Scolobates Gravenhorst, 1829

Taxonomic History / Nomenclature
Scolobates Gravenhorst, 1829: 357.

Type species: Scolobates crassitarsus Gravenhorst, 1829 (= Scolobates auriculatus (Fabricius, 1804)), by subsequent designation of Westwood (1840).

Gravenhorst, 1829: 357-360

(descriptions of genus and three species).
Nees von Esenbeck 1830: 1130-1150 (generic redescription).
Stephens 1835: 306 (generic and species redescriptions).
Westwood 1840: 57 (generic synopsis).
Brullé 1846: 322 (generic redescription).
Ratzeburg 1848: 76-77 (generic redescription; figures).
Foerster 1869: 157-159 (key to genera of Banchoidae).
Kriechbaumer 1877: 133-137 (discussion of S. longicornis morphology; collections holdings; species description).
Bridgman and Fitch 1884: 69-71 (key to genera).
Cresson 1887: 205-210 (key to genera of Ophioninae).
Davis 1897: 264 (key to genera, generic and species redescriptions).
Dalla Torre 1902: 283, 327-328 (catalog).
Schmiedeknecht 1907: 613-628 (keys to tribes and genera).
Morley 1911: 344 (generic and species redescriptions).
Schmiedeknecht 1912: 2357, 2490-2493 (key to Scolobatina, generic and species descriptions and redescriptions).
Morley 1913: 140 (generic and species descriptions and redescriptons).
Roman 1915: 4 (key to genera; discussion of genus morphology).
Meyer 1936: 7-8, 137-139 (keys to genera of Tryphonini and species of Scolobates: generic and species redescriptions).
Townes 1945: 480 (catalog).
Townes and Townes 1951: 321 (catalog).
Hopper 1959: 169 (etymological information).
Townes et al. 1965: 661 (catalog).
Townes 1970: 102, 104 (key to genera, generic redescription).
Carlson 1979: 592-593 (catalog).
Kasparyan 1981: 361-431 (key to genera, generic and species redescriptions).
Yu and Horstmann 1997: 455 (catalog).
Zhaurova and Wharton 2009: 1-77 (key to genera; generic redescription; analysis of relationships).

Aglyphus Giraud, 1872: 411.
Type species: Aglyphus nigripennis Giraud, 1872. Monobasic.
Giraud, 1872: 411 (description).
Dalla Torre 1902: 1055 (catalog, as genus incertae sedis).
Viereck 1914: 186 (type species).
Townes 1945: 480 (catalog; synonymy).
Townes and Townes 1951: 321 (catalog; synonymy).
Townes 1970: 104 (synonymy).
Yu and Horstmann 1997: 455 (catalog; synonymy).

Parabraconia Schmiedeknecht, 1914: 2803.
Type species: Parabraconia nigripennis Schmiedeknecht, 1925, by subsequent inclusion by Schmiedeknecht (1925) (= Scolobates nigripennis (Giraud, 1872)).
Schmiedeknecht, 1914: 2803 (description).
Schmiedeknecht 1925: 3121-3122 (first included species).
Meyer 1936: 340 (fauna of Russia and adjacent countries).
Townes 1970: 104 (synonymy).
Carlson 1979: 592-593 (catalog).
Yu and Horstmann 1997: 455 (catalog; synonymy).

Two genus-group names, Aglyphus Giraud and Parabraconia Schmiedeknecht, both apparently based on nigripennis Giraud, have been treated as synonyms of Scolobates (Townes 1945, 1970, Carlson 1979, Yu and Horstmann 1997). However, S. nigripennis (Giraud) is a junior secondary homonym of S. nigripennis Sichel, 1860, and de Gaulle (1908) also treated nigripennis Giraud as a junior objective synonym of nigripennis Sichel (Horstmann 1983). The catalog by Yu and Horstmann (1997) lists both nigripennis Giraud and nigripennis Schmiedeknecht as junior synonyms of nigripennis Sichel.

