Orussidae

The Orussidae or the parasitic wood wasps represent a small family of sawflies (" Symphyta"). Currently, about 93 extant and four fossil species are known.Blank, S.M., Groll, E.K., Liston, A.D., Prous, M. & Taeger, A. 2012: ECatSym - Electronic World Catalog of Symphyta (Insecta, Hymenoptera). Program version 4.0 beta, data version 39 (18. Dezember 2012). Digital Entomological Information, Müncheberg
/ref> They take a key position in phylogenetic analyses of Hymenoptera, because they form the sister taxon of the megadiverse apocritan wasps, and the common ancestor of Orussidae + Apocrita evolved parasitism for the first time in course of the evolution of the Hymenoptera. They are also the only sawflies with carnivorous larvae.

Description

Adults

The fully winged wasps are 2−23 mm long. They are predominantly black but species of ''Chalinus'', ''Mocsarya'' and ''Orussobaius'' are more or less metallic. Some species have a red thorax or abdomen and conspicuous white or golden pilosity. Many ''Orussus'' species bear white spots on the legs. The antennae of males are composed of 11, those of females of 10 articles. The modified distal antennal articles of females (article 9 enlarged, article 10 very small) are involved in vibrational sounding to detect suitable oviposition sites and the host larvae living concealed inside wood.Vilhelmsen, L., Isidoro, N., Romani, R., Basibuyuk, H.H. & Quicke, D.L.J. 2001: Host location and oviposition in a basal group of parasitic wasps: the subgenual organ, ovipositor apparatus and associated structures in the Orussidae (Hymenoptera, Insecta). Zoomorphology 121: 63-84. Contrary to other "Symphyta", the antennae insert near the lower edge of the compound eyes and close to the mandible. The mandibles are orthognathous and lack evident teeth. The number of palpomeres of the maxilla and the labium varies and is used as a taxonomic character. On the wings, some cross-veins are reduced in comparison with the more complete venation of other basal Hymenoptera. Similar as in most other sawflies, the wings are held at rest with a device called " cenchri". The complete body is strongly sclerotized and bears a species-specific microstructure, which is relevant for species identification. The ovipositor is several times as long as the body, and at rest it extends inside the body from the abdomen to the prothorax, where it is coiled, and back towards the tip of the abdomen again.

Orussidae and Stephanidae are the sole Hymenoptera in which the head bears a corona of erect teeth around the frontal ocelli. Contrary to Orussidae, the Stephanidae lack cenchri, and their mesosoma and metasoma are separated by a wasp waist, which is absent in Orussidae.

Larvae

Similar to the larvae of apocritan Hymenoptera, larvae of Orussidae have reduced some morphological features as a result of their parasitic life style inside the tunnels of wood-boring insects. They are white, subcylindrical, weakly sclerotized with a distinct head capsule. The mouthparts are hypognathous. Eyes and legs are reduced or completely absent. The mandibles are well developed and strongly sclerotized. Palps of maxilla and labium lack. The surface of the body is subdivided into distinct segments, each bearing a transverse row of 8−10 backward pointing spines.Rohwer, S.A. & Cushman, R.A. 1917: Idiogastra, a new suborder of Hymenoptera with notes on the immature stages of ''Oryssus''. Proceedings of the entomological Society of Washington 19(1-4): 89-98.

Biology and behavior

Orussid wasps have been collected only rarely. Over the years, their abundance has been subject to strong fluctuations.Burger, F. & Taeger, A. 1994: Aktuelle Nachweise von ''Orussus abietinus'' (Scopoli, 1763) (Hymenoptera, Orussidae). Brandenburgische Entomologische Nachrichten 2(1): 61-62. The species are thermophilous and wasps are active during the hottest hours of the day. Therefore, they are rarely found by the entomologists specialized in sawflies.

