The Arachnid Order Solifugae



Phylogeny of the Solifugae
Keys to FamiliesFamily Ammotrechidae
Family Ceromidae
Family Daesiidae
Family Eremobatidae
Family Galeodidae
Family Gylippidae
Family Hexisopodidae
Family Karschiidae
Family Melanoblossidae
Family Mummuciidae
Family RhagodidaeFamily Solpugidae
Catalog of the Solifugae


Phylogeny and Taxonomy

Twelve families of solifugids, comprising 141 genera and 1095 described species, are currently recognized.  Five monotypic fossil genera have been described (Petrunkevitch 1913; Poinar and Santiago-Blay 1989; Lebrun 1996; Mursch and Steffan 1996; Selden, and Shear 1996; Selden and Dunlop 1998; Dunlop and Rössler 2003; Dunlop, Wunderlichand Poinar 2004).  No suborders or superfamilies have been proposed within Solifugae and phylogenetic relationships are unknown.  Early attempts to produce a suprageneric classification were presented in the mid-late 1800s (Koch 1842; Simon 1879; Kraepelin 1899), but the modern classification was instigated in the 1930s by C.F. Roewer (1932, 1933, 1934, 1941), who described many new genera and species in a series of monographs that remain the most comprehensive treatments on the order.  Roewer’s classification is devoid of phylogenetic content (Harvey 2002, 2004) and has been repeatedly criticized (Birula 1938; Panouse 1961; Muma 1951; Lawrence 1955; Muma 1976; Simonetta and Delle Cave 1968; Lawrence 1976; Wharton 1981; Gromov 2000).   Several of his genera and subfamilies have been rejected (Birula 1938; Turk 1960; Delle Cave and Simonetta 1971).  Fundamental concepts of monophyly and diagnosibility are absent from the classification, which relies on a small set of variable characters for separating subfamilies and genera.  Some families (e.g., Galeodidae, Rhagodidae, Solpugidae), containing an abundance of arbitrarily defined genera that provide an inconsistent reflection of cladistic diversity, are in particularly dire need of revision.  Numerous genera are monotypic and narrowly delineated due to excessive ‘splitting’, while other genera are large and broadly  inclusive and appear to consist of multiple lineages that have been ‘lumped’ together.  As many as 50% of the current solifuge genera may be paraphyletic or polyphyletic, as could several families (e.g., Daesiidae, Gylippidae, Melanoblossidae), although some (e.g., Hexisopodidae, Rhagodidae) seem to be clearly defined by obvious autapomorphies.  The only published phylogenetic work conducted on any group of solifuges was a study of relationships in a single species-group of Eremobates (Brookhart and Cushing 2004).  Excepting Muma’s revision of the North American Eremobatidae (Muma 1951, 1962), no solifuge families have been revised since Roewer (1932, 1933, 1934, 1941), whose classification has been largely discredited.  The only regional faunas in relatively ‘good condition’ occur in the New World, where Roewer had little impact, and later researchers (e.g., Muma 1951, 1962, 1963, 1967, 1970, 1971, 1974, 1976, 1982, 1986, 1987, 1989; Muma and Muma 1988; Brookhart and Muma 1987; Muma and Brookhart 1988; Maury 1970, 1976, 1977, 1979, 1980a, 1980b, 1981, 1982a, 1982b, 1983, 1984, 1985, 1986, 1987, 1992, 1998) formulated a more meaningful classification based on the synthesis of multiple characters (although even here, there are significant problems with the existing classification).  Few large monographic syntheses of solifuge genera have appeared in the last 25 years.  Recent workers have tended to publish short papers, and these have tended to merely add new species to existing groups without addressing phylogenetic relations within or between those groups.  The rigor of traditional taxonomy has barely advanced in studies fo the Solifugae.  Few specimens are examined, few illustrations presented, and modern techniques (phylogenetics, molecular systematics, SEM, digital imaging, GIS, computerized databasing) are seldom applied.  Workable diagnostic keys to genera and species are available for few regions of the world (North and South America, southern Africa)(Muma 1951, 1970, 1976; Lawrence 1955; Maury 1984; Muma and Muma 1988; Armas 1996; Punzo 1998).  The order Solifugae is in urgent need of a higher-level phylogenetic analysis and monographic, family-scale revision to begin to sort out the confusion.  The described species diversity of Solifugae, just slightly lower than that of Scorpiones, is almost certainly a gross underestimate of the world fauna.  The regions where solifuges are most diverse (e.g., southern Africa, southwestern U.S.A.) have been best documented by past solifuge specialists (e.g., R.F. Lawrence, M.H. Muma).  However, these regions have not by any means been thoroughly surveyed: new species and distribution records continue to be discovered, e.g., in the southwestern U.S.A. (Brookhart and Cushing, 2002, 2004, 2005).  Many solifuge species are cryptic, seasonal, habitat-specific, and difficult to collect without appropriate methods (Muma 1966, 1970, 1975a, 1975b, 1979. 1980a, 1980b; Griffin 1990; Punzo 1994, 1997), and most regions where solifuges occur have not been surveyed in appropriate seasons or using appropriate methods.  Many species are known from one or a few specimens (often only the types), usually collected serendipitously at a single locality; hence little is known about their distributions.  The world’s solifuge fauna will probably double when all appropriate habitats are thoroughly surveyed.  Species delimitation in many families is largely based on secondary sexual characters of adult males (flagellum, cheliceral dentition)(e.g. Kraepelin, K. 1899; Roewer 1932, 1933, 1934, 1941; Muma 1951; Lawrence 1955; Wharton 1981), rendering females and immatures difficult or impossible to identify below genus.  Extensive field collecting, new morphological character systems, and molecular approaches for associating different sexes and life stages are needed to advance solifuge systematics.

This section of the website presents the current classification of the order Solifugae, along with a key to its contained families.  You may access this information through the links on the left side of the page.

Literature Cited:

Armas, L.F. de. 1996. Sistemática del orden Solpugida en Centroamérica. Situación actual y perspectivas. Revista Nicaraguense de Entomologia 36: 29–36.

Birula, A.A. 1938. Arachnides, Ordo Solifuga. In Faune de l’USSR. Vol. 1(3): i–vii, 1–173. L’Académie des Sciences de l’URSS: Moscow, Leningrad. (in Russian)

Brookhart, J.O. & Cushing, P.E. 2002. New species of Eremobatidae (Arachnida, Solifugae) from North America. Journal of Arachnology 30: 84–97

Brookhart, J.O. & Cushing, P.E. 2004. The systematics of the Eremobates scaber species-group (Solifugae, Eremobatidae). Journal of Arachnology 32: 284–312.

Brookhart, J.O. & Cushing, P.E. 2005. Three new species of Solifugae from North America and a description of the female of Branchia brevis (Arachnida, Solifugae). Journal of Arachnology 33: 127–133.

Brookhart, J.O. & Muma, M.H. 1987. Arenotherus, a new genus of Eremobatidae (Solpugida) in the United States. Cherry Creek High School Print Shop: Englewood, Colorado.

Delle Cave, L. & Simonetta, A.M. 1971. A tentative revision of the Daesiidae (Arachnida, Solifugae) from Ethiopia and Somalia. Monitore Zoologico Italiano, n.s., Supplemento 4: 37–77.

Dunlop, J.A. & Rössler, R. 2003. An enigmatic, solifuge-like fossil arachnid from the Lower Carboniferous of Kamienna Gуra (Intra-Sudetic Basin), Poland. Paläontologische Zeitschrift, Stuttgart 77(2): 389–400.

Dunlop, J.A., Wunderlich, J. & Poinar Jr, G.O. 2004. The first fossil opilioacariform mite (Acari: Opilioacariformes) and the first Baltic amber camel spider (Solifugae). Transactions of the Royal Society of Edinburgh: Earth Sciences 94: 261–273.

Griffin, E. 1990. Seasonal activity, habitat selection and species richness of Solifugae (Arachnida) on the gravel plains of the central Namib Desert. Pp. 77–82. In Namib ecology: 25 years of Namib research (M.K. Seely, ed.). Transvaal Museum Monograph 7.

Gromov, A.V. 2000. Solpugids of the genus Eusimonia Kraepelin 1899 (Arachnida: Solifugae, Karschiidae) of Central Asia. Pp. 79–86. In Proceedings of the 18th European Colloquium of Arachnology (P. Gajdoš & S.Pekár, eds.). Ekológia, Bratislava 19, Supplement 3.

Harvey, M.S. 2002. The neglected cousins: what do we know about the smaller arachnid orders? The Journal of Arachnology 30: 357–372.

Harvey, M.S. 2003. Catalogue of the Smaller Arachnid Orders of the World: Amblypygi, Uropygi, Schizomida, Palpigradi, Ricinulei and Solifugae. CSIRO Publishing: Collingwood, Australia.

Koch, C.L. 1842. Systematische Uebersicht über die familie der Galeoden. Archiv für Naturgeschichte 8: 350–356.

Kraepelin, K. 1899. Zur Systematik der Solifugen. Mitteilungen aus dem Naturhistorischen Museum in Hamburg 16: 195–259, taf. I, II.

Lawrence, R.F. 1955. Solifugae, scorpions and Pedipalpi, with checklists and keys to South African families, genera and species. Pp. 152–262. In South African Animal Life. Results of the Lund Expedition in 1950–1951. Vol. 1. Almquist and Wiksell: Stockholm.

Lawrence, R.F. 1976. Foreword Pp. i–ii In Muma, M.H. 1976. A review of solpugid families with an annotated list of western hemisphere solpugids. Publication of the Office of Research, Western New Mexico University, Silver City 2(1): 1–33.

Lebrun, P. 1996. [Cratosolpuga wunderlichi, a first Aptian solifuge from Brazil]. Mineraux et Fossiles 246:34–35. (in French)

Maury, E.A. 1970. Sobre la presencia de Guacha fasciata Mello-Leitao 1924 en la Argentina (Solifugae, Ammotrechidae, Mummuciinae). Physis, Buenos Aires 29(79): 357–362.

Maury, E.A. 1976. Nuevos solífugos Ammotrechidae de la Argentina (Arachnida, Solifugae). Physis, Buenos Aires 35: 87–104.

Maury, E.A. 1977. Notas sobre sa sistemática y distribución geográfica de Procleobis patagonicus (Holmberg 1876) (Solifugae, Ammotrechidae, Saronominae). Physis, Buenos Aires 36: 283–293.

Maury, E.A. 1979. Primera cita del orden Solifugae para el Uruguay (Arachnida). Neotropica 25: 159–160.

Maury, E.A. 1980a. Dos nuevos Pseudocleobis de la Patagonia (Arachnida, Solifugae, Ammotrechidae). Physis, Buenos Aires 39: 41–43.

Maury, E.A. 1980b. Presencia de la familia Daesiidae en America del Sur con la descripcion de un nuevo genero (Solifugae). Journal of Arachnology 8: 59–67.

Maury, E.A. 1981. Un nuevo genero de Daesiidae de la Argentina (Arachnida, Solifugae). Comunicaciones del Museo Argentino de Ciencias Naturales Bernardino Rivadavia. Serie entomología 1(5): 75–82.

Maury, E.A. 1982a. Solifugos de Colombia y Venezuela (Solifugae, Ammotrechidae). Journal of Arachnology 10(2): 123–143.

Maury, E.A. 1982b. Nota sobre un solifugo altoandino Dasycleobis crinitus Mello-Leito 1940 (Solifugae, Ammotrechidae). Neotropica 28(80): 183–187.

Maury, E.A. 1983. Los Pseudocleobis del oeste árido Argentino (Arachnida, Solifugae, Ammotrechidae). Physis, Buenos Aires 41: 169–174.

Maury, E.A. 1984. Los familias de Solifugos americanos y su distribucion geografica (Arachnida, Solifugae). Physis, Buenos Aires 42(103): 73–80.

Maury, E.A. 1985. Nota sobre los géneros Namibesia y Syndaesia (Solifugae, Daesiidae). Aracnologia 4:1–6.

Maury, E.A. 1986. The American solifugid families and their geographical distribution (Arachnida, Solifugidae). P. 318 [abstract]. In Proceedings of the Ninth International Congress of Arachnology (W.G. Egerhard, Y.D. Lubin & B.C. Robinson, eds.), Panama 1983. [9th International Congress of Arachnology, Panama City (Panama), 1–8 August 1983].

Maury, E.A. 1987. Consideraciones sobre algunos solifugos de Chile (Solifugae: Ammotrechidae, Daesiidae). Revista de la Sociedad Entomológica Argentina 44: 419–432.

Maury, E.A. 1992. Lista de los ejemplares tipicos de "Arachnida" (Opiliones, Scorpiones y Solifugae) depositados en el Museo Argentino de Ciencias Naturales "Bernardino Rivadavia". Suplemento I. Aracnol., Supl., 6: 1–10.

Maury, E.A. 1998. Solifugae. Pp. 560–568. In Biodiversidad de artrópodos argentinos (J.J. Morrone & S. Coscarón, eds). Ediciones SUR: La Plata.

Muma, M.H. 1951. The arachnid order Solpugida in the United States. Bulletin of the American Museum of Natural History 97(2): 35–141.

Muma, M.H. 1962. The arachnid order Solpugida in the United States, Supplement 1. American Museum Novitates 2092: 1–44.

Muma, M.H. 1963. Solpugida of the Nevada Test Site. Brigham Young University Science Bulletin. Biological series 3(2): 1–15.

