Belongs within: Tenebrionoidea.
Contains: Zopherini, Synchitini.
The Zopheridae, ironclad beetles, are a group of beetles usually found living under bark of dead or dying trees. Most feed on decaying plant tissue or fungi but species of the genus Nematidium are predators of the larvae of wood-boring weevils (Bouchard 2014).
Some authors have divided representatives between distinct families Zopheridae sensu stricto, with the first four abdominal ventrites connate, and Colydiidae, with fewer connate ventrites. The Usechini, found in western North America and Japan, are distinguished by antennal cavities on the anterolateral dorsal surface of the pronotum (Ivie 2002b).
Characters (from Ironclad ID): Antenna 9–11 segmented with a usually abrupt, 1–3 segmented club; antennal insertions concealed from above; mesocoxal cavities closed; trochanters heteromeroid; tarsal formula 4-4-4 or 5-5-4; male genitalia with tenebrionoid aedeagus.
<==Zopheridae [Pycnomerinae]
| i. s.: NotocoxelusE02
| HeterargusED04
| LatometiniI02b
| |--Orthocerodes Slipinski & Lawrence 1999I02b
| |--Notorthocerus Slipinski & Lawrence 1999I02b
| `--Latometus Erichson 1842I02b
| `--L. pubescens Erichs. 1842M86
|--ZopherinaeMW15
| |--ZopheriniB14
| `--UsechiniB14
| |--Usechus Motschulsky 1845I02b
| | |--U. lacerta Motschulsky 1845I02b
| | `--U. nucleatus Casey 1889I02b
| `--Usechimorpha Blaisdell 1929I02b
| |--U. barberi Blaisdell 1929I02b
| `--U. montanus Doyen & Lawrence 1979B14
`--Colydiinae [Adimeridae, Colydiidae, Monoedidae, Orthoceridae]MW15
| i. s.: EnarsusWH02
| DitomaC01
| |--D. crenataC01 [=Colydium (Didoma) crenatumG20]
| |--D. hilaris Blackb. 1887M96
| |--D. lineatocollis Blackb. 1887M96
| |--D. nivicola Blackb. 1891M96
| |--D. obscura Blackb. 1887M96
| |--D. parva Blackb. 1887M96
| |--D. perforata Blackb. 1887M96
| |--D. pulchra Blackb. 1887M96
| `--D. torrida Blackb. 1891M96
| Rhopalocerus Redtenbacher 1842 [Rhopalocerini]I02a
| SparactusLB91
| |--S. costatus Blackb. 1891M96
| |--S. elongatus Blackb. 1891M96
| |--S. proximus Blackb. 1891M96
| `--S. pustulosus Blackb. 1891M96
| TodimaLB91
| |--T. fusca Grouv. 1893M96
| `--T. rufula Grouv. 1893M96
| AblabusLB91
| DryptopsLB91
| Nematidium Erichson 1845 [Nematidiini]I02a
| |--N. (Nematidium) filiforme LeConte 1863I02a
| `--N. (Paleonematidium Heinze 1954)I02a
| Monoedus Horn 1882 [incl. Adimerus Sharp 1894; Adimerini, Monoedini]I02a
| `--M. guttatus (Horn 1882)I02a
| CiconesP92
| DiodesmaP92
| EndophloeusP92
| XylolaemusP92
| Sarrotrium [Sarrotriini]M96
| `--S. australe Blackb. 1891M96
| Phormesa epitheca Oll. 1889M96
| GempylodiniI02a
| |--Gempylodes tmetus Oll. 1889M96
| |--Aprostoma Guérin-Méneville 1939I02a
| `--Pseudendestes Lawrence 1980I02a
| |--P. australisLB91
| `--P. robertsiLB91
| Sympanotus australis Grouv. 1893M96
| Rhagodera Mannerheim 1843 [Rhagoderini]I02a
| `--R. costataI02a
| AcropiniI02a
|--ColydiiniI02a
| |--Colydium Fabricius 1792MW15, I02a
| | |--C. elongatumG20
| | `--C. lineolaI02a
| `--Aulonium Erichson 1845I02a, FS90
| |--A. bidentatum Fab. 1801FS90
| `--A. parallelopipedumI02a
`--+--Phellopsis LeConte 1862MW15, I02b [incl. Pseudonosoderma Heyden 1885I02b; Phellopsini]
| |--P. obcordata (Kirby 1837) [incl. P. montana Casey 1907, P. robustula Casey 1907]I02b
| `--P. porcataMW15
`--+--SynchitiniMW15
`--PycnomeriniI02b
|--Pycnomerodes Broun 1886I02b
|--Docalis Pascoe 1860I02b
| |--D. degener Pascoe 1860M86
| `--D. exoletus Pascoe 1860M86
|--Cotulades Pascoe 1860I02b
| |--C. fascicularis Pascoe 1860M86
| |--C. funerosa Hope 1845M86
| `--C. leucospila Hope 1845M86
`--Pycnomerus Erichson 1842MW15, I02b (see below for synonymy)
|--‘Penthelispa’ exarata Chv. 1863FS90
|--P. haematodesI02b
|--P. helmsiH15
|--P. longulus Sharp 1886M96
`--P. moestus Oll. 1889M96
Pycnomerus Erichson 1842MW15, I02b [incl. Dechomus Jacquelin du Val 1859I02b, Endectus LeConte 1863I02b, Penthelispa Pascoe 1863I02b, Pycnomorhus Motschulsky 1858I02b]
*Type species of generic name indicated
References
[B14] Bouchard, P. (ed.) 2014. The Book of Beetles: A lifesize guide to six hundred of nature’s gems. Ivy Press: Lewes (United Kingdom).
