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Containing group: Cucujoidea

Introduction

Biphyllidae is a relatively small family of small- to medium-sized mycophagous beetles.  The family is cosmopolitan in distribution, occurring in all zoogeographic regions except New Zealand, with highest diversity in the tropics.  However, its members are often difficult to collect and rarely appear in most collections.  The family currently includes six genera and approximately 200 species (Lawrence 1982, Goodrich and Springer 1992, Cline and McHugh 2010).

Cline and Shockley (2010) summarized the known biology and ecology of the family.  Both adults and larvae appear to be obligately mycophagous.  The most common method for collecting biphyllids is in association with their host Ascomycete fungus. Biphyllids have been collected in association with the following ascomycete fungi:  Cryptostroma, Daldinia, Hypoxylon, Nammularia, Tubercularia, and Xylaria (see Cline and Shockley 2010 for specific associations and references).  Specimens may also be collected by peeling bark from dead trees, mostly hardwoods, or by sifting moist leaf litter and other decaying plant material.  Biphyllids do not appear to be attracted to UV or mercury vapor lights, and are only infrequently encountered in malaise or flight intercept traps.

Characteristics

Adults of this family are easily recognizable among cucujoid beetles.  In general, the adults can be recognized by the presence of lateral and/or femoral lines on the first abdominal sternite and/or postcoxal lines on the metasternite.  In general, adults can be characterized by the following combination of characters (based on Lawrence et al. 1999a, Goodrich 2002, Cline and Shockley 2010): 

• Elongate-oval dorsal habitus, somewhat dorsoventrally flattened but still conspicuously convex
• Antennae 11-segmented with a 3-segmented club; antennal grooves present, each groove bearing a lateral pre-gular pocket
• Procoxal cavities closed behind, mesocoxal cavities open laterally
• All coxae distinctly, sometimes widely, separated
• Tarsal formula 5-5-5 with slender fleshy lobes on tarsomeres II and III
• Lateral and/or femoral lines present on abdominal sternite I (ventrite III)
• Elytra bearing distinct serially striate punctation

Larvae for this group are poorly known.  Biphyllid larvae most closely resemble those of Byturidae, but can be distinguished from byturids and other similar larvae by the following combination of characters (based on Lawrence 1991, Lawrence et al. 1999b, Goodrich 2002, Cline and Shockley 2010): 

• Body elongate, parallel-sided, cylindrical to slightly flattened
• Dorsum not heavily sclerotized, lightly to moderately pigmented, smooth
• Head prognathous, epicranial stem absent or short, frontal sutures lyriform
• 6 pairs of stemmata
• Antennae short, 3-segmented, sensory appendage relatively short (much shorter than antennomere III)
• Mandibles bearing an accessory ventral process and well-developed, sickle-shaped mola bearing a hyaline lobe at base
• Maxillary palpi 3-segmented; labial palpi 2-segmented
• Legs well-developed, 5-segmented with unisetose tarsungulus
• Urogomphi usually absent, but short and fixed if present

Taxonomy

Although Silfverberg (1992) and Goodrich and Springer (1992) credit Sharp (1900) for establishment of the family-group name, Lawrence and Newton (1995) pointed out that this assertion of family-group author assignment is incorrect.  The family-group name was actually first established by LeConte (1861), based on the genus Diphyllus.  Unfortunately, Diphyllus was an unjustified emendation of Biphyllus by Redtenbacher (1858).  Under the rule of priority, correction of this emendation and the subsequent synonymy of Diphyllus under Biphyllus makes LeConte's family-group name valid, albeit under the corrected name Biphyllidae.  Arrow (1929) is generally credited with formal elevation of the group to the family-level.  Most subsequent workers (Leng and Mutchler 1933, Schenkling 1934, Blackwelder 1945, Arnett 1963) have continued to treat biphyllids as a distinct family.

Relationships of Biphyllidae to Other Beetles

There has been considerable confusion regarding the phylogenetic relationship of Biphyllidae to other beetle families.  Over the years, species of Biphyllidae have been placed (always together as a group) within a number of other families including Mycetophagidae (LeConte 1861, Reitter 1877, Horn 1878, LeConte and Horn 1883), Cryptophagidae (Redtenbacher 1858, Thomson 1863, Reitter 1887, Casey 1900), Erotylidae (Ganglbauer 1899, Kuhnt 1911, Roberts 1958), and Byturidae (Falcoz 1926).  In addition, potential relationships to Bothrideridae, Nitidulidae, Protocucujidae and Cerylonidae have also been suggested (Crowson 1967, Lawrence 1991).  Cline and Shockley (2010) pointed out that the pre-gular pockets on the head and the presence of lateral/femoral lines on abdominal sternite I combine as potential synapomorphies to unite all members of the family.  Current consensus places Biphyllidae within Cucujoidea (Crowson 1967), sister to Byturidae (Falcoz 1926, Barber 1942, Crowson 1967, Goodrich & Springer 1992).

To date, there have been only two phylogenetic studies published that included members of Biphyllidae, offering the only insights into the evolutionary relationship of Biphyllidae to other cucujoid taxa.  Leschen et al. (2005) performed the first cladistic analysis of the basal cucujoid families (excluding the Cerylonid Series) based on adult and larval morphology.  Unfortunately, adult and larval morphology were incongruent in their placement of Biphyllidae.  Adult morphology supported a sister group relationship between Biphyllidae and Erotylidae, while larval morphology supported a relationship between Biphyllidae and Byturidae.

More recently, a molecular phylogenetic analysis of Coleoptera based on 3 genes (Hunt et al. 2007) recovered Biphyllidae as sister to Byturidae, placed within the superfamily Cleroidea.  However, this placement was not strongly supported and is likely an artifact of the inappropriate scope and taxon sampling used for the study, which was targeting the more basal nodes within the coleopteran phylogeny.  Nonetheless, the sister group relationship between Biphyllidae and Byturidae was strongly supported.

Discussion of Phylogenetic Relationships

No phylogenetic studies have yet been performed to assess intra-familial relationships at any level within Biphyllidae.

Other Names for Biphyllidae

• Diphyllidae

References

Arnett, R.H. 1963. The beetles of the United States. Catholic University of America Press, Washington, DC. 1112 pp.

Arrow, G.J. 1929. On the families of Coleoptera related to the Erotylidae, with descriptions of a new family, two new genera, and a few new species. Annals and Magazine of Natural History (ser. 10) 4: 305-322.

Barber, H.S. 1942. Raspberry fruitworms and related species. Miscellaneous Publications of the U.S. Department of Agriculture 468: 1-32.

Blackwelder, R.E. 1945. Checklist of the coleopterous insects of Mexico, Central America, the West Indies, and South America. Bulletin of the U.S. National Museum 185: 343-550.

Casey, T.L. 1900. Review of the American Corylophidae, Cryptophagidae, Tritomidae and Dermestidae, with other studies. Journal of the New York Entomological Society 8: 51-172.

Cline, A.R., and McHugh, J.V. 2010. New generic synonymy in Biphyllidae (Coleoptera: Cucujoidea) with a checklist of Anchorius Casey, 1900. The Coleopterists Bulletin 64: 98-99.

Cline, A.R., and Shockley, F.W. 2010. 10.6. Biphyllidae LeConte, 1861. Pp. 306-311. In: Leschen, R.A.B., Beutel, R.G., and Lawrence, J.F. (eds.), Handbuch der Zoologie/Handbook of Zoology. Band/Volume IV. Arthropoda: Insecta Teilband/Part 38. Coleoptera, Beetles. Volume 2. Morphology and Systematics (Polyphaga partim). W. DeGruyter, Berlin.

Crowson, R.A. 1967. The Natural Classification of the Families of Coleoptera. E.W. Classey Ltd., Middlesex, England. 214 pp.

Falcoz, L. 1926. Position systematique des genres Diphyllus Stephens and Diplocoelus Guérin. Pp. 69-74. In: Lesne, P. (ed.), Encyclopedie de Entomologique Serie B, Memoires et Notes. I. Coleoptera. Vol. 1. P. Lechevalier, Paris.