Only for few species the larval biology is known. Orussidae are parasitoids of xylobiontic larvae of beetles or Hymenoptera, particularly of the larvae of jewel beetles ( Buprestidae), long-horned beetles (Cerambycidae), and wood wasps ( Siricidae, Xiphydriidae).Rawlings, G.B. 1957: ''Guiglia schauinslandi'' (Ashmead) (Hym. Orussidae) a parasite of ''Sirex noctilio'' (Fabricius) in New Zealand. Entomologist 90(1125): 35-36.Kraus, M. 1998: Die Orussidae Europas und des Nahen Ostens (Hymenoptera: Orussidae). Pp. 283-300. In: Taeger, A. & Blank, S. M. (eds): Pflanzenwespen Deutschlands (Hymenoptera, Symphyta). Kommentierte Bestandsaufnahme. Goecke & Evers, Keltern. Orussids can be observed running around quickly on dead tree trunks. The females locate the host larvae living concealed inside wood by generating vibrations by tapping the tips of their antennae against the wood surface. The vibrations are picked up by the modified fore legs.Vilhelmsen, L. & Turrisi, G.F. 2011: Per arborem ad astra: Morphological adaptations to exploiting the woody habitat in the early evolution of Hymenoptera. Arthropod Structure & Development 40: 2-20. After locating the host, the female drills into the wood with the very long ovipositor and lays the egg. The egg is very elongate with a small expansion on the anterior end and a long expansion on the posterior end. The egg is coiled on the host. In some species the egg is possibly laid into the tunnel of the host if the host itself can not be reached, and the orussid larva itself crawls to the host. On its way to the host it may gnaw through the wood shavings left by the host. It has been disputed whether this material contributes to the nutrition of the larva. At least in ''Guiglia schauinslandi'', the larva lives externally for its first two instars and then enters the putrid(?) liquid in the dead host, where it remains until the adult emerges.

Systematics

Numerous morphological and genetic studies indicate that the Orussidae form the sister taxon of the Apocrita, the wasps, bees, and ants.Vilhelmsen, L. 2001: Phylogeny and classification of the extant basal lineages of the Hymenoptera (Insecta). Zoological Journal of the Linnean Society 131: 393-442.Sharkey, M.J., Carpenter, J.M., Vilhelmsen, L., Heraty, J., Liljeblad, J., Dowling, A.P.G., Schulmeister, S., Murray, D., Deans, A.R., Ronquist, F., Krogmann, L., Wheeler, W.C. 2012: Phylogenetic relationships among superfamilies of Hymenoptera. Cladistics 28: 80-112. Accordingly, parasitic life style has not evolved first in Apocrita but in the common ancestor of Orussidae + Apocrita.

Earlier, the Orussidae were sometimes put into a separate suborder, Idiogastra, but today they are classified in their own superfamily, Orussoidea.Taeger, A., Blank, S.M. & Liston, A.D. 2010: World Catalog of Symphyta (Hymenoptera). Zootaxa 2580: 1-1064
Abstract
/ref> Orussidae are demonstrably monophyletic. Tribes and subfamilies within the Orussidae have been abandoned, since such earlier subdivisions could not be corroborated in phylogenetic analyses.

The cladogram is based on Schulmeister 2003. The Orussoidea are sister to the Apocrita.

Orussidae genera and species of the World

An identification key for the genera of the World was published by Vilhemsen (2003).Vilhelmsen, L. 2003: Phylogeny and classification of the Orussidae (Insecta: Hymenoptera), a basal parasitic wasp taxon. Zoological Journal of the Linnean Society 139: 337-418. The following list summarizes the genera and species and their gross distribution together with indications on identification keys:

* '' Argentophrynopus'' Vilhelmsen & D.R. Smith, 2002: 2 species, Costa Rica and supposedly Mexico. Taxonomy: Vilhelmsen & Smith (2002).Vilhelmsen, L. & Smith, D.R. 2002: Revision of the 'ophrynopine' genera ''Argentophrynopus'' gen. n., ''Guiglia'' Benson, ''Kulcania'' Benson, ''Ophrella'' Middlekauff, ''Ophrynon'' Middlekauff, ''Ophrynopus'' Konow, and ''Stirocorsia'' Konow (Hymenoptera: Orussidae). Insect Systematics & Evolution 33(4): 387-420.
* †'' Baltorussus'' Schedl, 2011: 1 fossil species, '' Baltorussus velteni'' Schedl, 2011 from Baltic amber. Taxonomy: Schedl (2011).Schedl, W. 2011: Eine Orussidae aus dem baltischen Bernstein (Hymenoptera: Symphyta). Zeitschrift der Arbeitsgemeinschaft Österreichischer Entomologen 63: 33-36.
* '' Chalinus'' Konow, 1897: 9 Afrotropic species. Taxonomy: Vilhelmsen (2001),Vilhelmsen, L. 2001: Systematic revision of the genera ''Chalinus'' Konow, 1897 and ''Mocsarya'' Konow, 1897 (Hymenoptera: Orussidae). Insect Systematics & Evolution 32: 361-380 Vilhelmsen (2005).Vilhelmsen, L. 2005: ''Chalinus albitibialis'', a new species of Orussidae (Insecta, Hymenoptera) from Morocco. Zootaxa 880: 1-7
* ''Guiglia'' Benson, 1938: 7 Australian species, plus '' Guiglia chiliensis'' Benson, 1955 in Chile. Taxonomy: Vilhelmsen & Smith (2002).
* '' Kulcania'' Benson, 1935: 2 species in Colombia, Costa Rica, Mexico and the USA. Taxonomy: Vilhelmsen & Smith (2002).
* '' Leptorussus'' Benson, 1955: 2 Afrotropic species. Taxonomy: Vilhelmsen (2003), Vilhelmsen (2007).Vilhelmsen, L. 2007: The Phylogeny of Orussidae (Insecta: Hymenoptera) Revisited. Arthropod Systematics & Phylogeny 65: 111-118.
* †'' Mesorussus'' Rasnitsyn, 1977: 1 fossil species, '' Mesorussus taimyrensis'' Rasnitsyn, 1977. Taxonomy: Rasnitsyn (1977).Rasnitsyn, A.P. 1977: Novye pereponchatokrylye iz jury i mela Azii. ew Hymenoptera from the Jurassic and Cretaceous of Asia.(In Russian). Paleontologicheskij Zhurnal 9773): 98-108.
* †'' Minyorussus'' Basibuyuk, Quicke & Rasnitsyn, 2000: 1 fossil species, '' Minyorussus luzzii'' Basibuyuk, Quicke & Rasnitsyn, 2000. Taxonomy: Basibuyuk et al. (2000).Basibuyuk, H.H., Quicke, D.L.J. & Rasnitsyn, A.P. 2000: A new genus of the Orussidae (Insecta: Hymenoptera) from Late Cretaceous New Jersey amber. Pp. 305-311. In: Grimaldi, D. (ed.): Studies on Fossils in Amber, with Particular Reference to the Cretaceous of New Jersey. Leiden: Backhuys Publ.
* '' Mocsarya'' Konow, 1897: 2 species, '' Mocsarya metallica'' (Mocsáry, 1896) in Indonesia and Sri Lanka, '' Mocsarya syriaca'' Benson, 1936 in Greece and Turkey (not in Syria!). Taxonomy: Vilhelmsen (2001).,
* '' Ophrella'' Middlekauff, 1985: 3 species, ''O. amazonica'' (Westwood, 1874) in French Guiana, Brazil and Panama, ''O. eldorado'' Vilhelmsen, 2013 and ''O. seagi'' Vilhelmsen 2016 in French Guiana. Taxonomy: Vilhelmsen & Smith (2002), Vilhelmsen et al. (2013),Vilhelmsen, L., Blank, S.M., Costa, V.A., Alvarenga, T.M. & Smith, D.R. 2013: Phylogeny of the ophrynopine clade revisited: review of the parasitoid sawfly genera ''Ophrella'' Middlekauff, ''Ophrynopus'' Konow and ''Stirocorsia'' Konow (Hymenoptera: Orussidae). Invertebrate Systematics 27(4): 450-483
Abstract/PDF
/ref> Vilhelmsen (2016).Vilhelmsen, L. (2016) A new species of ''Ophrella'' Middlekauff, 1985 (Hymenoptera, Orussidae) from French Guiana. Journal of Hymenoptera Research 51: 171-185
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/ref>
* '' Ophrynon'' Middlekauff, 1983: 4 species in California (USA). Taxonomy: Vilhelmsen & Smith (2002), Blank et al. (2010).Blank, S.M. & Vilhelmsen, L. & Smith, D. R. 2010: ''Ophrynon'' (Hymenoptera: Orussidae) in California: diversity, distribution and phylogeny. Insect Systematics & Evolution 41: 3-27
Abstract
/ref>
* '' Ophrynopus'' Konow, 1897 (synonym: ''Stirocorsia'' Konow, 1897): 17 species in Japan, Indonesia, Laos, Malaysia, Philippines, Papua New Guinea and in the Neotropic realm. Taxonomy: Vilhelmsen & Smith (2002), Vilhelmsen et al. (2013).
* '' Orussella'' Benson, 1935: 1 species, '' Orussella dentifrons'' (Philippi, 1783), in Chile. Taxonomy: Vilhelmsen (2003).
* '' Orussobaius'' Benson, 1938: 9 species in Australia, Papua New Guinea and the Tanimbar Islands. Taxonomy: Schmidt & Vilhelmsen (2002).,Schmidt, S. & Vilhelmsen, L. 2002: Revision of the Australasian genus ''Orussobaius'' Benson (Hymenoptera: Symphyta: Orussidae). Australian Journal of Entomology 41: 226-235. Blank & Vilhelmsen (2016)Blank, S. M. & Vilhelmsen, L. 2016: Two new parasitoid wasp species of the Australasian genus ''Orussobaius'' (Hymenoptera: Orussidae). Arthropod Systematics & Phylogeny 74(1): 83-103
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* '' Orussonia'' Riek, 1955: 2 species in Australia. Taxonomy: Schmidt & Gibson (2001).Schmidt, S. & Gibson, G.A.P. 2001: A New Species of the Genus ''Orussonia'' Riek and the Female of ''O. depressa'' Riek (Hymenoptera: Symphyta, Orussidae). Journal of Hymenoptera Research 10: 113-118.
* ''Orussus'' Latreille, 1797: 29 species in the Holarctic and Oriental realms. Taxonomy: Vilhemsen (2003, species of the World), Blank et al. (2006, ''O. abietinus'' and ''O. smithi''),Blank, S. M., Kraus, M. & Taeger, A. 2006: ''Orussus smithi'' sp. n. and Notes on Other West Palaearctic Orussidae (Hymenoptera). Pp. 265-278. In: Blank, S. M.; Schmidt, S. & Taeger, A. (eds): Recent Sawfly Research: Synthesis and Prospects. Goecke & Evers, Keltern. Vilhelmsen et al. (2013, species of the World).Vilhelmsen, L., Blank S.M., Liu, Z.-W. & Smith, D.R. 2013: Discovery of new species confirms Oriental origin of ''Orussus'' Latreille (Hymenoptera: Orussidae). Insect Systematic & Evolution 44: 1-41
Abstract/PDF
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* '' Pedicrista'' Benson, 1935: 1 species, '' Pedicrista hyalina'' Benson, 1935, in Malawi, South Africa and Zimbabwe. Taxonomy: Vilhelmsen (2003).
* '' Pseudoryssus'' Guiglia, 1954: 2 species in the West Palaearctic. Taxonomy: Kraus (1998), Blank et al. (2006).

