Family Triaenonychidae version 1.0

Transcription

1 1 Family Triaenonychidae version 1.0 Adriano B. Kury Departamento de Invertebrados, Museu Nacional/UFRJ Quinta da Boa Vista, São Cristóvão, , Rio de Janeiro - RJ BRAZIL 1. Introduction The Triaenonychidae is a large family of a little less than 500 described species of small to medium-sized Laniatores distributed in the Northern and Southern Temperate Regions of the World. Recognizable by the posterior claws which are single with one or more pairs of lateral branches, exceptionally with more complex ramifications. Genitalia is complex, with many sclerites, all external. Definition of the family uses only symplesiomorphies, so it is not a surprise not to recover a monophyletic Triaenonychidae from phylogenetic analyses. They are the dominant element of the opiliofauna in Madagascar, South Africa and New Zealand. In Chile they share the importance with the Gonyleptidae. The biggest Triaenonychidae occur in Madagascar (Triaenonychinae) and South Africa (Adaeinae). There is a lot of undescribed genera and species in Chile, and many generic reviews are due, since different authors working with fauna of different continents did not establish a standard for descriptions and diagnostic characters. Intercontinental relationships are very interesting and little studied, telling part of the story of Temperate Gondwana (Kury 2004) Subtaxa included. The 121 genera and 478 species are organized in up to 6 subfamilies, although there is no consensus among the authors. Five subfamilies are small while the nominal subfamily concentrates most of diversity of the family (108 genera and 435 species) and is divided in three tribes. Four of the subfamilies are actually closest to the Travuniidae and Pentanychidae (Kury, 2004) and should not properly be included here, while the New Zealand Synthetonychiidae should be grouped inside Triaenonychidae. Two alternative classifications are shown below.

2 2 Compromise Conservative Classification Triaenonychidae Sørensen, 1886 Kaolinonychinae Suzuki, 1976 Nippononychinae Suzuki, 1976 Paranonychinae Briggs, 1971 Sclerobuninae Dumitrescu, 1976 Soerensenellinae Forster, 1954 Triaenonychinae Sørensen, 1886 Adaeini Pocock, 1902 Triaenobunini Pocock, 1902 Triaenonychini Sørensen, 1886 Bold Alternative Classification Triaenonychidae Sørensen, 1886 Adaeinae Pocock, 1902 Synthetonychiinae Forster, 1954 Soerensenellinae Forster, 1954 Triaenonychinae Sørensen, 1886 Triaenobuninae Pocock, 1902 (None of the others belongs to this family) 1.2. Systematic historical background Before the family was recognized, only a few species were described Packard (1877) described Scotolemon robustus from Colorado, USA, while Simon (1880) described the genus Equitius for Equitius doriae from Australia. Karsch (1880) described the genus Adaeum for Adaeum asperatum from South Africa considering it to be an intermediate between the families Cosmetidae and Gonyleptidae (!). Sørensen (1886) created the family Triaenonychoidae, with two more genera and two species genus Triaenonyx for Triaenonyx rapax Sørensen, 1886 from Polynesia, but actually from Chile and genus Triaenobunus for Triaenobunus bicarinatus Sørensen, Slowly other species started to be discovered. Banks (1893) described the genus Sclerobunus for Packard s species and a new one from Western USA and Loman (1898) the genera Acumontia for Acumontia armata from Madagascar, Larifuga from South Africa and two more species of Adaeum. 20th century brought the first great change, when Pocock (1902) split the young family in three different ones Adaeidae, Triaenobunidae and Triaenonychidae. At the same time Sørensen (1902) was describing more Chilean species, Pocock added some species from South Africa, Madagascar and Australia/New Zealand (1902; 1903) and Loman (1902) from New Zealand. Loman created without much explanation the new suborder Insidiatores for Triaenonychidae (only later including the subdivisions by Pocock), and in view of the heavy critics of Pocock, explained properly his views shortly later

