10.28. bothrideridae erichson, 1845

Hetschko, A. (1930): Pars 109. Cucujidae, Thorictidae (Supplement). Pp. 1–122 In Schenkling, S. (ed.). Coleopterum Catalogus. W. Junk, Berlin.

Horn, G. H. (1879): Revision of the Nitidulidae of the United States. – Transactions of the American Entomo-logical Society 7: 267–336.

Kirejtshuk, A. G. (1982): Systematic position of the genus Calonecrus J. Thomson and notes on the phy-logeny of the family Nitidulidae (Coleoptera). Ento-mologicheskoye Obozrenie 59: 833–851.

– (1998): Nitidulidae (Coleoptera) of the Himalayas and northern Indochina, Part 1: Subfamily Epu-raeinae, Theses Zoologicae, Vol. 28. Koeltz Scien-tifi c Books, Königstein. 489 pp.

Lawrence, J. F. (1982): Coleoptera. Pp. 482–553 in Parker, S. P. (ed.): Synopsis and Classifi cation of Living Organisms, Volume 2 McGraw-Hill, New York.

– (1991): Nitidulidae (Cucujoidea) (including Bra-chypteridae, Cateretidae, Cybocephalidae, Smicrip-idae): sap beetles, dried fruit beetles [pp. 456–460]. In: Immature Insects, Volume 2 (F. W. Stehr, editor). Kendall Hunt Pub. Co. Dubuque, Iowa, 975 pp.

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

– (1999 b): “Beetle Larvae of the World: Descriptions, Illus-trations, Identifi cation, and Information Retrieval for Families and Subfamilies.” CD-ROM, Version 1.0 for MS-Windows, CSIRO Publishing, Melbourne.

Lawrence, J. F. & Newton, A. F., Jr. (1995): Families and subfamilies of Coleoptera (with selected gen-era, notes, references and data on family-group names. Pp. 779–1006 in Pakaluk, J. & Slipinski, S. A. (eds.) Biology, Phylogeny, and Classifi cation of Coleop-tera: Papers celebrating the 80th Birthday of Roy A. Crow-son. Muzeum i Instytut Zoologii PAN, Warszawa.

LeConte, J. L. (1878): Additional descriptions of new species. – Proceedings of the American Philosophical Society 17: 373–434.

LeConte, J. L. & Horn, G. H. (1883): Classifi cation of the Coleoptera of North America. – Smithsonian Miscellaneous Collections 26: 1–567.

Leng, C. W. (1920): Catalogue of the Coleoptera of America, north of Mexico. 470 pp. John D. Sher-man Jr. Publisher, N.Y.

Leng, C. W. & Mutchler, A. J. (1933): Second and third supplements 1925–1932 (inclusive) to Catalogue of the Coleoptera of America, north of Mexico. 112 pp. John D. Sherman Jr. Publisher, N.Y.

Leschen, R. A. B., Lawrence, J. F. & Slipinski, S. A. (2005): Classifi cation of basal Cucujoidea (Cole-optera: Polyphaga): Cladistic analysis, keys and review of new families. – Invertebrate Systematics 19: 17–73.

Murray, A. (1864): Monograph of the family Nitidu-lariae. – Transactions of the Linnaean Society of London 24: 211–439.

Pakaluk, J., Slipinski, S. A. & Lawrence, J. F. (1995): Current classifi cation and family group names in Cucujoidea (Coleoptera). – Genus 5: 223–268.

Parsons, C. T. (1943): A revision of the Nearctic Niti-dulidae (Coleoptera). – Bulletin of the Museum of Comparative Zoology 92: 121–278.

Peck, S. B. (2005): A checklist of the beetles of Cuba with data on distributions and bionomics (Insecta: Coleoptera). Volume 18. Arthropods of Florida and neighbor-ing land areas. 241 pp. Florida Department of Agriculture and Consumer Services. Gainesville, Fl. Peck, S. B. & Thomas, M. C. (1998): A distribu-tional checklist of the beetles (Coleoptera) of Florida. Volume 16. Arthropods of Florida and neighboring land areas. 180 pp. Florida Department of Agriculture and Consumer Services, Gainesville, Fla.

Price, M. B. (2002): Smicripidae Horn 1879. Pp. 316–318 in Arnett, R. H., Thomas, M. C., Skelley, P. E. & Frank, J. H. (eds.) American Beetles. Volume 2. Polyph-aga: Scarabaeoidea through Curculionoidea. CRC Press, Boca Raton, Florida.

Reitter, E. (1876): Revision der Monotomidae (sensu LeConte). – Deutsche Entomologische Zeitschrift 20: 295–301.

Sharp, D. (1900): Monotomidae. Pp. 563–579 in: Godman, F. D. & Salvin, O. (eds.) Biologia Centrali-Americana. Insecta, Coleoptera. Volume II. Part 1. Porter, London.