During this study, we examined a specimen from Spain in BMNH determined as nigripennis Sichel by Aubert, who apparently compared it with the type. Based on this specimen, S. nigripennis Sichel is a valid species with black wings (Figs. 1, 2), which differs in several respects from other species of Scolobates that we have examined, most notably in the absence of pleural carinae. If this and other differences ultimately result in recognition as a separate genus, the types of Giraud and Sichel will need to be compared to verify the valid name for the type species of Aglyphus. Although this problem appears to be straightforward, recent catalogs have either ignored or overlooked the fact that Townes (1945) treated nigripennis Giraud as a junior objective synonym of S. auriculatus (Fabricius), a pale-winged species.

Scolobates consistently appears as a basal cluster in our analyses (Zhaurova and Wharton 2009, Figs. 55-62), but not as a monophyletic group in part because of our focus on capturing the full range of variation in the coding of the matrix. Based on our analyses, we treat the sharply pointed median tooth as independently derived relative to that of Onarion, and thus the best evidence for monophyly of Scolobates. Otherwise, Scolobates is characterized primarily by having the hind wing venation better developed than in other scolobatines, a plesiomorphic feature. Gauld (1997), in his treatment of the Costa Rican Scolobatini, noted that Onarion was characterized by the elongate malar space. The malar space is also very large in Scolobates (which is not known from the Neotropics), though the shape of the head is different in members of these two genera.

Included species:
S. auriculatus (Fabricius, 1804)
S. fennicus Schmiedeknecht, 1912
S. longicornis Gravenhorst, 1829
S. marshalli Vollenhoven, 1878
S. nigerrimus Ulbricht, 1922
S. nigriabdominalis Uchida, 1952
S. nigripennis Sichel, 1860
S. nigriventralis He and Tong, 1992
S. pyrthosoma He and Tong, 1992
S. ruficeps Uchida, 1932
S. rufiventris Ozols and Djanelidze, 1966
S. testaceus Morley, 1913