Ganglbauer, L. 1899. XXVI. Familie. Erotylidae. Pp. 629-739. In: Ganglbauer, L. (ed), Die Käfer Mitteleuropas. Wien, Germany.

Goodrich, M.A. 2002. 89. Biphyllidae LeConte 1861. Pp. 356-357. In: Arnett,Jr., R.H., Thomas, M.C., Skelley, P.E., and Frank, J.H. (eds.), American Beetles. Volume 2. Polyphaga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton.

Goodrich, M.A., and Springer, C.A. 1992. A Revision of the Family Biphyllidae (Coleoptera) for America, North of Mexico. The Coleopterists Bulletin 46: 361-377.

Horn, G.H. 1878. Synopsis of Mycetophagidae. In: Hubbard, H.G., and Schwarz, E.A. The Coleoptera of Michigan. Proceedings of the American Philosophical Society 17: 603-606.

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Kuhnt, P. 1911. Erotylidae. Coleopterorum Catalogus 15(34): 1-103.

Lawrence, J.F. 1982. Coleoptera. Pp. 482-553. In: Parker, S. P. (ed.), Synopsis and Classification of Living Organisms, Vol. 2. McGraw-Hill, New York.

Lawrence, J.F. 1991. Coleoptera. Family Biphyllidae (Cucujoidea). Pp. 475-476. In: Stehr, F.W. (ed.), Immature Insects, Vol. 2. Kendall/Hunt, Dubuque, Iowa.

Lawrence, J.F., and Newton, Jr., A.F. 1995. Families and subfamilies of Coleoptera (with selected genera, notes, references and data on family-group names). Pp. 779-1006. In: Pakaluk, J., and Ślipiński, S.A. (eds.), Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Museum i Instytut Zoologii PAN, Warszawa.

Lawrence, J.F., Hastings, A.M., Dalwitz, M.J., Paine, T.A., and Zurcher, E.J. 1999a. “Beetles of the World: A Key and Information System for Families and Subfamilies.” CD-ROM, Version 1.1 for MS-Windows. CSIRO Publishing, Melbourne.

Lawrence, J.F., Hastings, A.M., Dalwitz, M.J., Paine, T.A., and Zurcher, E.J. 1999b. “Beetle Larvae of the World: Descriptions, Illustrations, Identification, and Information Retrieval for Families and Subfamilies.” CD-ROM, Version 1.0 for MS-Windows. CSIRO Publishing, Melbourne.

LeConte, J.L. 1861. Classification of the Coleoptera of North America. Smithsonian Miscellaneous Collections 3: 1-208.

LeConte, J.L., and Horn, G.H. 1883. Classification of the Coleoptera of North America, vol. 507. Smithsonian Museum, Washington, DC.

Leng, C.W., and Mutchler, A.J. 1933. Second and third supplements 1925 to 1932 (inclusive) to the Catalog of the Coleoptera of America, north of Mexico. John D. Sherman, Jr., Mount Vernon, NY. 112 pp.

Leschen, R.A.B., Lawrence, J.F., and Ślipiński, S.A. 2005. Classification of basal Cucujoidea (Coleoptera: Polyphaga): Cladistic analysis, keys and review of new families. Invertebrate Systematics 19: 17-73.

Redtenbacher, L. 1858. Fauna Austriaca. Die Kafer. Nach der analytischen Methode bearbeitet. Zweite Auflage. C. Gerolds Sohn, Wein. 1017 pp.

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Reitter, E. 1887. Bestimmungs-Tebellen der europaischen Coleopteren. XVI Heft. Enthaltend die Familien: Erotylidae und Cryptophagidae. Verhandlungen des Naturforschenden Vereins in Brunn 26: 1-55.

Roberts, A.W.R. 1958. On the taxonomy of Erotylidae (Coleoptera), with special reference to the morphological characters of the larvae. Transactions of the Royal Society of London 110: 245-285.

Schenkling, S. 1934. Biphyllidae. Coleopterorum Catalogus 15(133): 1-7.

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Silfverberg, H. 1992. Enumeratio Coleopterorum Fennoscandiae, Danieae et Baltiae. Helsingin Hyonteisvaihtoyhdistys, Helsinki. 94 pp.

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