Deretaphrus Newman, 1842

Lord, Nathan P. & McHugh, Joseph V., 2013, A Taxonomic Revision of the Genus Deretaphrus Newman, 1842 (Coleoptera: Cucujoidea: Bothrideridae), The Coleopterists Bulletin (mo 12) 67, pp. 1-107 : 6-16

Deretaphrus Newman, 1842

Deretaphrus Newman, 1842: 403–404 . Type species: Deretaphrus fossus Newman, 1842: 404 .

Sigerpes Germar, 1848: 222–223 . Type species: Sigerpes piceus Germar, 1848: 223 . Synonymized by Lacordaire 1854: 377–378.

Derataphrus: Lacordaire 1854: 377–378, Lacordaire 1876: 8, pl. 20. Misspelling, no status.

Adult Diagnosis. Within the Bothrideridae , Deretaphrus is characterized by the loose, 3-segmented antennal club, strongly enlarged submentum which partially or completely conceals the palpi from view, externally closed coxal cavities, strongly heteromeroid trochanters, the intercoxal process of abdominal ventrite I acute, and the hind wing with a closed radial cell. Keys to separating New World bothriderid genera can be found in Stephan (1989) and Philips and Ivie (2002), and a key to the genera of Deretaphrini sensu Heinze can be found in Heinze (1943).

