioids by the presence of a distal swelling on tibia II that bears enlarged setae (Figure 6A, C), a sexually dimorphic structure found only in males. Both genera are fairly distinctive. Cryptomaster is easily identified as the largest (>2.5 mm body length) laniatorean in the Pacific Northwest of North America (Figure 6D) and largest member of Travunioidea, although two size forms exist (Starrett et al. 2016). Speleomaster species are restricted to lava tubes showing extreme levels of troglomorphy with complete absence of eyes, extremely reduced pigmentation, and leg elongation (Figure 6B). Although unrelated, Speleomaster and Speleonychia are both highly troglomorphic lava tube dwellers in the Pacific Northwest, found in Idaho and Washington, respectively. Aside from their disjunct geographic distribution, Speleomaster can be differentiated from Speleonychia by the absence of a free ninth tergite and lateral sclerites, and by the presence of bifurcating tarsal claws of the hind legs (Speleonychia with a peltonychium). The cryptomastrid genera can be distinguished from the eastern North American Cladonychiidae (Erebomaster + Theromaster) by the spination of the pedipalpal tarsus, previously noted by Briggs (1969, 1974). Cryptomastrids possess five prominent spines on the lateral margins of the pedipalpal tarsus, three on the prolateral margin and two on the retrolateral margin. Erebomaster and Theromaster possess three pairs of prominent lateral spines (in some Theromaster, the two apical retrolateral spines are fused at the base). The Cryptomastridae are unique in intestinal morphology, possessing a combination of an elongate and triangular DI (similar to Briggsus and Isolachus), and shorter OD2 and OD3 (similar to the Paranonychidae) (Suppl. material 2: Figure 2).

Included genera and species.

Cryptomaster. Described by Briggs (1969) and originally included only Cryptomaster leviathan Briggs, 1969 from the Coastal Range of southwestern Oregon. A second species, Cryptomaster behemoth Starrett & Derkarabetian, 2016, was described from the west-central Cascade Range of Oregon (Starrett et al. 2016).

Speleomaster. Briggs (1974) described the genus and both known species, Speleomaster lexi Briggs, 1974 and Speleomaster pecki Briggs, 1974, from lava tubes of the Snake River Plain in southern Idaho.

Morphology of Cryptomastridae. A) male Cryptomaster, arrow denotes sexually dimorphic swelling diagnostic of Cryptomastridae; B) Speleomaster, habitus; C) Scanning electron micrograph of tibial swelling, from Starrett et al. (2016); D) Cryptomastrid distribution.

Figures

Data type: molecular data

Explanation note:

Figure 1. Outgroup relationships for the 70% dataset. Nodes are fully supported (e.g., 100 bootstrap, 1.0 posterior probability, etc.) unless otherwise indicated. RAxML tree manually rooted according to BEAST topology. Asterisk in ASTRAL tree indicates node not given support values by default.

Figure 2. Representative midgut morphology. Only the right half of the midgut is illustrated. Drawings for Triaenonychoidea (Synthetonychia), Grassatores (Discocyrtus), Cladonychiidae, and Paranonychidae redrawn from Dumitrescu (1975, 1976). Abbreviations: D1 = diverticulum 1; OD2 = opisthosomal diverticula 2; OD3 = opisthosomal diverticula 3.

Figure 3. Representative genital morphology and distribution of Peltonychia in Europe. Records from Slovenia and Trieste, Italy (red) are in error. Genitalia from left to right: P. clavigera, redrawn from Thaler (1996); P. leprieurii, redrawn from Chemini (1985); P. gabria, redrawn from Martens (1978).

Figure 4. SEM penis morphology of Yuria and Paranonychidae. Scale bars 500 mm.

This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.Shahan Derkarabetian, James Starrett, Nobuo Tsurusaki, Darrell Ubick, Stephanie Castillo Marshal Hedin

Geographic distribution of travunioid genera. Colors correspond to classification proposed in this study. Abbreviations: wNA = western North America, eNA = eastern North America, EUR = central and southern Europe, JPN/KOR = Japan and South Korea.

StarrettJDerkarabetianSRichartCHCabreroAHedinM (2016) A new monster from southwest Oregon forests: Cryptomaster behemoth sp. n. (Opiliones, Laniatores, Travunioidea). ZooKeys 555: 1135. https://doi.org/10.3897/zookeys.555.6274 BriggsTS (1969) A new holarctic family of laniatorid phalangids (Opiliones). Pan-Pacific Entomologist 45: 35. BriggsTS (1974) Troglobitic harvestmen recently discovered in North American lava tubes (Travuniidae, Erebomastridae, Triaenonychidae: Opiliones). Journal of Arachnology 1: 205214. http://www.jstor.org/stable/3705196 DumitrescuDO (1975) Contribution a l’étude morphologique de l’appareil digestif (intestin moyen) des opilions. Proceedings of the 6th International Arachnology Congress, Amsterdam IV, April 1974. Vrije Universiteit at Amsterdam, 150–155. DumitrescuDO (1976) Recherches morphologiques sur l’appareil digestif (intestin moyen) des Gonyleptomorphi (Arachnida, Opilionida). Travaux du Muséum d’Histoire naturelle (Grigore Antipa) 17: 1730. ThalerK (1996) Neue Funde europäischer Krallenweberknechte. Berichte des naturwissenschaftlichen-medizinischen Verein Innsbruck 83: 135148. CheminiC (1985) Descrizione del maschio di Peltonychia leprieuri (Lucas) e ridescrizione di Mitostoma orobicum (Caporiacco) (Arachnida, Opiliones). Bollettino della Societá Entomologica Italiana 117: 7275. MartensJ (1978) Spinnentiere, Arachnida: Weberknechte, Opiliones. VEB Gustav Fischer Verlag, 464 pp.