Orussidae of the United States, Canada and Great Britain

Twelve species of Orussidae are distributed in the United States and in Canada:

* ''Kulcania mexicana'' (Cresson, 1879) (Florida), ''K. tomentosa'' (Middlekauff) (Arizona, California). Taxonomy: Vilhelmsen & Smith (2002).
* ''Ophrynon dominiqueae'' Blank, Vilhelmsen & D.R. Smith, 2010, ''O. levigatus'' Middlekauff, 1983, ''O. martini'' Blank, Vilhelmsen & D.R. Smith, 2010, ''O. patricki'' Blank, Vilhelmsen & D.R. Smith, 2010, all distributed in California. Taxonomy: Blank et al. (2010).
* ''Ophrynopus nigricans'' (Cameron, 1883) in Texas. Taxonomy: Middlekauff (1983)Middlekauff, W.W. 1983: A Revision of the Sawfly Family Orussidae for North and Central America (Hymenoptera: Symphyta, Orussidae). University of California Publications in Entomology 101: 1-46. (citing the species as ''Ophrynella nigricans''), Vilhelmsen & Smith (2002).
* ''Orussus minutus'' Middlekauff, 1983 (eastern USA), ''O. occidentalis'' Cresson, 1879 (western USA, in Canada from southern British Columbia eastward to Ottawa, Ontario), ''O. sayii'' Westwood, 1835 (eastern USA and Canada), ''O. terminalis'' Newman, 1838 (eastern USA and Canada), ''O. thoracicus'' Ashmead, 1898 (western USA). Taxonomy: Middlekauff (1983), Vilhelmsen et al. (2013).

In Great Britain, ''Orussus abietinus'' (Scopoli, 1763) was recorded by Stephens (1835)Stephens, J.F. 1835: Illustrations of British Entomology; or, a Synopsis of Indigenous Insects: containing their generic and specific distinctions;with an account of their metamorphosis, times of appearance, localities, food, and economy, as far as practicable. Mandibulata. Baldwin & Cradock, London 7: 1-312, tabs XXXV-XLVII. upon two specimens caught by William Elford Leach in Darenth Wood and Devonshire around 1817. Morice (1904)Morice, F.D. 1904: Help-notes towards the determination of British Tenthredinidae, &c. (8). Entomologist's Monthly Magazine, Second Series 40: 49-51. recorded a more recent specimen taken at Hastings about 1880, but after re-examination this turned out to be '' Xiphydria prolongata'' (Geoffroy, 1785) (Xiphydriidae).Benson, R. B. 1935: The alien element in the British sawfly fauna. Ann. appl. Ent. 22(4): 754-768.Benson, R.B. 1943: Some reputed British sawflies not found since Stephens's days (Hym., Symphyta). Entomologist's Monthly Magazine, Fourth Series 79(4): 5-7. Benson (1951)Benson, R.B. 1951: Hymenoptera, Symphyta. Handbooks for the Identification of British Insects 6(2a): 1-49. supposed ''O. abietinus'' "to have occurred in Britain in former times."

Fossils

The oldest members of Orussoidea, the group containing Orussidae and its close relatives, are known from the Middle Jurassic Karabastau Formation of Kazakhstan. The oldest known members of modern Orussidae are from the Late Cretaceous. ''Ophrynopus peritus'' Engel, 2008 was described from Dominican Amber,Engel, M. S. 2008: An orussid wood wasp in amber from the Dominican Republic (Hymenoptera: Orussidae). Transactions of the Kansas Academy of Science 111(1/2): 39-44. '' Baltorussus velteni'' Schedl, 2011 from Baltic Amber, ''Mesorussus taimyrensis'' Rasnitsyn, 1977 from the Late Cretaceous of Taimyr, Sibiria und ''Minyorussus luzzii'' Basibuyuk, Quicke & Rasnitsyn, 2000 from the Late Cretaceous of New Jersey.

References

External links

Waspweb
Excellent photographs of several ''Chalinus'' species.

ECatSym
Complete World catalog of sawflies and horntails, including the Orussidae.

{{Taxonbar, from=Q148072

Sawfly families
Parasitic wasps
Neogene first appearances