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Family Heteroceridae - Variegated Mud-loving Beetles

Heteroceridae - Heterocerus collaris Heterocerid - Heterocerus mollinus - male H. collaris-like - Heterocerus collaris mud beetle #6 Heteroceridae, lateral - Heterocerus - male Heterocerus texanus (Pacheco) - Heterocerus texanus Heterocerid - Heterocerus Tiny Beetle on Mullein - Heterocerus
Kingdom Animalia (Animals)
Phylum Arthropoda (Arthropods)
Subphylum Hexapoda (Hexapods)
Class Insecta (Insects)
Order Coleoptera (Beetles)
Suborder Polyphaga (Water, Rove, Scarab, Long-horned, Leaf and Snout Beetles)
Superfamily Byrrhoidea (Pill, Water and Toe-winged Beetles)
Family Heteroceridae (Variegated Mud-loving Beetles)
Synonyms and other taxonomic changes
for most recent treatment see(1)
Note: The classification proposed in(1) is reflected in the Guide. Summary of changes:
The genus Tropicus is retained in its current circumscription
Species currently treated in Augyles, Centuriatus, Explorator, and Microaugyles all go to the genus Augyles
The remaining New World species go (back!) to the genus Heterocerus
~40 spp. in 3 genera in our area (2)(1)
Initial separation into groups of genera and species is done using presence or absence of post-meso and post-meta coxcal lines (seen here in Am. Beetles illus.). The following is adapted from the key to the genera in Am Beetles, further modulated using the updated analysis, summarized above in the taxonomic changes section:

post-metacoxcal lines present:

Augyles Schiødte, 1866

A. auromicans Kiesenwetter 1851
A. blanda Miller 1994 - Alberta, Colorado, and Nebraska
A. canadensis Fall 1920
A. compactus Fall 1937
A. moleculus Fall 1920
A. mundulus Fall 1920

post-mesocoxal lines present, post-meta lines absent

Efflagitatus Pacheco 1964

E. selanderi Pacheco 1969 (Florida)

Heterocerus Fabricius 1792

H. labiatus Kiesenwetter 1851
H. miser Kiesenwetter 1851
H. substriatus Kiesenwetter 1851

"undatus" group (including Pacheco's Dampfius)
"Members of the undatus group are medium sized, more or less reddish colored insects with highly variable elytra markings." (Miller, 1988b)
H. collaris Kiesenwetter 1851 - east and central US
H. inciertus - across Canada, and northcentral US
H. mexicanus Sharp 1882 - western US, TX, LA, IL, and Quebec
H. pallidus Say 1823
H. schwarzi Horn 1890
H. subtilis Miller 1988b - northern central and east US and Canada
H. tenuis Miller 1988a
H. texanus (Pacheco) 1964
H. undatus Melsheimer 1844 - northern central and east US and Canada
H. unicus Miller 1988b - western US and Canada

"mollinus" group (sometimes lumped with gnatho group)
H. insolens Miller 1994 - southcentral US
H. brunneus Melsheimer 1844
H. gemmatus Horn 1890
H. mollinus Kiesenwetter 1851
H. parrotus (Pacheco) 1964
H. sinuosus (Pacheco) 1964

"fenestratus" group
H. fenestratus Thunberg (L. tristis Mannerheim 1853)

post-meso and post-metacoxal lines absent

"gnatho" group
H. angustatus Chevrolat 1864
H. aptus Miller 1994 - TX
H. fatuus Kiesenwetter 1851
H. glicki (Pacheco) 1963
H. gnatho LeConte 1863
H. infrequens Miller 1994 - northcentral US and across Canada
H. intermuralis (Pacheco) 1963
H. longilobulus (Pacheco) 1964
H. mirus Miller 1994 - Ontario
H. sandersoni (Pacheco) 1964

Tropicus Pacheco 1964

T. minutus (Fall) - single collection from southern Texas
T. nigrellus King and Lago 2012 - rare southeastern species
T. pusillus (Say) - one of the most common and widespread heterocerids in the Western Hemisphere
sandy or muddy shores of rivers, lakes, and ponds
Print References
King, JG and P.K. Lago. 2012. The variegated mud-loving beetles (Coleoptera: Heteroceridae) of Mississippi and Alabama, with discussion and keys to the species occurring in the southeastern United States. Insecta Mundi 0275: 1-53. (Full Text: (3))
King, J.G, J.R. Starr, and P.K. Lago.2011. Molecular data resolves relationships within Heteroceridae (Dryopoidea: Coleoptera). Systematic Entomology 36: 435-445.
Pacheco, F. 1964. Sistemática, filogenia y distribución de los Heteroceridos de América(Coleoptera: Heteroceridae). Monogràfías del Colegio de Postgraduados,Chapingo, Mexico. 115 pp.
Pacheco, F. 1969. A new species of Heterocerini (Coleoptera: Heteroceridae). Florida Entomol. 52:37-39.
Pacheco, M.F. 1978. A catalog of the Coleoptera of America north of Mexico. Family Heteroceridae. U. S. Dept. of Agric., Agric. Handbook Number 529-47. x + 9 pp. (4)
Miller, W.V. 1988a. A new species of Heterocerus recorded from Florida and Canada (Coleoptera: Heteroceridae). Florida Entomol. 71: 30-33.
Miller, W.V. 1988b. Two new species of Heterocerus from North America, with notes on related species (Coleoptera: Heteroceridae). Coleopterists Bulletin 42(4): 313-320.
Miller, W.V. 1994. New species of Heteroceridae (Coleoptera) from Canada and the United States. Coleopterists Bulletin 48(1): 11-18.
Schaeffer, K.F. and W.A. Drew. 1964. The Heteroceridae (Coleoptera) of Oklahoma. Proceedings of the Oklahoma Academy of Sciences 45: 69-71. (Full text)
Works Cited
1.Molecular data resolves relationships within Heteroceridae (Coleoptera: Dryopoidea)
J.G. King, J.R. Starr, P.K. Lago. 2011. Systematic Entomology 36(3): 435–445.
2.American Beetles, Volume II: Polyphaga: Scarabaeoidea through Curculionoidea
Arnett, R.H., Jr., M. C. Thomas, P. E. Skelley and J. H. Frank. (eds.). 2002. CRC Press LLC, Boca Raton, FL.
3.The variegated mud-loving beetles (Heteroceridae) of Mississippi and Alabama, with discussion and keys to se US spp.
King and Lago. 2012. Insecta Mundi 0275: 1-53.
4. A catalog of the Coleoptera of America north of Mexico. Family Heteroceridae.
Pacheco, F. 1978. USDA, Agric. Handbook. x + 9 pp. 1978. USDA.