Adult Description. Body: Length 4.9–12.9 mm. Width 1.15–3.6 mm. Elongate, parallel, slightly flattened to subcylindrical, reddish orange to black; surface luster matte to moderately shiny; glabrous except for minute setae that arise from the punctures. In general, adult Deretaphrus are heavily sclerotized. Head ( Figs. 18–20 ): Slightly to moderately declined ( Fig. 3 ), evenly convex to flattened, a little narrower than prothorax, elongate; with a weak lateral postocular (transverse) occipital impression that continues laterad and meets the submental suture ventrally, completely separating the vertex from the occiput. Occiput swollen, forming a bulbous posterior portion of the head capsule. Head not abruptly constricted posteriorly to form a neck. Occipital foramen small to moderate in size. Temples very short and shelf-like. Frons and vertex simple; stridulatory files absent. Frons sometimes with an evenly rounded to angulate expansion of the head capsule above and posterior to the eye (hereafter called supra-ocular ridges), often partially or completely concealing eyes from above. Frons anterolaterally emarginate (at frontoclypeal suture) to receive antennal scape. Frontoclypeal suture indistinct to distinct, arcuate to sharply incised medially. Clypeus with lateral and anterolateral emarginations forming 4 acute, dentate projections on anterior margin, antennal scape received in lateral emarginations at level of frontoclypeal suture. Anterior edge of clypeus straight to convex to sharply incised medially. Antennal insertions dorsolateral, moderately to widely separated and completely exposed from above. Subantennal groove well-developed, deep; present on ventrolateral surface, extending posteriad from the antennal base to posterior edge of the eye. Submentum forms anteroventral edge of head capsule. Submentum strongly produced and highly variable, slightly to strongly projecting anteroventrally, delimited posteriorly from the submental suture which lies at the bottom of a broad furrow. Laterally, submentum is margined by strongly expanded subgenal braces. Submentum and subgenal braces widely separated ( Figs. 50–51, 53 ), narrowly separated ( Fig. 48 ), touching ( Fig. 49 ), or partially fused ( Fig. 52 ). Submentum sometimes with paired setose pits anterior to submental suture, near connection of submentum and subgenal braces ( Figs. 49, 51, 53 ). Gena bearing a distinct pit near mandibular articulation (visible in Fig. 48 ), most likely the external manifestation of an apodeme for the attachment of the anterior arms of the corporotentorium. Gular sutures widely separated, narrowing anteriorly. Cervical sclerites absent. Corporotentorium narrow; with anterior arms and narrow tentorial bridge; lacking dorsal arms. Bulbous posterior portion of head capsule usually retracted into anterior region of the prothorax (as ball-and-socket joint). Cervical membrane with 2 short, transverse, dorsolateral rows of anteriorly projecting setae ( Fig. 18 ). Eyes: Moderately large, coarsely facetted, lacking interfacetal setae. Eyes evenly convex (most) to nearly angulate ( D. carinatus and D. parviceps , Figs. 36, 38 ), slightly to strongly protuberant. Posterior margin of eye straight to convex and bordered by a thickened strip of cuticle (most) to distinctly emarginate posteriorly by a cuticular triangular projection of the head capsule ( D. ocularis , Figs. 40–41 ). Antennae: Eleven-segmented with weak, loose, slightly flattened, 3-segmented apical club. Antennal scape moderately large, asymmetrical, more strongly inflated anteriorly, round, punctate, articulation with pedicel subapical; pedicel narrower and slightly elongate; segments 3–8 short, subcylindrical and subequal. Antennal club asymmetrical, inflated at leading edge (when antennae held out laterally), segments 9–11 subtriangular, segment 10 larger than 9 and 11. Apex of segments 9–11 with 1 or more sensilla-bearing depressions or cupules ( Figs. 46–47 ). Antennal club with minute setae ( Fig. 44 ), sparsely setose with short setae ( Fig. 46 ), or densely setose with long, erect setae ( Figs. 45, 47 ). Mouthparts: Labrum ( Fig. 21 ) partially concealed beneath clypeus, strongly transverse, apex subtruncate and slightly sclerotized, with a row of long, stiff setae projecting anteriorly; ventral surface with median, raised triangular portion (referred to as an epipharyngeal impression by Pal and Lawrence 1986), lateral margins of raised portion densely setose; mesal arms of tormae small, separate, oblique, projecting anteromedially; lacking brushlike epipharyngeal process (as is found in most Bothrideridae ). Maxilla ( Fig. 22 ) with distinct galea and lacinia. Galea subtriangular with apex broad and densely setose, lacking hooks or teeth. Lacinia long and narrow, with a bifid apex. Lacinia densely setose, setae stouter and thicker distally near the apex. Maxillary palp 4-segmented. Labial palp 3-segmented. Apical maxillary and labial palpomeres weakly fusiform. Ligula undivided. Mandible ( Fig. 23 ) short and broad; incisor edge simple; mola and