[C01] Csiki, E. 1901. Bogarak [Coleopteren]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 75–120. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.
[E02] Emberson, R. M. 2002. The beetle (Coleoptera) fauna of the Chatham Islands: additions and corrections. New Zealand Entomologist 25: 69–77.
[ED04] Ewers, R. M., & R. K. Didham. 2004. A first assessment of New Zealand beetles restricted to large forest areas. Weta 27: 33–40.
[FS90] Fleutiaux, E., & A. Sallé. 1890. Liste des coléoptères de la Guadeloupe et descriptions d’espèces nouvelles. Annales de la Société Entomologique de France, 6e série 9: 351–484.
[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.
[H15] Hedley, C. 1915. Presidential address. Journal and Proceedings of the Royal Society of New South Wales 49 (1): 1–77, pls 1–7.
[I02a] Ivie, M. A. 2002a. Colydiidae Erichson 1845. In: Arnett, R. H., Jr, M. C. Thomas, P. E. Skelley & J. H. Frank (eds) American Beetles vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea pp. 445–453. CRC Press.
[I02b] Ivie, M. A. 2002b. Zopheridae Solier 1834. In: Arnett, R. H., Jr, M. C. Thomas, P. E. Skelley & J. H. Frank (eds) American Beetles vol. 2. Polyphaga: Scarabaeoidea through Curculionoidea pp. 457–462. CRC Press.
[LB91] Lawrence, J. F., & E. B. Britton. 1991. Coleoptera (beetles). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 2 pp. 543–683. Melbourne University Press: Carlton (Victoria).
[M86] Masters, G. 1886. Catalogue of the described Coleoptera of Australia. Part IV. Proceedings of the Linnean Society of New South Wales, series 2, 1 (2): 259–380.
[M96] Masters, G. 1896. Catalogue of the described Coleoptera of Australia. Supplement, part II. Dytiscidae, Gyrinidae, Hydrophyllidae, Staphylinidae, Pselaphidae, Paussidae, Silphidae, Scaphididae, Histeridae, Phalacridae, Nitidulidae, Trogositidae, Colydiidae, Cucujidae, Cryptophagidae, Lathridiidae, Mycetophagidae, Dermestidae, Byrrhidae, Parnidae, Heteroceridae. Proceedings of the Linnean Society of New South Wales 21 (Suppl.): 695–754.
[MW15] McKenna, D. D., A. L. Wild, K. Kanda, C. L. Bellamy, R. G. Beutel, M. S. Caterino, C. W. Farnum, D. C. Hawks, M. A. Ivie, M. L. Jameson, R. A. B. Leschen, A. E. Marvaldi, J. V. McHugh, A. F. Newton, J. A. Robertson, M. K. Thayer, M. F. Whiting, J. F. Lawrence, A. Ślipiński, D. R. Maddison & B. D. Farrell. 2015. The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology 40 (4): 835–880.
[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.
[WH02] Worthy, T. H., & R. N. Holdaway. 2002. The Lost World of the Moa: Prehistoric life of New Zealand. Indiana University Press: Bloomington (Indiana).