306 Andrew R. Cline and Floyd W. Shockley

McHugh, J. V. (1993 a): A revision of Eurysphindus species are known from Europe (Reitter 1909; LeConte (Coleoptera: Cucujoidea: Sphindidae) and Winkler 1924). Within the Nearctic, two Dipla­ a review ofsphindid classification and phylogeny. coelus and one Anchorius Casey species are known -SystematicEntomology 18: 57-92. (Casey 1900; Goodrich & Springer 1992; Goodrich McHugh,]. V. (1993 b): First records ofparasiroids for 2002). Anchorius also includes undescribed spe­ slime mold beetles ofthefamily Sphindidae (Cole­ cies from Central and South America. In the Neo­ optera: Cucujoidea). - Entomological News 104(3): tropics, Anobocaelus Sharp contains four described 136-138. species (Schenkling 1934; Blackweldet 1945) and McHugh,J. V., Marshall, C. J. & Fawcett, F. L. (1997): an indigenous unnamed species on the Galapagos Astudy of adult morphology in Megalodacne heros Island ofSanta Cruz (Peck 2006), as well as 39 spe­ (Say) (Coleoptera: Erotylidae). - Transactions ofthe cies of Gonicoelus Sharp (Blackweldet 1945). From American Entomological Society 123: 167-223. 54 species known throughout the Neotropics, McHugh,J. V. & Lewis, C. N. (2000): Three new species no overlap of species occurs between Central and of carinisphindus McHugh (Coleoptera: Sphindi­ South Ametica (Blackwelder 1945). dae) from Bahamas, Florida, and Puerto Rico. - The caleopteristsBulletin 54: 143-153. Biology and Ecology. Adult and larval biphyl­ McHugh, J. V. & Kiselyova, T. G. (2003): First descrip­ tions for larval stages of Eurysphindus (Coleoptera: lids may be found in leaf litter, fruiting bodies Cucujoidea: 5phindidae). - The Coleopterists Bulletin of pyrenomycetous Ascomycota fungi, and under 57: 17-25. bark of dead trees or fallen branches. Anchorius Russell, L. K. (1979): Beetles associated with slime larvae were reported from under fermenting bark molds (Mycetozoa) in Oregon and California (Cole­ of mesquite (Prosopis sp.) (Lawrence 1991). Diplo­ optera: Leiodidae, Sphindidae, Lathridiidae). - The caelus bmnneus LeConte adults have been collected Pan-Pacific Entomologist 55: 1-9. from dead oaks infested with Hypoxylon fungi Sen Gupta, T. & Crowson, R. A. (1977): The cole­ (Donisthorpe 1935; Lawrence 1977; Crowson opteran family Sphindidae. - The Entomologist's 1981, 1984; Goodrich & Springer 1992; Downie Monthly Magazine 113: 177-19l. & Arnett 1996), and also have been sifted from SlipiIiski, S. A. (1998): Revision and phylogeny ofPro­ dead leaves of maple, beech and other hardwood tocucujidae (Coleoptera, Cucujoidea). - Annales trees. Diplocaelus rudis (LeConte) adults were found Zoologici 48: 275-298. under moist, loose batk of fallen oaks, hickory, Stephenson, S. L., Wheeler, Q D., McHugh, J. V. & and pines (Goodrich & Springer 1992). Some Fraissinet, P. R. (1994): New North American biphyllids, including Anchorius, some Diplocoelus associations of Coleoptera with Myxomycetes. and GonicoelllS in Costa Rica, have been collected -Journal ofNatural History 28: 921-936. 'en masse' at freshly cut stumps of hardwoods as well as on split stalks of palms (Iriartea sp.) spor­ ted with Ascomycota molds (Cline pers. obs.). GonicoelllS larvae also have been collected under bark of hardwoods (Lawrence 1991). Species of Biphyllus have been reported to feed on spores and 10.6. Biphyllidae LeConte, 1861 stromata ofDaldinia (Wollaston 1865; Ganglbauer 1899; Crowson 1955; Vogt 1967; Hingley 1971), Andrew R. Cline and Floyd W. Shockley whereas Diplocoelus have been taken in association with Tubercularia (Palm 1959) and Nammularia (Dajoz 1966). Hammond & Lawrence (1989) also Disttibution. Biphyllidae is cosmopolitan, indicated Biphyllus and Diplocoeills on Xylaria and occurring in all zoogeographic regions except Cryptostroma. Jones (2000) reported Biphylllls lllna­ New Zealand, with highest diversity in the trop­ tus (Fabricius) feeding on a Cryptostroma fungus ics. The family ineludes six genera and approxi­ specific to sycamore trees (Acer sp.) in England. mately 200 species (Lawrence 1982; Goodrich & Some Australian biphyllids have been found on Springer 1992). Schenkling (1934) provided the rotting flower stalks of Xallthorrhoea and rotting most recent world catalogue. In the Old World, cycad cones (Lawrence 1991). Biphyllus Dejean occurs with highest diversity in Africa and Japan (Wollaston 1873; Reittet 1889; Morphology, Adults (Fig. 10.6.1 A-E). Length Miwa 1931) but also occurs in Taiwan, Sumatra, 2.0-6.5 mm. Body oblong to elongate-oval, moder­ Australia (Schenkling 1934) and the Russian Far ately convex to flattened; color yellowish brown to East (Nikitsky 1992). Althaesia Pascoe contains reddish brown to black, typically uniform butocca­ three species from Australia (Blackburn 1894; Lea sionally bicolored; moderately to strongly pubes­ 1921), two from Papua-New Guinea (Pascoe 1860; cent, dorsal surface with erect and decumbent hairs Arrow 1929), and one from Indonesia (Grouvelle in more or less distinct striae, pubescence shorter 1913). DiplocOelllS Guerin-Meneville occurs sporad­ and subdepressed ventrally. ically in the Old World, with most species inhabit­ Head prognathous, visible from above, inserted ing Australia (Lea 1921 a, b, 1922); however, two in prothorax to base of eyes; surface punctate; Biphyllidae LeConte, 1861 307

~.~~·.,J':r U, , 8 1 L~~ t ,,-"".. ,

I 1.- s, _.i- t'd.'R G\;f

Fig.10.6.!. A,Biphyllusjrater Aube, adult, dorsal (modified from Delobel &Tran 1993; © Maurice Tran, ORSTOM); B, Gonicoellls unicornis Sharp, adult, dorsal (modified from Sharp 1902); C, Diplocaelusjagi Guerin-Meneville, adult dorsal (from Hansen 1950, © Danmarks Fauna and Zoological Museum, University of Copenhagen); D, Diplocoe­ Ius punetatlls Lea, adult, dorsal; E, Diplocoelus pllnctatus Lea, adult, ventral, (from Lawrence et al. 1999 a; © CSIRO); F,DipZocoelusamp!icollis Reitter, pupa, dorsal (from Costa etal. 1988; © Museu de Zoologia da Universidade de Sao Paulo, Brazil); G, Diplocoelu5 amplicollis Reitter, larva, dorsal (from Costa etal. 1988; © Museu de Zoologia da Uni­ versidade de Sao Paulo, Brazil); H, Diplocoelusjasciatus, larva, lateral (from Lawrence and Britton 1994, © CSIRO); I,Anchorius lineatus Casey, larva, lateral (from Lawrence 1991; © J. Lawrence). Lines = 1mm.

transverse occipital ridge present. Eyes large, terminal palpomere often securiform or subulate. globose, coarsely faceted, interfacetal setae pres­ Head ventrally with pair of distinct setose tubu­ ent but not elongate. Antennal insertions con­ lar invaginations opening laterally into base of cealed from above; subantennal grooves present snbantennal groove. Gular sutures either broadly between eyes and mandibular bases and extend­ separate or absent. Tentorial arms well-separated, ing behind eyes. Frontoclypeal sntnre absent. Lab­ corpotentorium narrow. rum transverse. Antennae 11-segmented, usually Pronotum transverse, with well-developed lat­ with 3-segmented clnb (2-segmented in Biphyl­ eral margins, sometimes finely crenulate or ser­ Ius). Mandibles moderately large, cnrved and api­ rulate, often with one or two pairs oflongitudinal cally bidentate, mola well-developed. Maxil1a with sublateral carinae (several pairs in Anchorius), and lacinia elongate, three times as long as wide, apex occasionally an additional pair of basal grooves or rounded; long setae present on medial and apical foveae present. Prosternal process parallel-sided. margins; galea wider than lacinia, twice as long Procoxae transversely oval, with concealed or as wide with long setae as in lacinia, apex of galea barely exposed trochantins. Procoxal cavities inter­ densely setose or spinose; maxillary palps 4-seg­ nally and externally closed, with very small lateral men ted, slender. Labium with mentum transverse, extension. Scutellar shield transverse, sides diverg­ trapezoidal; labial palps 3-segmented, slender, ingposteriorly.Elytra 1.2-2.2 times aslongas wide, 308 Andrew R. Cline and Floyd W. Shockley distinctly punctatewith10 rows ofpunctutedsttiae at base. Median endocarina absent. Six stemmata and usually aseutellarystriole. Epipleuragradually on each side. Fronroclypeal suture absent. Labrum narrowed and complete. Mesocoxal cavities ovate, partly fused to head capsule. Antennae shorr bur narrowly separated, open laterally (mesepimeron well-developed, 3-segmented; sensorium on seg­ reaching middle of coxal cavities), rfochantins ment 2 conical or palpiform, much shorter than exposed. Metavenrrite longer than abdominal terminal segment. Mandibles symmetrical, biden­ ventrite 1, slightlyconvex, discrimen present, sinu­ tate, with accessory ventral process and sometimes ate postcoxallines forming an axillaryspace on each with serrate incisor edge; mola well-developed and side (sometimes with an additional pair ofstraight sickle-shaped, transversely ridged, with asetose posrcoxal lines). Meracoxae transverse, slightly hyaline lobe at base; prostheca bearing a brush of grooved, well-separared. Metendosternite of typi­ complex comb-like. Ventral mourhparrs rerracred. cal cucujoid type with elongate lateral arms; ven­ Maxillary arriculating area well-developed. Max­ trolateral and anterior processes reduced or absent, illa with transverse cardo, elongate stipes, 3­ laminae well-developed. Hindwings well-devel­ segmented palps, and falciform mala bearing a oped; apical field with transverse linearsclerirejusr dorsal row of spines along inner edge. Labium distad ofradial cell, which is well-developed; cross­ free almost to base of mentum; ligula transverse; vein R3 oblique, R4 complete, sometimes with labial palps 2-segmenred, widely separated. Pre­ apical spur; rom loop moderarely broad; basal por­ mentum present, postmentum quadrate to ttap­ rion ofRP very short or absenr; medial spur almost ezoidal. Hypopharyngeal sclerome consisting of a reaching wing margin where there is aweakembay­ transverse bar. Hypostomal rods short and diverg­ menr; medial field wi th three or four free veins, the ing. Ventral epicranial ridges well-developed; gula anreriormost (MP,...J extending inro medial fleck; transverse. wedge cell presenr or absenr; anal lobe well-devel­ Legs well-developed, 5-segmented, nearly con­ oped, separared by deep, norch-like embaymenr. tiguous medially; clothed with short spines; pre­ Trochanters short, somewhat heteromeroid in tarsus with 1 seta. Mesotergum, metatergum, appearance; femora slender to moderately robust; and abdominal rerga I-VIII each with a transverse tibiae slender, broadening apically; protibiae usu­ carina near anterior edge. Tergum IX usually with­ ally with spines or spurs apically; tarsal formula our urogomphi, if present short and fixed; sternum 5-5-5, tarsomeres slender, 2-4 with slender IX well-developed, simple. Segmenr X more or less pubescent lobes, penultimate reduced, fifth as long cylindrical, forming a posteroventral pygopod. or longer than other tarsomeres combined. Claws Spiracles annular-biforous, raised on shorr nlbes; simple. posreriorly or posterodorsally placed on segment Abdomen with 5 free ventrites; ventrite 1 lon­ VIIl. [Costa IT af. 1988; Lawrence & Britron 1994; ger than second, with one or two pairs of straight, Lawrence 1991; Lawrence et al. 1999 b; Goodrich diverging postcoxal lines and narrowly rounded 2002.] to acute intercoxal process. Sternites VllI and IX in male with anterior struts. Aedeagus symmetri­ Morphology, Pupae (Fig. 10.6.1 F). Length cal. Anterior edge of tegmen wirh single median 2.0-4.0 mm. Adecricous and exarate. Color from strut and opposing paired srrurs. Parameres fused yellowish brown to light brown; moderately to phallobase, sometimes free from each other. pubescent with long erect hairs often associ­ Female sternite VIII with short spiculum ventrale. ared with distinct spicules. Head partially visible Ovipositor 3-6 times as long as wide, and almost from above with several minute spinose projec­ as long as ventrites 3-5 combined; gonocox­ tions. Pronotum transverse, somewhat to highly ites cylindrical; lengrh of combined gonocoxires convex; posterior angles prominent, apex shorter than paraprocrs, proximal lobe of gona­ rounded; dorsal surface with several to many large coxite and broad, distal lobe long and narrow; sub­ spicules with or wi thout associated setae. Meso­ apical gonostyli well-developed, wirh long setae and metatergum each with four pairs of promi­ subequal in length to srylus. Spermatheca sclero­ nent setose spicules, often arranged in transverse tized and with associated gland. [Lawrence 1999 a; bands. Wing pads long and wrapping around Lawrence & Britton 1994; Goodrich & Springer abdomen to ventral midline. Abdominal seg­ 1992; Goodrich 2002.] ments I-VIII with two pairs of setose spicules dor­ sal and two pairs lateral; segment IX with a pair Morphology, Larvae (Fig. 10.6.1 G-I). Length of distinct, minute urogomphi and two to four 2.5-10 mm. Body elongate, parallel-sided,cylindri­ pairs of tubercles laterad of urogomphi, one pair cal ro slightly flattened, straight or slightly curved distinctly ventral; segment I with large dorso­ ventrally; dorsal surface sclerotized and lightly to lateral annular spiracles; spiracles on segments moderately pigmented, smooth, with scattered, II-VI smaller than rhose on segment I [Costa et al. simple setae; no asperites present, but protergum 19881· with one or more sclerotized plates. Head protracred and prognarhous, moderarely Phylogeny and Classification. Although biphyl­ broad, slightly flattened. Epicranial stem absent lids were always considered to be a well-defined or very short; frontal arms lyriform, contiguous group, sister-group relationships and placement 8iphyllidae LeConte, '86' 309 within Cucujoidea were historically contentious. Acknowledgements Early 19th and 20th century coleopterists placed biphyllids within Mycetophagidae (LeConte 1861, We thank Danmarks Fauna and Zoological 1863; Reitter 1877; Horn 1878; LeConte & Horn Museum, University of Copenhagen, the Com­ 1883), Cryptophagidae (Redtenbacher 1858; Jac­ monwealth Scielllific and Industrial Research quelin DuVal 1859; Wollaston 1862; Thomson Organization (CSIRO), the Museu de Zoologia da 1863; Seidlitz 1891; Casey 1900; Reitter 1909; Universidade de Sao Paolo, and John Lawrence for Leng 1920) and Erotylidae (Ganglbauer 1899; use of copyrighted illustrations. Special thanks to Winkler 1924; Robetts 1958; Crowson 1955; Vogt John Lawrence for providing informarion on fig­ 1967). 1n the mid-20th century, biphyllids again ure sources and contacts and to John Lawrence and were erroneously placed in Tenebrionoidea (Crow­ Richard Leschen for manuscript suggestions. Joe son 1960; Abdullah & Abdullah 1966; Abdullah McHugh (Department of Entomology, University 1973) based on superficial aedeagal and trochan­ of Georgia) provided logistical suppon to FWS [cral similarities; despite the non-hereromeroid through an NSF/pEET grant (DEB-03291l5) to J. tarsi and work by Falcoz (1926) that used adult McHugh, M. Whiting and K. Miller. Drs. Alexey characters and proposed an affinity with Bytllrlls Tishechkin and Martin Hauser aided in Russian (Byturidae). Lawrence and Newton (1982) noted and German translations, respectively. the biphyllid aedeagus was more similar to the cleroid sheath type than the tenebrionoid type and suggested a relationship to languriine Erotylidae Literature (e. g., Cryptophillls). Shared features of larvae and adulrs also have indicated potential relationships Abdullah, M. (1973): The improvement of an exist­ ro Bothrideridae, Nitidulidae, Protocucujidae and ing modern classification in biology. - Zoologische euxestine Cerylonidae (Crowson 1955; Lawrence Beitriige 19: 13-41. 1991). Crowson (1981,1984) suggested the unique Abdullah, M. & Abdullah, A. (1966): Byturidae and biphyllid pre-gular pits might be a type of mycan­ Biphyllidae (Coleoptera), two primitive families of gium; however, no evidence suppons this claim. the Hcreromera not the Clavicornia - a new inter­ pretation ofsome old observations. - Entomological The pits do appear to present a potential synapo­ News 77(3): 63-69. morphy for Biphyllidae, in combination with the Arrow, G. J. (1929): On the families of Coleoptera meraventral and/or abdominal coxal lines, uniting related to the Erotylidac, with descriptions of a members of this family. new family, two new genera, and a few new spe­ Some family-level taxonomic confusion sur­ cies. - Allnals and MagazineofNatllral History 4 (10): rounded this group until the late 20th century. 305-322. Lawrence and Newton (1995) noted the incor­ Barber, H. S.(1942): Raspberry fruirwormsand related rect family-group author assignment used in species. - Miscellaneous Publications a/the u.s. Depart­ Silfverberg (1992) and that Goodrich & Springer mentofAgriCllltllre468: 1-32. (1992) had refetted to Biphyllidae Sharp (1900) Blackburn, C. T. (1894): Further notes on Australian in which Sharp apparently assumed Biphylllls Coleoptera with descriptions of new genera and Dejean and Dipilylllls Berthold were independent. species. - Transactions a/the Royal Society ojSouthAlts­ LeConte (1861) used the genus Diphylllls to estab­ tralia (Partxvt) 18: 200-240. lish rhe family-group name, but Diphylllls was an Blackwelder, R. E. (1945): Checklist of the coleopter­ unjustified emendation of Biphylllls by Redren­ ous insects of Mexico, Central America, the West bacher (1858). Under rule of priority, correction Indies, and South America. Pan 3. - Bulletin. ofthe of this emendation and subsequent relegation of U.S. National Mllsellm 185: 343-550. Diphylllls to junior synonym made LeConte's fam­ Brirton, E. B. (1970): Coleoptera (beetles). Pp. 495­ ily-group name valid under the cortected name 621 in C.S.I.R.O. Division of Entomology (ed.) Tile Biphyllidae. Current consensus places Biphyllidae Insects ofAustralia: aTextbookfor Students alld Research within Cucujoidea (Crowson 1955), sister to Byru­ Workers. Melbourne University Press, Carlton ridae (Falcoz 1926; Barber 1942; Crowson 1955; (Victoria). Casey, T. 1. (1900): Review of the American Goodrich & Springer 1992). Leschen et al. (2005) Corylophidae, Cryptophagidae, Tritomidae and recently performed a robust cladistic analysis of Dermestidae, with other studies. - Journal of tile basal cucujoid families based on adult and latval New York Entomological Society 8: 51-172. morphology that, like the above previous efforts, Costa, C., Vallill, S. A. & Casari-Chen, S. A. (1988): lar­ yielded conflicting results. Adult morphology sup­ vas de Coleoptera do Brasil. 282 pp. Museu de ZooID­ ported a sister group relationship to Erotylidae, gia, Universidad de Sao Paulo, Sao Paulo. but larval morphology supported a relationship to Crowson, R. A. (1955): Tile Natural Classification ofthe Byturidae. A combined molecularand morpholog­ Families of Coleoptera. 187 pp. Nathaniel Lloyd, ical dataset may be required to resolve the proper London. placement of this family within Cucujoidea. No - (1960): The phylogeny of the Coleoptera. - Annllal phylogenetic srudies have yet been perfotmed Review ofEntomology 5: 111-134. to assess intra-familial relationships at any level - (1981): The Biology of tile Coleoptera. 802 pp. Aca­ within Biphyllidae. demic Press, London. 310 Andrew R. Cline and Floyd W. Shockley

- (1984): The associations of Coleoptera with Asco­ Entomologist's Records and Joumal of Variation 11Z: mycetes. Pp. 256-285. in Wheeler, Q & Blackwell, 177-178. M. (cds.) Fungus Insect Relationships: Perspectives in Lawrence. J. F. (1977): Coleoptera associated with a Ecology and Evolution. Columbia University Press, Hypoxyfon species on oak. - The Coleopterists Bulfetin New York. 31: 309-312. Dajoz, R. (1966): Ecologie er biologie des coJeopteres - (1982): Coleoptera. Pp. 482-553 in Parker, S. P. dela Hetraie. - VieetMilieu (C) 17: 525-763. (ed.) Synopsis and Classificalioll of Livillg Organisms, Delobel, A. & Tran, M. (1993): Les CoJeopteres des Vol. 2. McGraw-HilI. New York. denrees Alimentaires entreposees dans les Regions - (1991): Coleoprera. Family Biphyllidae (Cucujoi­ Chaudes. Faune Tropicale xxxn. 424 pp. Orstom deal. Pp. 475-476 in Stehr, F. W. (ed.) Immature Editions, Paris. Insects. Vol. 2. Kendall/Hunt. Dubuque. Iowa. Donisthorpc, H. 1935. The British fungicolous Lawrence, J. F. & Britton, E. B. (1994): Australian Coleoprera. - Entomologists Monthly Magazine 71: Beetles. 192 pp. Melbourne University Press, 21-31. Melbourne. Downie, N. M. & Arnett, R. H., Jr. (1996): Family 94. Lawrence, J. F. & Newton, A. F.,]r. (1982): Evolution Biphyllidae. The False Skin Beetles. Pp. 1075 in and classification ofbeetles. - Amllial Review oj£col­ Downie, N. M. & Arnert, R. H., Jr. (eds.) The Beetles ogy and Systematics 13: 261-290. of Northeastern North America, Volume II: Polyphaga, - (1995): Families and subfamilies of Coleoptera Superfamily Bostrichoidea through Curculionoidea. The (with selected genera, notes, references and data Sandhill Crane Press, Gainesville. on family-group names). Pp. 779-1006 in Paka­ Falcoz, 1. (1926): Position systematique des genres luI<, J. & Slipiriski, S. A. (eds.) Biology, Phylogeny, Diphyllus Stephens and Diplocoeills Guerin. Pp. and Classification of Coleoptera: Papers Celebrating the 69-74 in Lesne, P. (ed.) Encyclopedie de En tomologiq ue 80th Birthday ofRoy A. Crowson. Museum i Instytut Serie B, Memoires et Notes. I. Coleoptera. Vol. 1. P. Lech­ Zoologii PAN, Warszawa. evalier, Paris. Lawrence,J. F., Hastings, A. M.• Dalwitz. M. J., Paine, Ganglbauer, L. (1899): XXVI. Familie. Erotylidae. T. A. & Zurcher, E. J. (1999 a): Beetles of the World: Pp. 629-739 in Ganglbauer, L. Die Kafer Mitteleuro­ A Key and Information System for Families and Subfami­ pas. Wien, Germany. lies. CD-ROM, Version 1.1 for MS-Windows. CSlRO Goodrich, M. A. (2002): 89. Biphyllidae LeConte Publishing, Melbourne. 1861. Pp. 356-357 in Arnett, R. H., Jr., Thomas, - (1999 b): Beetle Larvae of tI" World: Descriptions, M. C., Skelley, P. E. & Franlr, J. H. (eds.) Ameri­ Illustrations, Identification, and Information can Beetles. Volume 2. Polyphaga: Scarabae­ Retrieval for Families and Subfamilies. CD-ROM, oidea through Curculionoidea. CRC Press. Boca Version 1.0 for MS-Windows. CSIRO Publishing, Raton. Melbourne. Goodrich, M. A. & Springer, C.A. (1992): A Revision of Lea, A. M. (1921 a): On Coleoptera, mostly from the Family Biphyllidae (Coleoptera) for America, Queensland. - Memoirs ojthe Queensland Museum 7: North of Mexico. - The Coleopterists Bulletill 46: 182-240. 361-377. - (1921 b): Descriptions of new species of Australian Grouvelle, A. H. (1913): Descriprion d'un Althaesia Coleoptera. Part xvii. - Proceeding of the Lill11aean ICol. Diphyllini] appartenant au British Museum. Society ofNew South Wales 46: 351-369. - Bulletin de la Societe Entomologique de France 20: - (1922): On Australian Coleoptera. Part IV. Fam­ 497-S00. ily Chrysomelidae. - Records of (he South Australian Hammond, P. M. & Lawrence, J. F. (1989): Appendix: Museum 2: 271-308. mycophagy in insects: a summary. Pp. 275-324 in LeConte, J. L. (1861): Classification of the Coleoptera Wilding. N .• Collins. N. M.• Hammond. P. M. & ofNorrh America. -Smithsonian Miscellaneous Collec­ Webber, J. F. (eds.) Insect- FUllgus Interactions. Aca­ tions 3: 1-208. demic Press. London. - (1863): New Species of North American Coleop­ Hansen, V. (1950): Danmarks Fauna Bd. 55. Biller tera. Part 1. - Smithsonian Miscellaneous Collections 6: XIII. Clavicornia. Vol. 1. 278 pp. Del. G. E. C. Gads 1-92. Forlag. Copenhagen. LeConte, J. L. & Horn, G. H. (I883): Fam. XXVII. Hingley, M. R. (1971): Theascomycete fungus Daldinia Mycerophagidae Pp. 138-40 in LeConte, J. L. & concentrica as a habitat for animals. - Journal ofAni­ Horn, G. H. Classification of Ihe Coleoptera of North mal Ecology 40: 17-32. America. Smithsonian Miscellaneous Collections Horn, G. H. (1878): Synopsis of Mycetophagidae. Pp. 507. Smithsonian Institution, Washington. 603-606 in Hubbard, H. G. & Schwarz, E. A. (eds.) Leng, C. W. (1920): Catalog of tile Coleoptera of The Coleoptera of Micbigan. - Proceedings of the America North ofMexico. 470 pp. John D. Sherman, American Philosophica.l Society 17. Jr., Vermont. Jacquelin DuVal, P. N. C. (1859): Manuel entomologique. Leschen, R. A. B., Lawrence, J. F. & Slipinski, S. A. Genera des Coliopteres d'Europe. Vol. II. 286 pp. Dey­ (2005): Classification of basal Cucujoidea (Cole­ rolle. Paris. optera: Polyphaga): Cladistic analysis, keys and Jones, R. A. (2000): Biphyllus lunatllS (Fabr.) (Col.: review of new families. - InvercebraceSystematics 19: Biphyllidae) and Annomatus duodecimstriatus 17-73. (Muller, P. W. J.) (Col.: Bothrideridae) both feed­ Miwa, Y. (1931): A Systematic Catalogue of Formosan ing on sycamore Acer pseudoplatanus bark. - The Coleoptera. 359 pp. Department of Agriculture, Erotylidae Leach, 181 5 311

Government Research Institute (Report No. 55), 10.7. Erotylidae Leach, 1815 Formosa. Nikitsky, N. B. (1983): Species of the genus Biphyllus Richard A. B. Leschen, Paul E.Skelley and (Coleoptera; Biphyllidae) in the east Palearctic. - Zoologicheskiy Zhumal62: 695-706. Joseph V. McHugh - (1992): Biphyllidae. Pp. 279-284 in Lehr, P. A. (ed.) Identification Key to the Insects of the For East of the USSR. Vol. 3. Part 2. Nauka Pren, St. Petersburg. Distribution. There are approximately 258 pres­ - (1993): New species ofDerodontidae and Biphylli­ ently recognized genera and over 3500 described dae(Coleoptera) from theCaucasus. - ZOologicheskiy species occurring worldwide and present mainly in Zhumal 72 (2): 152-156. forested environments where fungi or host plants Palm, T. (1959): Die Holz-und Rinden-Kafer det siid­ occur. Apartfrom some members ofCryptophilinae und mittelschwedischen Laubbaume. - Opuscula and stored product pests, the family is absent from Entomologica. Supplementum 16, 374 pp. oceanic islands. The group is divided into six sub­ Pascoe, F. P. (1860): Notices of new or little known families and ten tribes. The subfanlily Xenoscelinae genera and species of Coleoptera. - Journal a/Ento­ (8 genera, 10 spp.) is widespread, but absent from mology 1: 98-132. temperate South America and the Nearctic. The Peck, S. B. (2006): The Beetles of the Galapagos Islands, subfamily Pharaxonothinae (5 genera, 24 described Ecuador. Evolution, Ecology, and Diversity (Insecta: Cole­ species) is widely distributed with genera that are optera). 