10.28. Bothrideridae Erichson, 1845

Adam Slipinski, Nathan Lord and John F. Lawrence

Distribution. Bothrideridae are found in all major zoogeographical regions of the world. About 400 species are placed in 38 genera and four subfamilies according to a classifi cation by Pal & Lawrence (1986) with modifi cations by Slipinski (2007). Xylariophilinae (Xylariophilus Pal & Law-rence) is widespread throughout the Oriental and Australian regions with several undescribed spe-cies found on the Pacifi c islands. Within Teredi-nae, the tribe Teredini is known mostly from the Palaearctic (Teredus Shuckard, Oxylaemus Erichson, Teredolaemus Sharp), Nearctic (Oxylaemus, Rustleria Stephan) and Afrotropical (Teredolaemus, Teredo-morphus Heinze) regions with few Teredolaemus species known from Madagascar, India, Indonesia and Australia. The teredine tribe Sysolini com-prises the single genus Sysolus Grouvelle with two described species from India and Vietnam, but unnamed species are known from New Guinea, Fiji, Panama and Mexico. The endemic and prac-tically unknown Malagasy Sosylopsini (=Sosylopsis Grouvelle) is also classifi ed in this group (Dajoz 1977). Anommatinae consists of three genera and about 110 species widely spread throughout Europe and northern Africa (Dajoz 1965, 1968, 1969, 1977). A. duodecimstriatus (Müller) has been introduced into North and South America, New Zealand (Kuschel 1979), Australia (Lea 1904, 1905) and Chile, and there is a single record of A. vallombrosae Dieck occurring in North America (Nichols & Wheeler 1984). The genus Deretaphrus

Bothrideridae Erichson, 1845 411

Newman is represented by a single Nearctic spe-cies, D. oregonensis Horn, several endemic species in Australia, one in New Caledonia, and one in Bolivia (Lord 2008). Bothriderinae is a large cos-mopolitan group (27 genera, about 270 described species), but the majority of the diversity is found in the tropics (Slipinski et al. 1989).

In the New World, little taxonomic work has been accomplished except for Stephan (1989) and Philips & Ivie (2002), covering the eight genera and 18 species found in the United States and Canada. North American Bothrideridae are clearly Holarc-tic, sharing most genera with the Palaearctic region. Two South American genera (Lithophorus Sharp, Pro-lyctus Zimmerman) reach the Southwestern United States. Deretaphrus, with one species occurring in the Western United States, is the only genus shared exclusively with Australia, New Caledonia, and Bolivia.

The Mesoamerican and South American both-riderid fauna is relatively depauperate, including no native Teredinae, Xylariophilinae, or Anom-matinae, with Bothriderinae being represented by four genera only with Prolyctus and Lithophorus endemic to the New World. Also, this fauna has not been studied since Hinton’s (1935, 1936) addition of several new Sosylus and Bothrideres Dejean species to the classical treatment of Colydiidae by Sharp (1894 –95) in the Biologia Centrali Americana.

Old World Bothrideridae are distinctly more diverse with representatives from 33 of the 38 cur-rently recognized genera. Within Bothriderinae, 23 of 27 genera and about 180 of the 270 species are represented. The family is poorly represented in the Palaearctic region except for Anommatinae, all 90 native species of which are confi ned to the Old World. All genera are Holarctic or broader in distribution except for Teredus, distributed only in Europe and northern Africa (Dajoz 1977; Nikitsky 1985; Slipinski 2007). Four large genera, Teredol-aemus, Pseudobothrideres Grouvelle, Ascetoderes Pope and the subcosmopolitan Sosylus, which contain jointly about 145 species, dominate the Afrotropi-cal, Oriental and Australian faunae (Heinze 1943; Pope 1961; Slipinski et al. 1989). Madagascar has a very similar generic composition of Bothrideridae to Africa and both share all genera except for the enigmatic Sosylopsis Grouvelle (Dajoz 1980). The most peculiar element of Australian Bothrideridae (Carter & Zeck 1937; Pal & Lawrence 1986; Lord 2008) is the speciose genus Deretaphrus, which is widely spread in the dry sclerophyll zone. Other-wise most of the genera are clearly of Indo-Pacifi c or broader tropical origin.

Biology and Ecology. All Bothrideridae except for Anommatinae are found on or under the bark of trees and are usually associated with wood bor-ing insects. Anommatinae are found in compost, forest litter, grass cuttings and the in upper stra-tum of organic rich soil. Anommatus larvae have been found in the same habitat but their biology is

unknown. The few known larvae of Xylariophili-nae and Teredinae are free living and apparently mycophagous. Xylariophilines feed nocturnally on the surfaces of various Xylariaceae (Ascomycetes), while larvae of Teredolaemus leai (Grouvelle) (Teredi-nae) are known to be commensals in tunnels of ambrosia beetles (Curculionidae: Platypodinae) on Nothofagus. Larvae of this species do not construct cocoons for pupation. The larvae of the Palaearctic Teredus are undescribed, but adults and putative larvae have been collected from galleries of various wood boring beetles belonging to the families Ano-biidae, Cerambycidae and Curculionidae (Scolyti-nae) (Horion 1961).

All known larvae of Bothriderinae are hyper-metamorphic with an active triungulin stage that seeks out and attaches itself to its host. The later instars are grub-like and modifi ed as ectoparasit-oids of larvae and pupae of wood boring beetles and hymenopteran wood wasps (Xiphydria sp.) and carpenter bees (Xylocopa appendiculata (Piel 1938)). Two distinct biological groups are present among the Bothriderinae that utilize different hosts and apply slightly different strategies. Adults of various Sosylus are found in galleries of numerous Platypo-dinae (Curculionidae) (Browne 1962; Roberts 1969, 1980) where they prey on both adults and larvae. The fi rst instar larva of Sosylus is well sclerotized, fl attened and distinctly spinose with relatively

Fig. 10.28.1. Ascetoderes taeniatus Pascoe, adult, dorsal (from Britton 1970, as Bothrideres equinus Pascoe; © CSIRO Australia), length = 5 mm.

412 Adam Slipinski, Nathan Lord and John F. Lawrence

Fig. 10.28.2. Bothrideridae adults, dorsal: A, Sosylus sp., length = 4.6 mm; B, Prolyctus n. sp., length = 4.4 mm; C, Deretaphrus oregonensis Horn, length = 11.3 mm; D, Oxylaemus californicus Crotch, length = 3.8 mm; E, Deretaphrus fossus Newman, length = 10.5 mm; F, Bothrideres geminatus Say, length = 3.0 mm; G, Teredolaemus leae (Grouvelle), length = 2.8 mm; H, Ascetoderes musivus Pascoe, length = 3.3 mm; I, Xylariophilus bicoloripennis Pal & Lawrence, length = 1.7 mm; J, Teredolaemus setipennis Pope, length = 2.8 mm; K, Dastarcus kurosawai Sasaji, length = 6.1 mm; L, Antibothrus major Slipinski, length = 6.3 mm. (© Nathan Lord.)


larger legs. Upon fi nding the pupal stage host, it attaches itself on its ventral side of the abdomen near segment II and starts feeding. After subse-quent moulting, the larva constructs a waxy cell in which pupation occurs. Asosylus rhysodoides (Pascoe) behaves similarly, but it attacks pupae of a bren-tid host Carcinopisthius kolbei (Senna) that is a nest parasite of Platypus jansoni (Chapuis). Fragmented observations on biology of Deretaphrini and Both-riderini (Burke 1919; Craighead 1920; Lieu 1944;

Negru 1954; Svácha 1994) suggest that the larvae utilize a wider range of hosts compared to the pre-vious group. Svácha (1994; personal communica-tion) found the fi rst instar larvae of the European species Bothrideres bipunctatus (Gmelin) sucking on the surface of dead/dying larvae of the siricoid hymenopteran larvae of Xiphydria Latreille, and, in other instances, on soft xylophagous coleopteran larvae of the families Cerambycidae, Buprestidae, Curculionidae and Anobiidae. Later instars of

Fig. 10.28.3. Bothrideridae adults, dorsal: A, Ascetoderes signatus Grouvelle, length = 5.0 mm; B, Asosylus rhysodoides (Pascoe), length = 5.6 mm; C, Lithophorus succineus (Pascoe), length = 4.7 mm; D, Teredolaemus sp., length = 4.1 mm; E, Ogmoderes sculpticollis (Thomson), length = 6.9 mm; F, Roplyctus minutus Pope, length = 2.2 mm. (© Nathan Lord.)


Bothrideres, Ascetoderes, Deretaphrus and Dastarcus Walker have been observed feeding externally on various larvae of wood inhabiting beetles, Hymenop-tera and Lepidoptera. Full grown larvae spin waxy or silken cocoons for pupation (Figs. 10.28.7 B, E). It is not clear from published records whether or not there is an endoparasitic stage or how the larva reaches the host. Stephan (1989) found sev-eral partially empty cocoons clustered together (Fig. 10.28.7 A) and adults of Bothrideres cryptus Stephan nearby in an empty cerambycid pupal chamber. This points to the possibility that sev-eral parasites may be associated with a single, large host.

Morphology, Adults (Figs. 10.28.1–4, 5 A–C, H–J,6 A–B). Length 1.4–12.0 mm. Body usually dis-tinctly elongate, 2.5 to more than 4 times as long as wide (somewhat shorter and broader in Dastarcus and relatives, less than twice as long as wide in some Xylariophilus), cylindrical or slightly fl at-tened. Colour yellow to black, rarely with irregular pattern (Xylariophilus, Teredolaemus), red apices or spots (Teredolaemus). Surface glabrous, clothed with fi ne hairs (Teredini, Bothrideres) or scales (Dastarcus, Emmaglaeus Fairmaire).

Head slightly declined, not abruptly constricted posteriorly, or constricted by a weak lateral post-ocular occipital impression (Deretaphrus). Occipital region without stridulatory fi le. Eyes entire (absent in Anommatinae), strongly protuberant; facets variable, without or with very short interfacetal hairs. Antennal insertions exposed from above;

subantennal grooves between eye and mandibular articulation usually present, rarely extending beyond posterior edge of eye. Frontoclypeal suture distinct, arcuate. Labrum lightly sclerotized, transverse, with sclerotized rods; anterior margin straight or with slight emargination (Sosylopsis). Antenna 9- to 11-segmented, usually with compact club composed of 1–3 segments (club relatively loose in Sysolus, Sosylopsis, and less distinct in Deretaphrus). Mandible bidentate, often with well-developed ventral subapical tooth; incisor edge simple; mola well-developed and transversely ridged in most of the groups, rarely reduced and replaced by a fl eshy lobe (some Bothriderini, e. g., Dastarcus); prostheca with well-developed setal fringe. Galea broad, setose; lacinia long with one or two apical spines. Maxillary palps short, apical palpomere fusiform. Ligula slightly expanded and truncate; apical labial palpomere fusiform or rarely shorter and apically truncate. Gular sutures widely separated and short (Teredini, Anommatini, Xylariophilini), strongly convergent or anteriorly fused (Bothriderinae), or externally absent (Sysolini). Tentorium absent in many Bothriderinae. Corporotentorium if present narrow, with (Xylariophilus, Teredus, Sosylopsis) or without (Anommatus Wesmael, Sysolus) median pro-cess. Cervical sclerites reduced and hardly visible.

Pronotum about 0.7–1.2 times as long as wide; base usually as broad as elytral bases; sides not explanate; lateral pronotal carinae complete or incomplete (Deretaphrus); disc variable, smooth, with median groove (Asosylus Grouvelle, Craspe-dophilus Heinze, Deretaphrus), two to four carinae (Machlotes Pascoe, Erotylathris Motschulsky), paired

Fig. 10.28.4. Bothrideridae adults, dorsal: A, Emmaglaeus raffrayi Grouvelle, length = 5.6 mm; B, Sysolus antennatus Grouvelle, length = 4.2 mm; C, Anommatus duodecimstriatus (Müller), length = 1.5 mm.


basal impressions (Oxylaemus); central island with complex channels (Pseudobothrideres Grouvelle) and rarely a complex system of carinae and transverse grooves (e. g., Cosmothorax Kraatz). Prosternum in front of coxae at least two times as long as short-est diameter of coxal cavity. Prosternal process varies from very narrow (Teredini, Xylariophilini, Deretephrus, Sysolus) to very broad in many Both-riderinae. Prosternal process always complete, par-allel-sided or weakly to strongly expanded apically. Notosternal sutures complete, rarely absent (Dere-taphrus). Procoxa not or weakly projecting below prosternum, with well-developed internal exten-sions. Trochantins not visible externally. Procoxal cavities rounded, variably separated, externally open (Teredini, Xylariophilini), narrowly closed (Sosylopsis, Teredus, Bothrideres), or broadly closed (most Bothriderinae, Deretaphrus); without lateral extensions; internally open, or closed by a slen-der bar (Sosylopsis). Scutellar shield not abruptly elevated, anteriorly simple. Elytra elongate; dor-sal surface with ten variously marked puncture rows; scutellary striole absent; elytral surfaces smooth (most of Teredini, Xylariophilini), with apical declivity (some Teredolaemus, Sysolus, Sosylus) or variously developed costae; apices meeting at the suture; epipleura narrow, incomplete at very apex; often with closing mechanism composed of outer elytral fl ange fi tting in expanded margin of last abdominal ventrite. Mesoventrite separated by complete sutures from mesanepisterna, which are broadly separated at midline; anterior edge on same plane as metaventrite, without distinct procoxal rests. Mesocoxa not projecting; contiguous or very narrowly separated (Teredini, Sosylopsini, Dere-taphrini and Xylariophilini), or narrowly to broadly separated (Bothriderinae, Sosylopsis). Trochantins hidden, rarely visible (Sysolus). Mesocoxal cavities circular, closed or very narrowly open laterally (Sysolus); external closure apparently incomplete in some Xylariophilus; mesepimeron small, transverse, often strongly reduced. Mesometaventral junction simple, with single metasternal knob or incom-plete (Anommatus, several Teredini). Metaventrite often with short discrimen (absent in Sosylopsis); postcoxal lines usually absent, rarely present (some Bothriderinae, e. g., Pseudobothrideres); exposed por-tion of metanepisternum very long and narrow; metepimeron visible externally near metacoxae or strongly reduced, fused or covered from below by metanepisternum (Bothriderinae). Metacoxae transversely oval, narrowly or broadly separated. Metendosternite with short broad stalk (narrow in Deretaphrus), long lateral arms, laminae well-devel-oped but narrow (absent in Sosylopsis), anterior tendons broadly separated and located near base of laminae. Hind wings absent in Anommatinae; if present without radial cell (except for Dereteph-rus Fig. 10.28.7 B); weak medial embayment pres-ent; medial fi eld with fi ve free veins (Deretaphrus), three to four (Teredini, Sysolini, Bothriderinae), or one (Xylariophilus, Sosylopsis); divided medial fl eck

present in Teredini, Sysolini and Xylariophilini, absent in Sosylopsis; anal embayment (jugal lobe) usually present, deep and notch-like (absent in Sos-ylopsis). Legs well-developed; trochanterofemoral joint strongly oblique (Teredinae, Anommatinae), truncate (Sosylopsis), or completely internalized and externally invisible (Bothriderinae); tibial api-ces narrow (Sosylopsis) or widened and sometimes strongly spinose at apex; two subequal to strongly unequal spurs always present; tarsi 3-3-3 (Anom-matini) or 4-4-4 in both sexes; tarsomeres simple; pretarsal claws simple; empodium reduced.

Abdomen with fi ve free ventrites. Ventrite 1 lon-ger than 2, without or with rudimentary postcoxal lines; intercoxal process acute, narrowly rounded or truncated. Apical ventrite in Bothriderinae with expanded margins fi tting into interlocking devices on elytra. Functional spiracles on segments I–VII located on pleural membrane. Aedeagus symmet-rical and of cucujiform type; tegmen usually long with articulated, setose parameres and a very long and narrow anterior ventral strut; penis long and narrow, its anterior edge usually with paired, slen-der struts (without paired struts in Sosylopsis). Ster-nite VIII in female with long spiculum ventrale. Ovipositor variable, sometimes very long (Dastarcus) with proctigeral, paraproctal and transverse coxital baculi; paraprocts subequal in length to coxites, which are subdivided into broad basal and slender apical lobes; styli long and subapical. Internal tract with lightly sclerotized capsular spermatheca (sper-matheca apparently absent in Teredinae). [Sharp 1885, 1894, 1895; Grouvelle 1910; Binaghi 1941; Pope 1961, 1980; Dajoz 1965, 1977; Slipinski & Pal 1985; Burakowski & Slipinski 1986; Pal & Lawrence 1986; Stephan 1989; Slipinski et al. 1989; Lawrence et al. 1999 b; Phillips & Ivie 2002.]

Morphology, free living larvae (Xylariophilini, Teredini, Anommatini; Sysolini unknown) (Figs. 10.28.6 C–H). Length 2–4 mm Body elongate and more or less parallel-sided, slightly fl attened. Lightly pigmented including ventral surfaces. Dor-sal surfaces granulate, asperate and tuberculate; dorsal vestiture consisting of simple, expanded, and short scale-like setae.

Head prognathous, narrower than thorax. Pos-terior edge of head capsule arcuate. Epicranial stem short (Anommatus, Teredolaemus) or absent (Xylariophilus); frontal arms lyriform (indistinct in Xylariophilus). Endocarinae absent. Stemmata absent (Anommatus) or fi ve (six) on each side (fi ve well-developed lenses forming two vertical rows, three anterior and two posterior, plus a smaller eye spot at the ventral end of posterior row). Frontoclypeal suture weak (Anommatus) or absent (Teredolaemus, Xylariophilus). Labrum free. Anten-nae separated from mandibular articulation by very narrow strip of cuticle, 0.15–0.2 times as long as head width, 3-segmented; conical sensorium slightly longer than apical antennomere. Man-dibles symmetrical, broad at base and tridentate;


Fig. 10.28.5. Bothrideridae SEM images: A, Deretaphrus erichsoni Newman adult, head, dorsal; B, Deretaphrus erichsoni Newman adult, head, ventral; C, Deretaphrus erichsoni Newman adult, protibia and protarsus; D–G, Bothrideres bipunctatus (Gmelin), fi rst instar larva: D, head, dorsal (© P. Svácha); E, head, dorsolateral; F, abdominal apex, dor solateral; G, head and thorax, ventrolateral; H–J, Deretaphrus adults: H, D. puncticollis Lea, prosternum; I, D. erichsoni Newman, pro- and mesosternum; J, D. erichsoni Newman, antenna. (E–G from Svácha 1994; © Taylor & Francis, London.)

ventral accessory process large or reduced (Anom-matus); incisor edge smooth or with short tooth; mola well-developed, tuberculate; prostheca reduced or absent. Ventral mouthparts retracted; maxillary articulating area present. Cardo trans-verse; stipes about as long as wide; mala truncate (Xylariophilus, Anommatus) or falciform (Teredol-aemus) and setose; palp 3-segmented. Labium consisting of prementum, mentum and submen-tum; mentum only basally connate with maxillae; ligula short, rounded, shorter than labial palps, which are 2-segmented and separated by less than width of fi rst palpomere. Hypopharyngeal sclerome present. Hypostomal rods moderately long and diverging posteriorly; ventral epicranial ridges absent. Gular sutures separate; gula wider than long, fused to labium.

Prothorax not longer than meso- and metatho-rax combined. Thoracic and abdominal terga with granules, tubercles and rarely sharp asperities (Xyl-ariophilus), without distinct sclerotized plates or lateral tergal processes. Prosternum without arma-ture. Legs well-developed, 5-segmented; pretarsus with one seta.

Abdomen less than twice as long as thorax. Terga sometimes extending laterally beyond edges of sterna and forming short processes; terga with-out sclerotized plates; each with transverse row

of scale-like setae or rows of asperities, especially prominent on segments V–IX (Anommatus). Ter-gum IX complex with median elevation (Xylariophi-lus), a pair of dorsolateral processes with upturned urogomphi and two or three additional pairs of processes at the base of each urogomphus; sternum IX simple, not enclosed by sternum. Segment X oval, ventrally oriented. Spiracles annular in Anom-matus; annular-biforous in Xylariophilus and Teredol-aemus, with accessory openings much longer than main openings and placed on the sides of spiracular tubes. [Dajoz 1968, 1977; Klausnitzer 1975; Law-rence 1985; Pal & Lawrence 1986, 1991; Lawrence et al. 1999 a.]

Morphology, Ectoparasitic Larvae (Bothrideri-nae) (Fig. 10.28.6 I). Length 3–18 mm. Body elon-gate, cylindrical with abdomen variously enlarged (physogastric), lightly sclerotized. Vestiture of short, simple setae.

Head protracted, prognathous, distinctly nar-rower than thorax. Posterior edge of head capsule not emarginate. Epicranial stem absent; frontal sutures absent or straight and widely separated. Endocarinae absent. Stemmata absent. Anten-nae about 0.10 times as long as head width, 1- or 2- segmented, separated from mandibular articu-lation by broad strip of cuticle; sensorium long,


inserted on segment 1. Frontoclypeal suture absent. Labrum usually free. Mandibles symmetrical, broad at base and unidentate or bidentate apically; ventral accessory process absent; incisor edge smooth; mola absent; prostheca absent; mandibular base simple or with fi xed acute process. Ventral mouthparts retracted; maxillary articulating area present. Cardo transverse or oblique; stipes about as long as wide; mala falciform (Deretaphrus) or rarely absent; palp 2-segmented; in Sosylus cardo, stipes and articulating area indistinct, mala and palps absent or strongly reduced. Labium mostly free to base of mentum with ligula usually longer than 2-segmented, short and rounded palps; in Sosylus labium not subdi-vided and palp and ligula absent. Hypopharyngeal

sclerome absent. Hypostomal rods absent; ventral epicranial ridges absent. Gular sutures indistinct; gula longer than wide.

Thorax relatively narrow and short. Thoracic and abdominal terga without granules, asperities or distinct sclerotized plates. Legs well-developed, 5-segmented in Deretaphrus, reduced in other taxa; pretarsus if present with 1 seta.

Abdomen more than twice as long as thorax. Tergum IX simple or with short upturned urogom-phi (Deretaphrus, Lithophorus); sternum IX simple. Segment X oval, posteroventrally oriented. Spira-cles annular, usually with heavily sclerotised peri-treme, not raised on tubes; absent on thorax and abdominal segment VIII in Sosylus.

Fig. 10.28.6. A–B, Bothrideridae adults, dorsal: A, Anommatus duodecimstriatus (Müller) (from Klimaszewski & Watt 1997; © Landcare Research New Zealand), length = 2.1 mm; B, Teredolaemus leai (Grouvelle) (from Lawrence & Britton 1991; © CSIRO Australia), length = 3.5 mm; C–I, Bothrideridae larvae; C, Xylariophilus honoratus Pal & Lawrence, dorsal, length = 3 mm; D–E, Teredolaemus leai; D, late instar, dorsal, length = 2.6 mm; E, early instar, pos-terior port of abdomen, dorsal; F–H, Xylariophilus honoratus larval parts: F, lateral portion of segment VI showing frayed setae and spiracle; G, right mandible, ventral; H, portion of right maxilla and labium, ventral; I, Deretaphrus oregonensis Horn, larva, lateral (from Lawrence 1991; © CSIRO Australia), length = 15 mm. (Figs. 10.28.3 C–H from Pal & Lawrence 1986; © Australian Entomological Society.)


Morphology, Triungulin Stage of Ectoparasitic Larvae (Sosylus, Bothrideres, Dastarcus) (Figs. 10.28.5 D–G). Length 1–1.8 mm. Body elongate, fusiform, moderately to strongly fl attened (Sosylus), more heavily sclerotised than mature larvae. Vestiture of sparse simple setae or stout dorsal and ventral spines (Sosylus).

Head strongly transverse (Sosylus) to somewhat elongate (Bothrideres). Labrum and clypeus fused to head capsule. One or two (Dastarcus) well vis-ible stemmata on each side. Antenna 2-segmented with antennomere 1 very short and antennomere 2 elongate (longer than digitiform sensorium) with long terminal seta. Mandibles narrow and falcate, strongly curved and sickle-shaped, without mola or prostheca. Ventral mouthparts protracted; max-illary articulating area absent. Maxilla with cardo and stipes distinct (Sosylus) or fused with labium into single plate (Dastarcus); mala broadly rounded (Sosylus) to strongly reduced (Dastarcus); palps long, 2- or 3-segmented. Labial palps 2-segmented and very close to each other; ligula indistinct.

Thoracic segments, with long, 5-segmented legs; trochanter with long seta; pretarsus simple with single seta.

Segment IX about as long as VIII, without urogomphi but with pair of long terminal setae (very long in Bothrideres and Dastarcus). Segment X long and posteriorly oriented. Spiracles annular, on mesothorax and abdominal segments I–VIII; placed on short tubes in Dastarcus.

Phylogeny and Taxonomy. Members of Both-rideridae were originally classifi ed as distinct

tribes or subfamilies of Colydiidae (= Zopheridae). Within Colydiidae, they were often associated with the tribes Cerylini or Cerylonini (currently Cerylonidae) based on shared exposed antennal insertions in both groups (Erichson 1845; Horn 1878; Reitter 1882; van Emden 1928; Hetschko 1930; Heinze 1943). The subfamily Anommatinae was transferred to Latridiidae by Jacquelin du Val (1859) because of their trimerous tarsi, but was relocated by Ganglbauer (1899) to a redefi ned sub-family Cerylinae that included the tribes Cerylini (currently Cerylonidae), Bothriderini, Deretaph-rini (= Teredini) and Anommatini (currently Both-rideridae), further acknowledging the similarity between Cerylonidae and Bothrideridae. This con-cept was further elaborated by van Emden (1928) who added to that group the tribe Euxestini from Erotylidae. Cerylonidae family status was formally recognized by Crowson (1955), while Bothride-ridae, in spite of suggestions by Craighead (1920), continued to be treated as a colydiine subfamily. That classifi cation remained until Lawrence (1980) demonstrated they clearly belonged to Cucujoidea rather than to Tenebrionoidea. Within Cucujoi-dea, Bothrideridae are placed next to Cerylonidae in a crown group called the Cerylonid Series (Crow-son 1955; Pal & Lawrence 1986; Slipinski & Paka-luk, 1991). Except for recent works by Hunt et al. (2007) and Robertson et al. (2008), there has been no phylogenetic analysis for this assemblage and most of the placements are based on intuition alone. In spite of concerns about the monophyly of this group by Slipinski & Pakaluk (1991), the molecular data provided by Hunt et al. (2007) and

Fig. 10.28.7. Bothrideridae biology and morphological characters: A, Bothrideres cryptus Stephan coccons (Red Oak, Oklahoma), length of one cocoon = 5.4 mm; B, Deretaphrus puncticollis Lea cocoon mass and emerging adult attached to larval host, Trichaulax philipsii; C, Anommatus duodecimstriatus (Müller) adult, head exhibiting anophthalmy; D, Deretaphrus piceus (Germar) hindwing, exhibiting closed radial cell; E, Deretaphrus xanthorrhoeae Lea cocoon, length = 10.6 mm.


Robertson et al. (2008) support a monophyletic Cerylonid Series.

A rigorous morphological analysis of the family has not yet been conducted, and the group is not supported by any exclusive morphological char-acter. Hunt et al. (2007) included three of the four bothriderid subfamilies (Anommatinae, Xylari-ophilinae and Teredinae), and the analysis resulted in a paraphyletic Bothrideridae. Although Both-rideridae was recovered as monophyletic by Robert-son et al. (2008), the results are not very informative at the family level as all bothriderid taxa in the dataset belong to a single subfamily (Bothrideri-nae). Both molecular studies lacked adequate taxon sampling to draw defi nitive conclusions pertaining to the classifi cation of the family. Both suggest a close relationship between Bothrideridae and Cery-lonidae. However, the support for this association was weak. Based on results from Robertson et al. (2008) we can only infer that Ceryloninae and Both-riderinae may be sister taxa while the placement of Deretaphrus remains to be studied. From a character chart and thorough discussion in Pal & Lawrence (1986), it is clear that Cerylonidae (Euxestinae, Metaceryloninae) are almost inseparable from free living Bothrideridae (Teredinae) and their relation-ships need further testing. Teredinae, Xylariophili-nae and Anommatinae probably belong together with their free living mycophagous larvae, pupae not enclosed in cocoons, and adults with narrowly separated coxae, normal trochanters, and wings with a medial fl eck. Most of the listed characters are shared by Euxestinae (Cerylonidae) but their potential sister group relationships (suggested by Pal & Lawrence 1986) need further research. The hypermetamorphic larvae and modifi ed trochan-ters are exclusive characters for Bothriderinae (including Deretaphrus), but their internal relation-ships remain to be studied.

Acknowledgements

CSIRO Australia is acknowledged for support of research (for SAS and JFL) and CSIRO Austra-lia, Landcare Research New Zealand and Taylor and Francis and for allowing us to use copyrigh-ted images. Portions of this Chapter benefi ted from research carried out with the support of a National Science Foundation grant “Assembling the Beetle Tree of Life” (DEB-0531768) and NSF PEET grant (DEB-0329115) to J. V. McHugh, M. F. Whiting and K. B. Miller. The following art-ists are acknowledged: F. Nanninga (Fig. 10.28.1), D. Helmore (Fig. 10.28.6 A), S. Montieth (Fig. 10.28.6 B), S. P. Kim (Figs. 10.28.6 C–E), A. Hastings (Fig. 10.28.6 F–H), A. Atkins (Fig. 10.28.6 I). Anne Hastings is gratefully acknowledged for composing the plates. Thanks are also given to Dr. Petr Svácha for providing prints of the images in Figs. 10.28.7 D–G.

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10.29. Cerylonidae Billberg, 1820

Adam Slipinski and John F. Lawrence

Distribution. Cerylonidae are rather small family comprising about 450 described species classifi ed in 52 genera divided into fi ve subfamilies. They are found in all zoogeographical regions of the world but are more abundant in the tropical and subtropical forested areas with relatively few spe-cies in temperate or open habitats. Cerylonids are found on all habitable continents and larger oce-anic islands. They may occur at elevations as high as 4000 m in the Himalayas. Native Cerylonidae are absent from most Atlantic islands except for Madeira and the West Indies. They are also absent from most mid Pacifi c islands except for the almost cosmopolitan species Euxestus erithacus Chevro-lat, which is found on Hawaii, the Marquesas and Samoa. Most of the larger Pacifi c islands are inhabited by Cerylonidae, and they occur as far as Society and Cook Islands in SE Polynesia, and Cha-tham Islands east of New Zealand. Large Pacifi c

islands like Fiji, New Caledonia and New Guinea have a rich fauna for their size, but the majority of species represent three widely distributed genera of Ceryloninae: Philothermus Aubé, Philothermopsis Heinze and Australiorylon Slipinski. Euxestinae are cosmopolitan with maximum diversity in the Oriental Region (seven of the eleven known gen-era) but are almost absent from Europe and Japan (except for the cosmopolitan Euxestus Wollaston). Endemic austral euxestine genera include Hypodac-nella Slipinski from Australia, New Caledonia and New Zealand and Protoxestus Sen Gupta & Crowson from southern Queensland. The Caucasian Hypo-dacne edithae (Reitter) and eastern North American H. punctata LeConte are the only true Holarctic members of this group. Metacerylon Heinze has numerous species in Africa and South America with single isolated species in Australia and Indo-nesia. Euxestoxenus Arrow is the largest genus in this subfamily, with most species in Africa and a few offshoots in continental Asia, while the sec-ond largest genus, Euxestus Wollaston, has numer-ous endemic species in the Afrotropical, Oriental and Australian regions. E. erithacus Chevrolat has probably been introduced to various parts of the world including Florida and South America. The monotypic, Oriental Loebliorylon Slipinski (subfam-ily Loeblorylinae) comprises nine known species from higher elevations in Himalayas, northern

Fig. 10.29.1. Philothermopsis alienigenus (Blackburn), adult, dorsal (from Britton 1970; © CSIRO Australia), length = 1.8 mm.

422 Adam Slipinski and John F. Lawrence

10.28. bothrideridae erichson, 1845

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