prostheca absent. Mandibular base punctate; apex smooth, bidentate apically, some species with a subapical tooth on inner mesal margin. External (dorsal) face of mandible with short setae, few long setae arising from punctures medially, or with median setose groove bearing long setae. Prothorax: Pronotum quadrate to elongate, widest at middle or anterior half, narrowing posteriorly; dorsal surface convex (e.g., Fig. 64 ), flattened (e.g., Fig. 58 ), or slightly depressed at middle (e.g., Figs. 62 , 75, 77–78, 80 ). Pronotal disc punctate, punctures variable in size and density. Anterior margin of pronotum straight or arcuate; anterior angles rounded and indistinct (e.g., Fig. 73 ) to distinct and obtuse (e.g., Fig. 65 ). Base of pronotum slightly to moderately narrower than elytral bases, weakly sinuate to arcuate between posterior angles; posterior angles distinct, often with single small denticle (e.g., Fig. 67 ). Pronotal disc usually with median longitudinal depression, groove, or canal; canal sometimes interrupted, forming an anterior fovea and a posterior canal, both separated by an elevated, usually impunctate portion of the pronotal disc (e.g., Figs. 59 , 70 , 76 ). Sides of prothorax more or less straight (e.g., Figs. 58 , 77 ), slightly arcuate to subtly sinuate (e.g., Figs. 69, 73 ), or narrowing towards base (e.g., Figs. 59 , 70, 74 ). Pronotum margined laterally by an incomplete carina with a raised margin or bead; when viewed laterally, carina gradually to abruptly curved anteroventrally, ending before anterior margin of prothorax ( Fig. 3 ). Pronotum strongly deflexed laterally, forming the hypomeron. Hypomeron sparsely punctate, with long, narrow postcoxal projections that meet under the prosternal process. Tergosternal suture straight to slightly sinuate, originating at anterolateral margin of procoxa, extending anteriorly, parallel to the posterolateral margin of the prothorax, terminating before anterior margin. Prosternum with a transverse, sinuate groove anterior to the procoxae ( Fig. 54 ); groove incised for entire length anterior to coxae or on either side of midline and smooth medially, groove extends laterally and ends at tergosternal suture lateral to procoxae, separating prothoracic basisternum from prosternal process. Prothoracic basisternum quadrate to elongate, flat to moderately convex, always longer than prosternal process. Prosternal process complete, narrowed then strongly and abruptly expanded behind coxae, expansion meeting postcoxal projections. Prosternal process ( Fig. 54 ) slightly elevated posterior to the coxae, broadly rounded, slightly overlapping mesoventrite. Prosternum bearing row of short, golden setae at anterior margin that project anteriad opposite the base of the head; margination continues laterally and dissipates near anterior pronotal angles. Posteriorly, entire basal margin of prothorax bears row of short, golden setae that project posteriad. Procoxae narrowly separated. Procoxal cavities round, narrowly separated at middle, internally open, externally broadly closed by postcoxal projections. Prothoracic trochantin concealed. Pterothorax: Mesoscutellum visible, abruptly elevated, shield-shaped. Mesoscutum shield-shaped, with fine, crenulate microsculpturing. Mesoscutum with anterolaterally projecting arms and postnotal elytral process. Lateral margin of mesoscutum bearing sharp angulate projection, locking into grooves on elytral shelf. Mesoscutum with weakly visible longitudinal mesothoracic suture, represented internally by the longitudinal mesothoracic ridge. The internally projecting yolk plates of the mesoscutum are pointed and sclerotized. Anterior edge of mesoventrite at midline on different plane than mesosternal process and metaventrite. Mesothorax usually retracted into posterior region of the prothorax. Mesocoxae ( Fig. 27 ) round, countersunk, narrowly separated, closed laterally by meso- and metaventrite. Mesoventral process extending to middle of mesocoxae. Meso-metaventral junction between mesocoxae a notched fitting, apex of metaventrite overlapping mesoventrite ( Fig. 27 ). Mesepisterna distinctly separated, variously punctured. Mesoventrite variously punctured. Metaventral discrimen deeply impressed, tapering and becoming weaker anteriorly, terminating near midpoint of metaventrite. Metaventrite longer than first abdominal ventrite, variously punctured. Metepisternum long, narrow. Metepimeron highly reduced, represented only weakly anteriorly in the arm of the notal wing process, obsolete posteriorly. Metacoxae ovoid, transverse, countersunk, narrowly separated. Lateral furcal arms of metendosternite moderately long, apices swollen. Metendosternite ( Figs. 26–27 ) with anterior process absent; apical edge not emarginated; laminae well-developed, narrowed, projecting slightly posteriad from beneath bases of furcal arms; ventrolateral processes absent; metafurcal ventral flange well-developed; anterior tendons widely separated, arising from short, pointed, anteriorly projecting arms. Elytra ( Figs. 82–131 ): Elongate, completely concealing abdomen. Elytron with 9 distinct puncture rows. Scutellary striole absent. Elytral punctures range from inconspicuous (e.g., D. analis ) to deep and strongly marked (e.g., D. ignarus ). Interstitial intervals (interstriae) flat (e.g., Figs. 100–101 ), to feebly raised (e.g., Figs. 86–87 ), to strongly carinate (e.g., Figs. 122–123 ). In some species, apex of interstitial intervals with micropunctures, each bearing a minute seta (e.g., D. puncticollis ). In most species, interstitial intervals 3, 5, and 7 elevated at base and apex of elytron. In some, elytral interstitial carinae join near apex (e.g., D. puncticollis , Fig. 122 ). Carina of interstitial interval 9 begins at humeral angle, extends posteriad and merges with elytral apex. Inner margin of elytra forms tongue and groove locking apparatus. True basal margin of elytron strongly depressed, forming large elytral shelf. Grooves on inner margin of shelf articulate with and lock into corresponding notches on lateral margin of mesoscutum. Apex of humeral angles with knob-like or finger-like callosities or protuberances, sometimes merging with elytral interstitial carina. Callosities sometimes extend anterolaterally at the basal margin of elytron to epipleuron. Epipleuron gradually set off from lateral margin of elyron, narrowing posteriorly until dissipating near the elytral apex. Mesal edge of epipleuron delimited by a sharply impressed longitudinal groove; groove confluent with the sutural and apical elytral carinae. Lateral edge of elytron straight. Basal edge of elytron (when viewed laterally), weakly sinuate. Hind wing ( Fig. 25 ): With well-developed, elongate radial cell. Inner posterior angle formed at base of radial cell 90°. Median area of hind wing with 4 free veins. Medial fleck present, weakly developed, partially bisected by a free vein. Wing veins in medial area variable inter- and intra-specifically. Wedge cell present in some species, minute or absent in others. Anal lobe present. Legs ( Figs. 27 , 54–56 ): Trochanterofemoral attachment of the heteromeroid type; trochanter partially concealed within excavations in proximal end of femur, femur nearly touching coxa. Trochanter and coxa sometimes bearing sparse setae. Femur glabrous, swollen near middle; at apex, anterior and posterior walls with expanded, downward-projecting semicircular plates, concealing the femorotibial articulation from the sides ( Figs. 27 , 55 ). Tibia slightly expanded apically, outer edge with row of small denticles, inner edge sparsely to moderately setose; outer apical angle produced into large, outwardly facing tooth (mucro) and crown of smaller teeth; with 2 subequal, glabrous tibial spurs, 1 spur much larger and thicker; curved Figs. 55–56 ). Tarsal formula 4-4- 4 in both sexes; tarsomere I distinctly longer than II. Tarsomeres sparsely setose, some with pockets of stout, bristle-like setae on ventral surface arising from transverse slits near anterior margin of tarsomeres ( Fig. 55 ). Tarsal claws simple and subequal. Empodium reduced. Procoxa cylindrical, countersunk, not or slightly projecting below prosternum. Abdomen ( Figs. 28 , 132–139 ): Abdominal tergites completely concealed by elytra. With 5 subequal ventrites. Intercoxal process of ventrite I narrowed, slightly pointed anteriorly, apex notched to interlock with metathorax. Lateral edges of abdominal ventrites I–V (= sternites III–VII) upturned (laterosternites). A parasternite is also present between the pleural membrane and laterosternite of ventrite I (= sternite III). Ventrites variously punctate. In most species, abdominal ventrite V simple. In some, abdominal ventrite V bears 2 depressions, 1 on each side of midline, resulting in raised, median, V-shaped area near posterior margin ( Figs. 132–133 ), present in both sexes. In others, ventrite V is sexually dimorphic, with a bulbous swelling near posterior margin (females of D. erichsoni , Figs. 134–135 ), or a notched posterior face (females of D. xanthorrhoeae , Figs. 138–139 ). Tergite VII with functional spiracle in pleural membrane; lateral and apical margin slightly more sclerotized and darkly pigmented than remainder of tergite; apical margin fringed with setae. Male terminalia ( Fig. 29 ): Tergite VIII concealed dorsally by tergite VII. Tergite VIII and sternite VIII fused at base, forming a ring; sternite VIII narrowly divided at middle, connected by membrane; apical margin of tergite VIII slightly emarginate; apical margins of tergite and sternite VIII fringed with setae. Tergite IX concealed dorsally by tergite VIII and tergite VII. Anterior edge of tergite IX not developed into spiculum gastrale. Tergite IX divided into two separated laterotergites, attached to sternite IX (ventrolaterally) and tergite X (dorsolaterally); sternite IX divided into two separate, well-sclerotized subtriangular plates, each produced anteriorly into narrow struts; both plates connected by a membrane medially. Tergite X (= proctiger) welldeveloped, free, lightly sclerotized, with weakly indicated paired longitudinal baculi; apical margin shallowly