1. Scolobates nigripenni...
2. Scolobates nigripenni...
Diagnosis and Relationships
Unlike other scolobatines (as well as most other ctenopelmatines), both Scolobates and Onarion possess a small, median, sharply pointed tooth on the apical margin of the clypeus (Figs. 1, 2). Onarion has an unusually elongate labiomaxillary complex, Scolobates does not. The occipital carina (Fig. 3) is also much better developed in Scolobates.
1. Scolobates auriculatus fa...
2. Scolobates auriculatus show...
3. Scolobates auriculatus back...
Length: body (Fig. 1) 5.3-10.4 mm, fore wing: 4.6-9.8 mm. Clypeal margin blunt laterally, with a narrow, sharp, slightly protruding median tooth (Fig. 2). Clypeus about 2.3-3.2 times as broad as long, apically protruding and basally slightly concave in profile, not separated from face by a distinct sulcus dorsomedially (Fig. 2). Anterior tentorial pits oval, with lateral corners downturned. Malar space about 1.2-1.4 times longer than basal width of mandible. Mandible with ventral tooth slightly longer than dorsal tooth, ventral tooth distinctly separated into an upper, more basal blunt area and a lower extended apex. Mandible triangular, wide at base and narrowing at apex (Fig. 2). Mouthparts normal, labiomaxillary complex not elongate (Fig. 5). Face 1.3-1.4 times as wide as long, smooth or lightly punctate, lightly to quite strongly pubescent, with a small median tooth or tubercle dorsally. Inter-antennal area flat to slightly raised medially, anterior margin of torulus situated at about 0.6-0.7 of eye height. Widest diameter of torulus slightly wider than widest diameter of median ocellus. Area between lateral ocelli flat to slightly depressed, distance between lateral ocelli 0.7-1.2 times their widest diameter, distance from lateral ocellus to eye margin 1.3-2.1 times widest diameter of lateral ocellus. Area behind ocelli sharply declivitous. Antennae with 33-46 flagellomeres, as long as to distinctly longer than body. First flagellomere with a small tyloid laterally (Figs 3, 4), length of second flagellomere 0.6-0.7 times length of first. Occipital carina incomplete, present on ventral 0.4 of head, occipital and hypostomal carina join at mandibular base, or hypostomal carina terminates just before mandibular base. Pronotum dorsally with distinctly impressed transverse groove, anterior margin of pronotum weakly to strongly emarginate and exceptionally narrow dorsomedially. Lateral groove of pronotum varying from entirely absent to complete (extending from middorsal to midventral margins in lateral view). Pronotum impunctate. Mesoscutum glabrous, pubescence sparse or absent; notauli absent. Epicnemial carina extending dorsally along ventral 0.3 of pronotum, never reaching anterior margin of mesopleuron. Mesopleuron pubescent, smooth to lightly punctate. Propodeum with pleural carinae usually complete, absent in at least one species, lateral longitudinal carina varying from absent to present over apical 0.6, median longitudinal carina absent; propodeum unsculptured, largely impunctate, varying from pubescent (especially laterally) to bare (especially medially). Posterolateral edge of propodeum upcurved. Hind trochanter 1.8-2.0 times as long as basally wide, apical margin of hind trochanter reaching apical margin of trochantellus. Claws fully pectinate (Fig. 6). Fore wing with abscissa of Cu1 between 1m-cu and 2cu-a about 0.6-0.8 times length of 2cu-a. Marginal cell about 3.0-3.2 times as long as wide. Hind wing with distal abscissa of 1A reaching at least half way to wing margin, but never reaching the margin in the form of a tubular vein. T1 about 2.0-2.7 times as long as posteriorly broad, spiracles flat. Cerci variable, usually quite prominent.
1. Scolobates auriculatus ...
2. Scolobates auriculatus fa...
3. Scolobates auriculatus ...
4. Scolobates auriculatus c...
5. Scolobates sp. from Japan: mout...
6. Scolobates sp. pectinate...
Widespread; Holarctic and Oriental Regions. New World: previously recorded in Canada and USA, as far south as Texas (Davis 1897, Carlson 1979). The southernmost record we have seen is from Fredricksburg in central Texas. Old World: Western Europe through Japan, and south from Turkey to Indonesia.
No referenced distribution records have been added to the database for this OTU.
Biology / Hosts
Three species have been reared from sawflies, two from the genus Arge Schrank (Argidae) (Pschorn-Walcher and Kriegl 1965, Carlson 1979) and one from Cimbex (Cimbicidae) (Rudow 1917). The latter needs verification. Three of the specimens of S. auriculatus that we examined from Ontario, Canada (CNC) and three others reared by Dyar (USNM) were mounted with cocoons and host remains. Scolobates cocoons were inside the sawfly cocoons.

There are no specimens currently determined for this OTU, or those specimens determined for this OTU are not yet mappable.

This page was assembled by Bob Wharton. It is part of a revision of the genera of Westwoodiini and Scolobatini conducted by Kira Zhaurova as part of her M. S. thesis in Entomology at Texas A&M University, completed in 2005, and published in the Contributions of the American Entomological Institute (Zhaurova and Wharton 2009). Page last updated April, 2011.

This work would not have been possible without the groundwork provided by Ian Gauld’s study of the Costa Rican and Australian faunas, and we are particularly grateful for his assistance in many aspects of this study. We also thank the following curators and researchers for extended loans of the material used for this revision: David Wahl (AEIC), John LaSalle (ANIC), Ian Gauld and Gavin Broad (BMNH), Andy Bennett (CNC), Gabriel Melo (DZUP), Anders Albrecht and Pekka Malinen (FMNH), Ronald Zúñiga (INBio), Ken Walker (MVMA), Hege Vårdal (NHRS), Chris Burwell (QM), and Dave Furth (USNM). Matt Yoder provided considerable assistance along the way, particularly with databasing. Images used here were obtained through the combined efforts of Kira Zhaurova, Heather Cummins, and Patricia Mullins. Our use of PURLs ( follows the example of their use in publications by Norm Johnson. This content is based upon work conducted at Texas A&M University and supported by the National Science Foundation’s PEET program under Grant No. DEB 0328922 and associated REU supplement nos DEB 0723663 and DEB 0616851.

This material is based upon work supported by the National Science Foundation under Grant Number DEB 0328922 with REU supplements DEB 0723663 and 0616851.
Any opinions, findings, and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation.