Hodostates Förster, 1869

Taxonomic History / Nomenclature
Hodostates Foerster, 1869: 202.
Type species: Hodostatus brevis Thomson, 1883 by subsequent monotypy based on inclusion by Thomson (1883).

Gnesia Foerster, 1869: 202. Preoccupied by Doubleday. Synonymized by Townes (1970): 70.
Type species: Gnesia caliroae Rohwer, 1915.
Designated by Rohwer (1915); first included species.

Hodostatus Thomson, 1883: 929. Emendation.

Esigna Perkins, 1962: 425. New name for Gnesia Förster.

Remarks
The tribal placement of Hodostates is problematic, and in the diagnosis given above, we have also included characters useful for separating Hodostates from genera in other tribes of the Ctenopelmatinae. Townes (1970) specifically questioned his own placement of Hodostates in Pionini because of the possession of a “weak” subapical notch on the ovipositor. The Pionini are essentially characterized by a slender ovipositor lacking a subapical notch Townes (1970) but this feature is also found within Perilissini (e.g. some Lathrolestes Foerster), Mesoleiini (Anoncus Townes), and several Ctenopelmatini. Townes (1970) also characterized most pionines on the basis of two u-shaped grooves between the metanotum and the propodeum (Townes noted Asthenara as exceptional in this regard). Hodostates is unfortunately interspecifically variable for at least one of these grooves. Additionally, these grooves are found in several other non-pionine Ctenopelmatinae and in some Tryphoninae. Thus, the characters used by Townes (1970) to delimit the tribe Pionini are homoplastic and would appear to be of little utility.

The following valid species were included by Yu and Horstmann (1997).
Hodostates brevis (Thomson,1883)
Hodostates palustris (Habermehl, 1925)
Hodostates rotundatus (Davis, 1897)
Hodostates schaffneri (Hinz, 1996)

Diagnosis and Relationships
Hodostates is presently included in the tribe Pionini. It is distinguished from all other Pionini by the combination of 1) distinct subapical notch on the ovipositor (Figs 14, 16); 2) completely areolate propodeum (Fig. 11) with posterior field sharply declivitous (Figs 9, 10); 3) presence of a distinct glymma subbasally (Fig. 12); and 4) long notauli extending posteriorly as distinctly impressed grooves at least over anterior 0.75 of mesoscutum (Fig. 8). Hodostates also resembles certain euryproctine genera with a short, areolate propodeum and a broader T1, but differs from all euryproctines in the possession of a glymma. Hodostates differs from mesoleiine genera by the combination of a completely areolate propodeum, evenly margined clypeus (Figs 5, 6), presence of fore wing areolet (Figs 3, 7), and the exceptionally long, distinctly impressed notauli (Fig. 8). Hodostates can be distinguished from all Perilissini by the combination of the small glymma, deep notauli and clypeus without distinctly impressed margin.

Figures 1 and 9 are of the lectotype of H. brevis; Figs 2, 4, 6, 7, and 10 are of the holotype of palustris Habermehl, a synonym of brevis; Figs 3, 11, and 14 are of the lectotype of H. rotundatus; Figs. 5, 12, and 13 are of other determined specimens of H. rotundatus; Fig. 8 is of the holotype of caliroae Rohwer, a synonym of rotundatus; Figs 15 and 16 are of another determined specimen of H. brevis.

6246_mximage
1. Hodostates brevis Thomso...
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2. Hodostates brevis dorsal...
10207_mximage
3. Hodostates rotundatus (Davis) ...
6029_mximage
4. Hodostates brevis dor...
5975_mximage
5. Hodostates rotundatus ...
6031_mximage
6. Hodostates brevis clypeus and ...
6030_mximage
7.Hodostates brevis wings.
6244_mximage
8. Hodostates rotundatus sh...
6248_mximage
9. Hodostates brevis showin...
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10. Hodostates brevis propod...
6039_mximage
11. Hodostates rotundatus pr...
6014_mximage
12. Hodostates rotundatus ...
5008_mximage
13. Hodostates rotundatus ov...
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14. Hodostates rotundatus oviposito...
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15. Hodostates brevis ventra...
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16. Hodostates brevis dorsal valve...
 
Description
Face densely sculptured. Inner margins of eyes subparallel; not or only weakly emarginate. Clypeus separated from face by a distinct groove throughout; flat to weakly convex in profile with apical margin not or only very weakly protruding; in frontal view apical margin varying in appearance from shallowly and evenly convex to very weakly and broadly truncate medially; uniformly rounded, never sharply impressed nor thickened medially. Mandible not excavated basally; very gradually and evenly narrowing from base to middle; approximately parallel-sided from middle to apex, not twisted; teeth subequal, with ventral tooth appearing slightly longer depending on angle of view. Labial and maxillary palps shorter than head height. First flagellomere not or only very slightly longer than second, with 2-3 irregular rows of 6-8 placoid sensilla in lateral view, these not clustered to form a distinct tyloid. Ocellar triangle not equilateral, posterior ocelli more widely spaced. Occipital carinae complete, joining hypostomal carina ventrally distinctly before reaching mandible. Epomia absent. Notaulus strong, extending at least over anterior 0.75 of mesoscutum as a distinctly incised groove. Dorsal end of epicnemial carinae distinctly separated from anterior edge of mesopleuron (by at least half basal width of mandible); sternaulus completely absent. Scutellum in lateral view varying from conical to more evenly rounded. Pleural carina distinct, well developed throughout; u-shaped notch between lateral longitudinal carina of propodeum and metanotum variable. Propodeum areolate, with complete complement of carinae; all carinae distinctly elevated; petiolar area elongate (distinctly more than half as long as propodeum) and sharply declivous. Hind coxal cavities confluent with propodeal cavity. Tarsal claws not pectinate. Fore wing with areolet present, 2m-cu received nearer 3rs-m than 2rs-m, often with 2m-cu received at distal corner, areolet petiolate or not, 3-5 sided depending on position of 2m-cu and whether petiolate or not; stigma short and broad; 1cu-a inclivous, meeting Cu1 distad of Rs&M; first subdiscal cell longer than tall, parallel-sided, not expanded distally. Hind wing with cu-a vertical or nearly so; basal abscissa of Cu1 varying from slightly shorter to distinctly longer than cu-a; M+Cu weakly but distinctly bowed. First tergite moderately stout, rugulose, dorsal profile arched; glymma small, pit-like, near base, lateral to basal-median depression; dorsal median carinae more or less parallel-sided and widely spaced, extending as strongly elevated ridges at least 0.5 length of petiole (at level of spiracle), weakening posteriorly, occasionally discernible to apex; dorsal median and dorsal lateral carinae converging at base but not meeting; dorsal lateral carinae sharp, extending from base to apex; S1 very short, extending less than half distance to T1 spiracle. T2 without carina extending from base to spiracle; T2 and T3 lacking thyridia; T2 extensively rugulose, at least medially, T3 smooth or nearly so. Cerci short, somewhat broadly triangular, not or only weakly protruding. Ovipositor straight, not up-curved, dorsal valve, where visible, with a deep sub-apical dorsal notch; ovipositor sheath very short.
Distribution
Hodostates brevis Thomson was originally described from Sweden, and this is the only locality given by Kasparyan (1981) and Kasparyan and Khalaim 2007. Kazmierczak 1991 also records it from Austria. The records from Bulgaria (Kolarov 1983, Aubert 2000) are for a species that we do not include in Hodostates. Hodostates kotenkoi Kasparyan is known only from the southern Kurile Islands. The New World species Hodostates rotundatus is known from northeastern U.S.A. and southeastern Canada south to northwestern South Carolina.
Distribution
No referenced distribution records have been added to the database for this OTU.
Biology / Hosts
Known host records are confined to the Tenthredinidae and include the woolly caterpillar Eriocampa umbratica (Klug) (subfamily Allantinae) (Thomson 1888), and slug caterpillars in the genus Caliroa (subfamily Heterarthrinae) (Rohwer 1915, Carlson 1979, Kasparyan and Khalaim 2007). Carlson 1979 indicates that the host plant for the North American species is American chestnut, Castanea dentata, which was incorrectly recorded in the original description by Rohwer 1915.
Map

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

Additional references
Hodostatus brevis Thomson, 1883: 929 (original description); Thomson 1888: 1258 (host); Perkins 1962: 427 (as type species of Hodostates); Fitton 1982: 40 (lectotype designation and label data).
Hodostates brevis: Dalla Torre 1901: 307 (catalog); Perkins 1962: 387, 427 (Thomson emendations); Fitton 1982: 40 (correct name for Hodostatus brevis); Townes 1970: 70 (as type species of Hodostates); Kasparyan 1981: 323 (brief description in key); Kasparyan and Khalaim 2007 : 490 (key, updated host records).
Hodotastes (sic) brevis: Meyer 1936: 289-290 (key, genus redescription).
Polyblastus palustris Habermehl 1925: 10-11 (original description); Meyer 1936: 1-340 (key, Russian fauna).
Hodostates palustris: Kasparyan 1996: 196 (transfer to Hodostates as probable synonym of brevis); Yu and Horstmann 1997: 448 (catalog, as valid species); Yu et al. (2005) (electronic catalog, as synonym of H. brevis).
Hodostates kotenkoi Kasparyan 2007: 490 (key, original description).
Trematopygus rotundatus Davis, 1897: 277 (key, original description); Cresson 1928: 24 (lectotype designation).
Gnesia rotundata: Townes 1945: 505 (catalog); Townes and Townes 1951: 330 (catalog).
Gnesia caliroae Rohwer 1915: 220 (host record, original description); Townes 1945: 505 (catalog); Townes and Townes 1951: 330 (catalog); Townes 1970: 70 (as synonym of rotundatus; inclusion in Hodostates).
Hodostates rotundatus: Townes 1970: 70, 71 (transfer to Hodostates, synonymy); Carlson 1979: 584 (correction of host record for Rohwer’s specimen).
Acknowledgements
This page was assembled by Bob Wharton, and is part of a revision of Hodostates undertaken by Mika Cameron as part of her M.S. thesis completed at Texas A&M University in 2009. The revision has been submitted for publication in The Canadian Entomologist. We are most grateful to the following curators or collection managers and institutions for providing material on loan used in this study: David Wahl (AEI), Jason Weintraub (ANSP), Gavin Broad (BMNH), Andy Bennett (Canadian National Collection, Ottawa), Jens-Peter Kopelke (NHMS), Robert Kula (USNM), Roy Danielsson (MZLU), and Stefan Schmidt (ZSM). We are especially grateful to Stefan Schmidt for discovery of an additional specimen of H. brevis and permission to dissect the ovipositor. We also thank John Cameron and Matt Yoder for their support, Klaus Horstmann for suggestions on location of specimens in European museums, and Patricia Mullins, Heather Cummins, and Cheryl Hyde for considerable assistance with imaging. This work was funded by the National Science Foundation Partnerships for the Enhancement and Education in Taxonomy (NSF-PEET) Grant DEB 0328922 and associated REU supplement 0723663.

This material is based upon work supported by the National Science Foundation under Grant Number DEB 0328922 with REU supplements DEB 0723663.
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.