313 pp. NRC Research Press, Ottawa. restricted to Japan and southern Europe and the Redtenbacher,1. (1858): Fa unaAustriaca. Die Kafer. Nach genus Pharaxollotha Reitter found in the tropi­ der analytischea Methode bearbeitet. Zweite Auftage. cal New World and Asia and possibly Africa. Two 1017 pp. C. Gerolds Sohn, Wein. species, Leucohimatium arundinaceum (Forskal) and Reitter, E. (1877): Coleoprorum species novae. - Ver­ Phartvronotha kirschi Reitter, are widespread stored hondlungen der k. k. Zoologisch-Botanischen Gesellschaft product pests and there are many undescribed spe­ in Wien 27: 165-194. cies in the largest genus Phoraxollotha (13 spp.). The - (1909): Cryptophagidae Pp. 54--73 in Reitter, E. subfamilyLoberinae(5 genera, 79spp.) is widely dis­ Fauna Gennanjca. Vol. m. K. G. Lutz Verlag, Stutt­ tributed with genera restricted to Madagascar, New gart, Germany. Roberts, A. W. R. (1958): On the taxonomy ofEroryli­ Caledonia, and Central America. The world wide dae (Coleoptera). with special reference to the mor­ genus LoberltS LeConte (the largest loberine genus phological characters of the larvae. - Transactions of with 75 spp.) and the New Caledonian genus Paphe­ theRayalSocietyofLondOllllO: 245-285. zia Zablotny & Leschen (1 sp.) contain many unde­ Schenkling, S. (1934): Coleopterorum Catalogus. Pars scribed species. One ofthe largest and most diverse 133. Biphyllidae. 7 pp. W. Junk, Berlin. subfamilies is the phytophagous Languriinae with Seidlirz, G. (1891): FOlino Baltica, Die Kdfer (Coleop­ three tribes. Hapalipini (3 gen., 74 spp.) is predomi­ tera) der Deutschen Ostseeprovinzen. Russlands. 818 nantly pantropical, wirh HapalipsReitter the largest pp. Hattungsche Verlagsdrueckerei, Konigsberg genus (57 spp.). Languriini (56 genera, >750 spp.) (Germany). is mainly tropical, and absent from Europe, New Sharp, D. (1902): Fam. Cryptophagidae. Pp. 579-624, Zealand, and Chile. Thalisellini (4 genera, 25 spp.) PI. 18 in Insecta, Coleoptera. Vol. 2 (1). Biologia is Neotropical and contains many undescribed spp. Cmtrali-Americano. Vol. 2 (1). Taylor and Francis, Thesubfamily Cryptophilinaecontains three tribes. London. Empocryptini (3 genera, 17 spp.) occurs in the SiIfverberg, H. (1992): Enumeratio Coleopterorum Neotropics and temperate South America. Crypto­ Fennoscandiae, Danieae et Baltiae. 94 pp. Helsin­ philini (5 genera, 33 spp.) is mainly pantropical, but gin Hyontcisvaihtoyhdiscys, Helsinki. the widespread genus Cryptophilus Reitter has some Thomson, C. G. (1863): Skandinaviens Coleoptera, Holarctic species and Cryptophilus integer (Heer) is a synopiskt bearbetade. Volume 5. 340 pp. Tryckt widespread vagrant. Toramini (5 genera, 58 spp.) uti Lundbergska Boktrycheriet, Lund. is widespread, mainly tropical with few species in Vogt, H. (1967): 54. Familie: Erorylidae. Pp. 104--109 temperate regions, and absent from much of the in Freude, H., Harde, K. W. & Lohse, G. A. Die Kafer Pacific, Australia, and New Zealand. The largest Mitteleuropas. Band 7. Clavicornia. Goeke & Evers, and most widespread genus is Toramus(43 spp). The Kerfeld. Winkler, A. (1924): Erotylidae. Pp. 715-720 in Win­ mycophagous subfamily Erorylinae is the largest kler, A. (ed.) Cotaloglls Coleopterorum Regionis Palearc­ consisting of five tribes, 172 genera, and 2563 spe­ ticae. Volume 1. cies. Dacnini (14 genera, 92 spp.) has a worldwide Wollaston, T. V. (1862): On the Euphorbia-infesting distribution, occurs primarily in rhe Holarctic and Coleoptera of the Canary Islands. - Transactions of Austral-Asian regions, with few members from the the Royal Entomological Society ofLondon (series 3) 1: New World and African tropics. Encaustini (13 gen­ 136-214. era, 203 spp.) is pantropical and occurs primarily in - (1865): Coleoptera Atlantidllm, Being an Enumeration southeastern Asia-Indonesia, with few African and ofthe Coleopterous Insects ofthe Madeiras, Salvages, and Central American taxa. Megalodacnini (sometimes Callaries. 526 pp + 140 pp Appendix. Taylor and considered part of Encaustini by W~grzynowicz Francis Publishers, London. 2002) (28 geneta, 323 spp.) is distributed worldwide