emarginate. Laterotergites IX (positioned laterally), sternite IX (positioned ventrally), and tergite X (positioned dorsally) form a ring around the aedeagus. Male genitalia ( Figs. 30–31 , 140–172 ): A brief description and illustrations of the aedeagus of D. ignarus were presented by Sharp and Muir (1912). Phallobase of cucujiform type, forming a ring that completely surrounds median lobe (= penis). Parameres attached by a membrane to distal edge of phallobase, free from one another at base. Tegmen (dorsal) and basal piece (ventral) reduced, mostly membranous, with slightly more sclerotized areas medially. Tegmen and basal piece normally connected laterally by membrane, although may be connected by slightly more sclerotized portions in some species. Tegmen with well-developed, elongate, heavily sclerotized, rod-like anterior strut that sits dorsal to the penis, loosely attached medially by a membrane. Tegmen consisting of 2 sclerotized plates, often meeting at midline; anterior margin of tegmen usually evenly curved, but may be produced into anteriorly– projecting sclerotized lobes ( D. analis , D. gracilis , D. iridescens , D. piceus ; Figs. 142 , 149, 154 , 159 ). In most species, the basal piece is emarginate anteromedially, resulting in an evenly curved anterior margin and the lateral margins produced into two short, anterolateral struts; struts straight (e.g., Figs. 153, 156 ), to slightly curved laterad (e.g., Figs. 151 , 163 ) to strongly recurved (e.g., Figs. 149 , 159 ). In D. oregonensis , the basal piece is projecting and pointed apically (lacking lateral struts; Fig. 157 ). Parameres separate (in most species) to broadly fused to one another at base (e.g., Figs. 142 , 149, 154 , 159 ); degree of separation variable; shape variable, with apices narrowly rounded to truncate; size varying from small to large (in relation to phallobase). Parameres evenly sclerotized (in most species) to possessing areas of greater sclerotization (e.g., Figs. 142 , 150 , 160 ); apical and/or margins usually bearing setae. Parameres with paired, hook-like lobes (in most species, hereafter referred to as “parameral processes”) attached to dorsal surface of parameres basomedially, serving as a penis guide. Parameral processes well-separated to nearly touching medially; orientation variable, transverse to vertical; shape variable, straight to strongly curved; size variable, reduced to nearly as large as parameres. Apices of processes evenly rounded to slightly swollen to curved and pointed. Penis (= median lobe; Figs. 165–172 ) elongate, tubular to slightly flattened, straight in dorsal/ventral view, evenly curved to subtly sinuate in lateral view; apex narrows gradually to a point (most) to swollen and pointed ( D. bucculentus , D. parviceps ); apex open dorsally, opening (= median orifice of Sharp and Muir (1912)) of variable size, closed basally by fusion of margins of penis, suture of variable length, but completely obsolete in basal half. Base of penis open, slightly to greatly swollen, usually trumpet-shaped, bearing paired anterior struts; struts positioned dorsally (in most species) to dorsolaterally; length short to long; shape truncate to narrow, straight to curved mesally to nearly touching at apices. Penis dark red to brown, unicolored to lighter near base (in most species) and appearing bicolored. Penis slides through ring-shaped phallobase and ventrad to parameral processes. Internal sac undifferentiated. Female terminalia ( Fig. 32–33 ): Sternite VIII and tergite VIII separate, connected by a thin portion of membrane, forming a ring; sternite VIII entire, not divided at middle, apical margin truncate medially, fringed with setae; basal margin thickened (as weakly indicated transverse baculi), evenly curved, with welldeveloped, elongate, heavily sclerotized, rod-like spiculum ventrale, loosely attached medially by a membrane. Tergite VIII with sparse setae in medial area, apical margin fringed with setae and with slight medial emargination. Ovipositor ( Fig. 32 ): Elongate, with well-defined but weakly sclerotized segments. Tergite IX completely divided into 2 lateral paraprocts. Paraproct with long baculus, about 2X as long as gonocoxite. Tergite X (= proctiger, epiproct) weakly sclerotized, situated between paraprocts; apical margin of tergite X rounded, extending slightly past junction of paraproct+ gonocoxite. Gonocoxite divided into proximal (= valvifer) and distal (= coxite) lobes; proximal lobe with weakly indicated transverse basal baculi; distal lobe not more heavily sclerotized than proximal lobe; apex of distal lobe with few setae, with well-developed, palpiform gonostylus attached sub-apically; gonostylus with few setae at apex. Female reproductive tract ( Fig. 34 ): Vagina membranous; common oviduct flattened, fan-shaped and connecting to vagina apically; spermatheca membranous, sacular, annulate, with spermathecal gland attaching near the base; spermathecal gland tubular, elongate and narrow, with minute, dense, microtubules covering surface, base of gland with sclerotized ring.

Larvae. No larvae were examined, as they are rarely collected. For description of a larva ( D. oregonensis ), see Craighead (1920) and Böving and Craighead (1930).

Etymology. The genus-group name Deretaphrus ( Dere: Greek “neck,” – taphrus: Greek “a trench”) refers to the transverse, postocular constriction of the head capsule forming a bulbous posterior portion ( Borror 1961).

Taxonomic History. Deretaphrus was first described by Newman (1842) from insects collected at Port Phillip in Victoria, Australia. A limited genus-group description and short descriptions of four species ( D. fossus , D. puteus , D. illusus , and D. vittatus ) were provided. Erichson (1845) later placed the genus in Bothriderini within Colydiidae . He also moved two species ( D. illusus and D. vittatus ) into the genus Bothrideres Dejean.

Germar (1848) described Sigerpes , containing one species, S. piceus . Lacordaire (1854) synonymized Sigerpes Germar with Deretaphrus and provided a more detailed description of the genus.

Wollaston and Newman (1855) supported Erichson’ s placement of the genus within Colydiidae , but also stated that Deretaphrus has several characters potentially allying it with other taxa. Wollaston recognized its close relationship with Bothrideres and presented a fairly detailed description of the genus, mentioning several previously undescribed characters to support his hypothesis (Wollaston and Newman 1855). In the same paper, Newman conceded Erichson’ s move of two of his original species into Bothrideres . He also described two new species, Deretaphrus wollastoni and Deretaphrus erichsoni . Newman closed the paper with an updated list of the species, noted the Australian distribution, and stated that they are found “…feeding under the bark of trees, principally Eucalyptus .”

Based on his observation of the type specimen at the BMNH, Pascoe (1862) moved Deretaphrus puteus (one of Newman’ s originally described Deretaphrus species ) into the genus Bothrideres . Contrary to Wollaston, Pascoe stated “The affinity of the two genera, Deretaphrus and Bothrideres , seems to be by no means close…” Pascoe described four new species ( D. ignarus , D. viduatus , D. bakewellii , and D. colydioides ) but was unable to study D. wollastoni and could not locate the type specimen. Pascoe provided the first key to the species of the genus. Macleay (1871) described Deretaphrus pascoei .

In the Catalogue of the Coleoptera of Australia, Masters (1872) listed 10 species of Australian Deretaphrus and their localities, apparently unaware of Pascoe’ s placement of D. puteus within Bothrideres some 10 years prior.

Horn (1873) described D. oregonensis from the western United States, noting the odd distribution of the genus. Reitter (1877) described D. granulipennis from New South Wales. Horn (1878) treated the North American Colydiidae and erected the tribe Deretaphrini on the basis of a deflexed head and the contiguous (or nearly so) anterior coxae. Within the tribe, he included Oxylaemus Erichson , Deretaphrus , and Sosylus Erichson and provided a brief generic description and species description for D. oregonensis , comparing and contrasting this sole North American representative with the type species, D. fossus .

Lea (1898) described four new species: D. analis , D. xanthorrhoeae , D. parviceps , and D. puncticollis . Grouvelle (1903) described D. interruptus from New Caledonia, only the second non-Australian species in the genus at that time. Fauvel (1903) noted D. interruptus in New Caledonia, and tallied 16 species in the genus.

One of the major works on Deretaphrus was by Blackburn (1903), who discussed the ambiguity of several species descriptions preventing confident identifications. He synonymized Deretaphrus granulipennis Reitter with Deretaphrus erichsoni Newman and indicated that D. wollastoni probably did not belong in the genus. Blackburn also mentioned that it was difficult to identify the type species, D. fossus , because the original description was inadequate to differentiate it from several congeners. He discussed some variation in taxonomic characters (color, size, elytral punctation) and provided several characters that he believed were better at diagnosing Deretaphrus species. He described eight new species ( D. thoracicus , D. gracilis , D. iridescens , D. cordicollis , D. popularis , D. aequaliceps , D. sparsiceps , and D. cribriceps ) and provided an identification key. Blackburn relied on locality in his delimitation of species, which later resulted in the synonymization of most of his species.