3 3 (Loman, 1903). Hogg (1910) described a few new Zealand species. Roewer (1915) did the first monograph on the family, with an update 16 years later (Roewer 1931). The excellent papers by Lawrence (e. g. 1931; 1938a; 1963) greatly enhanced the knowledge on South African Fauna. Forster, in a review of Laniatores from New Zealand (Forster 1954), described a lot of new species and genera and created a much influential new system. Hickman (1958) created a great number of new genera and species for Tasmania. Briggs (1967; 1971) reviewed most USA species and proposed a new subfamily. Suzuki (1975; 1976) followed Briggs paradigm and proposed yet two new subfamilies of Japanese/Korean Triaenonychidae. Dumitrescu (1976) based on studies of the midgut diverticules proposed another subfamily, ignored by all other authors. Maury (e. g. Maury & Roig, 1985) started a fine series of papers devoted to the review of South American Triaenonychidae, containing high-quality illustrations of genitalia and including a significant discussion on the systematics of the superfamily Travunioidea by occasion of the description of a strange Argentinean troglomorphic species (Maury, 1988). This series was interrupted by his untimely death. Hunt (e. g. Hunt, 1985; Hunt & Hickman, 1993) reviewed some Australian taxa and made objections to current generic cuts. Unfortunately this author also passed away prematurely Natural history A review of cavernicolous members of the family is given by Hunt (1972). Lawrence (1938b) studied devices of spreading repugnatory fluid. Austral Triaenonychidae are typically found under rotten logs and in leaf litter. Some species possess a cover of earth particles adhered to the complexly ornamented tegument. Forster (1954) separated a whole subfamily, Soerensenellinae based on maternal care of the eggs, contrasting with the other(s) in which such behavior was never observed: Eggs laid in a single group under a log or stone and guarded by female until hatched. Some South African species possess a stridulatory grate on the mesal surface of cheliceral hand, but actual stridulation has never been reported. 2. Characterization SIZE. Medium-sized Laniatores, body length typically 3 to 5 mm, although some South African Triaenonychinae can be much smaller (down to 1.5 mm) and on the other side some Adaeinae are much larger (up to 9 mm). Legs I-IV almost always short 4-7/6-12/4-8/6-10 mm long. DORSUM. Dorsal scutum width increasing backwards without major constriction. Mesotergum seldom clearly divided into areas by grooves, usually areas are marked by arrangement of tubercle rows. No areas fused. Armature of areas and tergites

4 4 usually weak, formed by small paired acuminate spiniform tubercles. Common eye mound usually present, mostly very narrow and high with unpaired armature. Sometimes common eye mound lacking and eyes are sessile placed close together. Eyes elevated, much higher than the level of carapace. Anterior margin of carapace with rows of spines. Ozopores hidden by a pad-shaped apophysis of coxa II. VENTER. Coxae more or less parallel, coxa IV not greatly enlarged. Stigmata sometimes concealed by tubercles. Shape of sternum much variable. CHELICERA. Cheliceral hands usually not swollen, and basichelicerite rarely with dorsal ornamentation of tubercles, but mesal rows of pointed tubercles are common. PEDIPALP. Pedipalps large, much stronger than legs, not crossed in the region of trochanter. Armed with ventro-mesal and ventro-ectal spines in patella-tibia-tarsus. Femur variedly armed with apophyses and spines, ventrally in many genera with strip of fine bead-like granules. Tibia and tarsus do not form a subchela. LEGS. Legs usually short, substraight and armed only with tubercle rows. Large elaborate apophyses never present. Coxa I ventrally with strong frontal apophyses. Femur I in many species armed with ventral and/or dorsal rows of setiferous spiniform processes. Metatarsus I may be notched in males and provided with strong setae. Tarsi I-II with a single claw, III-IV with a multifurcate (usually trifurcate) claw. Distitarsus I undivided or 2 jointed. Distitarsus II typically 2 jointed, may reach 4 joints. Tarsal counts typically 2-3/2-8 (reaching more than 20)/3-4/3-4. COLOR. Color background usually orange-brown to dark-brown, with black mottling and reticulation. No white markings on dorsal scutum. GENITALIA. Truncus penis filled with a single muscle. Glans very complex, with a full complement of sclerites, rarely found together in a same species. Sometimes stylus is extremely elongate and plates and spines may be fused with it. MALE DIMORPHISM (POECILANDRY). Reported by Forster (1954) and Hunt (1985), some males have secondary dimorphic features only weakly developed, though genitalia are normal. SEXUAL DIMORPHISM. Sexual dimorphism is manifested typically in pedipalps, which in male are much stronger and incrassate. In a few species, chelicerae of male have supplementary spatulate mesal apophyses. The number of tarsal counts of leg I in female may be a little lower. In many species males possess a notch in metatarsus I. 3. Distribution USA, Japan, Korea, Australia, New Zealand, Madagascar, Chile, Argentina and southern Brazil. It is interesting to note that some genera of Triaenonychidae are distributed across the Austral continents, not to mention the possibly non-triaenonychidae of the Boreal temperate that also cross continents (Paranonychinae, see Shear, 1986).