Muma, M.H. 1966. The life cycle of Eremobates durangonus (Arachnida: Solpugida). The Florida Entomologist 49: 233–242.

Muma, M.H. 1967. Scorpions, whip scorpions and wind scorpions of Florida. Arthropods of Florida and Neighboring Land Areas 4: 1–28.

Muma, M.H. 1970a. An improved can trap. Notes of the Arachnologists of the Southwest 1: 16–18.

Muma, M.H. 1970b. A synoptic review of North American, Central American, and West Indian Solpugida (Arthropoda, Arachnida). Arthropods of Florida and Neighboring Land Areas 5: 1–62.

Muma, M.H. 1971a. A new Ammotrechella Roewer (Solpugida: Ammotrechidae) from Jamaica. The Florida Entomologist 54(1): 97–99.

Muma, M.H. 1971b. The Solpugids (Arachnida: Solpugida) of Chile with descriptions of a new family, new genera, and new species. American Museum Novitates 2476: 1–23.

Muma, M.H. 1974. An annotated list of solpugids of New Mexico. New Mexico Academy of Sciences Bulletin 15(2): 13–16.

Muma, M.H. 1975a. Long term can trapping for population analyses of ground-surface, arid-land arachnids. The Florida Entomologist 58: 257–270.

Muma, M.H. 1975b. Two vernal ground-surface arachnid populations in Tularosa Basin, New Mexico. The Southwestern Naturalist 20: 55–67.

Muma, M.H. 1976. A review of solpugid families with an annotated list of western hemisphere solpugids. Publication of the Office of Research, Western New Mexico University, Silver City 2(1): 1–33.

Muma, M.H. 1979. Arid-grassland solpugid population variations in southwestern New Mexico. The Florida Entomologist 62(4): 320–328.

Muma, M.H. 1980a. Comparison of three methods for estimating solpugid (Arachnida) populations. Journal of Arachnology 8: 267–270.

Muma, M.H. 1980b. Comparison of ground-surface spider populations in pinyon-juniper and arid-grassland associations in southwestern New Mexico. The Florida Entomologist 63(1): 211–222.

Muma, M.H. 1980c. Solpugid (Arachnida) populations in a creosotebush vs. a mixed plant association. The Southwestern Naturalist 25(2): 129–136.

Muma, M.H. 1982. Solpugida. Pp. 102–104. In Synopsis and classification of living organisms (S.P. Parker, ed.). Vol. 2. McGraw-Hill: New York.

Muma, M.H. 1986. New species and records of Solpugida (Arachnida) from Mexico, Central America, and the West Indies. Novitates Arthropodae 2(3): 1–31.

Muma, M.H. 1987. New species and records of Solpugida (Arachnida) from Mexico, Central America and the West Indies. Pp. 1–24, and 4 plates. Southwest Offset: Silver City, New Mexico.

Muma, M. H., & Muma, K. E. 1988. The arachnid order Solpugida in the United States (Supplement 2, a biological review). Pp. 1–35 and 2 plates. Southwest Offset: Silver City, New Mexico.

Muma, M. H. 1989. New species and records of Solpugida (Arachnida) from the United States. 60 pp. Douglas Print Shop: Douglas, Arizona.

Muma, M.H. & Brookhart, J.O. 1988. The Eremobates palpisetulosus species-group (Solpugida: Eremobatidae) in the United States. Cherry Creek High School Print Shop: Englewood, Colorado.

Muma, M. H., & Muma, K. E. 1988. The arachnid order Solpugida in the United States (Supplement 2, a biological review). Pp. 1–35 and 2 plates. Southwest Offset: Silver City, New Mexico.

Muma, M.H. & Nazario, M.L. 1971. New solpugids (Arachnida, Solpugida) from Puerto Rico. The Journal of Agriculture of the University of Puerto Rico 55(4): 505–512.

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Panouse, J.B. 1961. Variation with the age of characters used for the systematics of solipugids. Pp. 258–262. In Verhandlungen der XI. Internationaler Kongress für Entomologie, Wien, 17. bis 25. August 1960. (H. Strouhal & M. Beier, eds). Vol. 1.

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Punzo, F. 1994. Intraspecific variation in response to temperature and moisture in Eremobates palpisetulosus Fichter (Solpugida, Eremobatidae) along an altitudinal gradient. Bulletin of the British Arachnological Society 9: 256–262.

Punzo, F. 1997. Dispersion, temporal patterns of activity, and the phenology of feeding and mating in Eremobates palpisetulosus Fichter (Solifugae, Eremobatidae). Bulletin of the British Arachnological Society 10: 303–307.

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