5 5 4. Relationships In the chapter of evolution in this book and elsewhere (Kury, 2004), reasons are given to split the Triaenonychidae as traditionally conceived in at least two different families. The Boreal genera should be grouped with the Travuniidae while the Austral genera represent the Triaenonychidae sensu stricto and may include the strange Synthetonychiidae. 5. References Banks, Nathan, The Phalangida Mecostethi of the United States. Trans. American Entomol. Soc., 20(2): Briggs, Thomas S., An emendation for Zuma acuta Goodnight & Goodnight (Opiliones). Pan-Pacific Entomologist, 43: 89. Briggs, Thomas S., The harvestmen of family Triaenonychidae in North America (Opiliones). Occ. Pap. Calif. Acad. Sci., 90: Dumitrescu, Dan, Recherches morphologiques sur l appareil digestif (intestin moyen) des Gonyleptomorphi (Arachnida, Opilionida). Trav. Mus. Hist. Nat. Gr. Antipa, 17: Forster, Raymond R The New Zealand harvestmen (sub-order Laniatores). Canterbury Museum bulletin, Christchurch, 2: Hickman, V. V Some Tasmanian harvestmen of the family Triaenonychidae (suborder Laniatores). Papers and proceedings of the Royal Society of Tasmania, Hobart Town, 92: Hogg, Henry R Some New Zealand and Tasmanian Arachnidae. Transactions of the New Zealand Institute, Wellington, 42(33): Hunt, Glenn S A new cavernicolous harvestman from Western Australia (Arachnida: Opiliones: Triaenonychidae). Journal of the Australian Entomological Society, Brisbane, 11(3): Hunt, Glenn S Taxonomy and distribution of Equitius in Eastern Australia (Opiliones, Laniatores, Triaenonychidae). Records of the Australian Museum, Sydney, 36: Hunt, Glenn S., Revision of the genus Holonuncia Forster (Arachnida: Opiliones: Triaenonychidae) with description of cavernicolous and epigean species from Eastern Australia. Records of the Australian Museum, Sydney, 44:

6 6 Hunt, Glenn S. & John L. Hickman A revision of the genus Lomanella Pocock and its implications for family level classification in the Travunioidea (Arachnida: Opiliones: Triaenonychidae). Records of the Australian Museum, Sydney, 45: Karsch, Ferdinand IX. Neue Phalangiden des Berliner Museums, pp In Arachnologische Blätter (Decas I). Tafel 12. Zeistschrift fur die gesamten Naturwissenschaften, Berlin, 53(6): Kury, Adriano B., Intercontinental relationships of Southern Hemisphere Triaenonychidae. Cimbebasia, Windhoek [ in review]. Lawrence, Reginald F The harvest-spiders (Opiliones) of South Africa. Annals of the South African Museum, Cape Town, 29: Lawrence, R. F., New harvest-spiders from Natal and Zululand. Annals of the Natal Museum, Pietermaritzburg, 8(2): , 11 fig. Lawrence, Reginald F. 1938a. Harvest-spiders of Natal and Zululand. Annals of the Natal Museum, Pietermaritzburg, 8(3): Lawrence, Reginald F. 1938b. The odoriferous glands of some south african harvestspiders. Transactions of the Royal Society of South Africa, Cape Town, 25(4): Lawrence, Reginald F The Opiliones of the Transvaal. Annals of the Transvaal Museum, Pretoria, 24: Loman, Jan Cornelis Christiaan Beiträge zur Kenntniss der Fauna von Süd-Afrika. Ergebnisse einer Reise von Prof. Max Weber im Jahre IV. Neue Opilioniden von Süd-Afrika und Madagaskar. Zoologische Jahrbücher, Jena, Abteilung für Systematik, Geographie und Biologie der Tiere, 11: Loman, Jan Cornelis Christiaan Neue aussereuropaische Opilioniden. Zoologische Jahrbücher, Jena, Abteilung für Systematik, Ökologie und Geographie der Tiere, 16: , pr. 9. Loman, Jan Cornelis Christiaan On the classification of Opiliones. Tijdschrift der Nederlandsche dierkundige vereeniging, Rotterdam, 8: Maury, Emilio A., Triaenonychidae sudamericanos V. Un nuevo genero de opiliones cavernicolas de la Patagonia (Opiliones, Laniatores. Mémoires de Biospéologie, 15: Maury, Emilio A., Triaenonychidae sudamericanos. VII. Redescripcion de Araucanobunus juberthiei Muñoz Cuevas 1973 (Opiliones, Laniatores). Boletin sociedad biologica Concepción, 64:

7 7 Maury, Emilio A. & Arturo H. Roig A., Triaenonychidae sudamericanos I. El género Ceratomontia Roewer, 1915 (Opiliones: Laniatores). Historia Natural, 5(11): Packard Jr., Alpheus S., On a new cave fauna in Utah. Bull. U.S.A. Geol. Geogr. Surv. Terr., 3(1): Pocock, Reginald Innes. 1902a. Some points in the morphology and classification of the Opiliones. Annals and Magazine of Natural History: Zoology, Botany and Geology, London, 7th Series, 10: Pocock, Reginald Innes. 1902b, On some new harvest-spiders of the order Opiliones from the Southern Continents. Proceedings of the Zoological Society of London, London, 2 (6): Pocock, Reginald Innes. 1903a. Fifteen new species and two new genera of tropical southern Opiliones. Annals and Magazine of Natural History: Zoology, Botany and Geology, London, 7th Series, 11: , 2 pl. Roewer, Carl-Friedrich. 1915a. Die Familie Triaenonychidae der Opiliones-Laniatores. Archiv für Naturgeschichte, Berlin, Abt. A, Original-Arbeiten, 80(12): Roewer, Carl-Friedrich. 1931b. Über Triaenonychiden (6. Ergänzung der Weberknechte der Erde 1923). Zeitschrift für wissenschaftliche Zoologie, Leipzig, 138(1): Shear, William A., A cladistic analysis of the opilionid superfamily Ischyropsalidoidea, with descriptions of the new family Ceratolasmatidae, the new genus Acuclavella, and four new species. American Museum Novitates, New York City, 2844: Simon, Eugène. 1880a. Premier supplément au travail intitulé essai d une classification des Opiliones Mecostethi, etc (première Partie). Comptes rendus des séances de la Société Entomologique de Belgique, Bruxelles, 1880: Sørensen, William E. 1886a. Opiliones. In Ludwig Koch & E. von Keyserling, Die Arachniden Australiens nach der Natur beschrieben und abgebildet, 2(33): 53-86, pl Nürnberg: Bauer & Raspe. Sørensen, William E Gonyleptiden (Opiliones Laniatores). Ergebnisse der Hamburger Magalhaensischen Sammelreise, 5: Suzuki, Seisho The harvestmen of family Triaenonychidae in Japan and Korea. Journal of Sciences of the Hiroshima University, series B1 (Zoology), 26: Suzuki, Seisho. 1976c. Two triaenonychid harvestmen from the Northeast Japan. Journal of Sciences of the Hiroshima University, series B1 (Zoology), 26:

8 a b c d e 6 a b c Triaenonychidae. Figs Holonuncia katoomba Hunt, 1992 from Australia, habitus, lateral view (from Hunt, 1992); 2-3. South African Adaeinae, schematic granulation of Adaeulum (left) and Larifuga (from Lawrence, 1933); 4-5. Araucanobunus juberthiei Muñoz-Cuevas, 1973 from Chile, coxae, sternum and genital opercle of male and female (from Maury, 1993); 6. Typification of outlines of sternum of the three tribes of Triaenonychinae, from left, Triaenonychini, Adaeini, Triaenobunini (redrawn from Forster, 1954); 7. Further elaboration on the sternum outline of genera of South African Adaeini, from left, Montadaeum, Larifuga, Paradaeum, Adaeulum, Cryptadaeum (redrawn from Lawrence, 1931); 8-9. Gen. sp. of Triaenobuninae from Chile showing complex ornamentation (photos A.B.Kury).

9 Triaenonychidae. Figs Araucanobunus juberthiei Muñoz-Cuevas, 1973 from Chile, penis ventral and lateral views (from Maury, 1993); 11. Ceratomontia mendocina Maury & Roig, 1985, from Argentina, penis lateral and ventral (from Maury & Roig, 1985); Graemontia natalensis Lawrence, 1937 from South Africa chelicera mesal view, pedipalpus mesal view and leg I, mesal view (from Lawrence, 1937); 15. Ankaratrix illota Lawrence, 1959 from Madagascar, eye mound, lateral view (from Lawrence, 1959); 16. Pristobunus henopoeus Forster, 1954 from New Zealand, habitus, dorsal view (from Forster, 1954); 17. Gen. sp. from Madagascar (USNM) ozopore and pad of coxa II (photo A.B.Kury); 18. Holonuncia cavernicola Forster, 1955 from Australia, metatarsal notch (from Hunt, 1992); 19. Lomanella spp. posterior claws (from Hunt & Hickman, 1993); 20. Gen. sp. from Madagascar (USNM) typical 3-pronged claw (photo A.B.Kury).