Craighead (1920) addressed the taxonomic position of the Colydiidae based on his study of larval morphology, noting dissimilarities between bothriderine taxa and the rest of the family. He employed the familial name Bothrideridae but formally retained the group within Colydiidae as a “specialized development” of the colydiids due to their parasitic nature. Craighead also described and illustrated the larva, pupa, and cocoon of D. oregonensis .

Elston (1923) described D. bucculentus . Perhaps the most significant work on the genus Deretaphrus was Carter and Zeck’ s (1937) revision of the Australian Colydiidae . Carter and Zeck mentioned the unfortunate loss of the type specimen for the type species, D. fossus , and again discussed the many synonyms that had been created due to ambiguous species descriptions. They synonymized several species: D. bakewellii Pascoe and D. thoracicus Blackburn with D. piceus (Germar) , D. pascoei Macleay with D. ignarus Pascoe , D. cordicollis Blackburn with D. fossus Newman , and D. bucculentus Elston with D. puncticollis Lea. Carter and Zeck also stated that D. colydioides Pascoe is probably a small D. viduatus Pascoe , and D. sparsiceps is most likely a junior synonym of D. viduatus . Carter and Zeck (1937) also described two new species, D. alveolatus and D. incultus .

Heinze (1943) revised the tribe Deretaphrini , stating that he considered the species accounts of Blackburn (1903) and Carter and Zeck (1937) to be inadequate for correct identifications.

In The Natural Classification of the Families of Coleoptera ( Crowson 1955) , Deretaphrus was classified within Colydiidae . Crowson considered Deretaphrus as “possibly the most primitive genus” in the group and allied it with other current bothriderid genera. He also illustrated the mesothorax and metendosternite, representing the first and only morphological study of the genus in any detail.

Since the redefinition of Bothrideridae ( Lawrence 1985, 1991; Pal and Lawrence 1986), Deretaphrus has been considered a member of the tribe Deretaphrini within the Bothriderinae in all major works on the family ( Stephan 1989; Lawrence and Britton 1991; Lawrence and Newton 1995; Philips and Ivie 2002; Ślipiński et al. 2010).

Natural History. Although Deretaphrus is the most commonly encountered genus of Australian Bothrideridae (Lawrence and Britton 1991) , the biology of most species is poorly known. The larvae are ectoparasitic on a variety of hosts, predominantly wood-boring beetles in the families Cerambycidae and Buprestidae ( Burke 1919; Craighead 1920, 1950; Böving and Craighead 1931; Balduf 1935; Clausen 1940; Furniss and Carolin 1977; Stephan 1989). Larvae have also been collected inside the nests of solitary bees (S. A. Ślipiński, personal communication). Members of Deretaphrus have hypermetamorphic development, existing as a triungulin form during the first instar (S. A. Ślipiński, personal communication) and a grub-like form in the later instars (Böving and Craighead 1931; Lawrence 1991). This hypermetamorphic lifestyle is consistent with an ectoparasitic life history. During the last instar, Deretaphrus larvae migrate away from the host and spin elongate-oval silken cocoons ( Figs. 8–17 ). The cocoon is composed of a threadlike, fibrous material that may be loose or tightly layered. It is not known whether the secretions are emitted from the buccal cavity or the anal glands of the larvae ( Balduf 1935). The cocoons are attached near or directly onto the host, and the larvae pupate within their last larval skin inside the cocoon ( Fig. 8 ). The construction of a silken cocoon is uncommon among Coleoptera , having been reported in some species of only five other families ( Brachypsectridae , Tenebrionidae , Curculionidae , aleocharine Staphylinidae , and Cerylonidae ) ( Ashe 1981; Lawrence and Britton 1991; Leschen 1991; S. A. Ślipiński, personal communication). The duration of the immature stage of Deretaphrus is not known, however, label data from reared specimens suggest pupation lasts for a minimum of two months (e.g., D. rodmani, WAMP 43NL – larva 21.x.80, adult 23.xii.80).

Adults have coarsely facetted eyes and are commonly collected at mercury-vapor and/or UV lights or are found crawling on the surface of logs at night (M. Caterino, personal communication), suggesting a nocturnal lifestyle. Most adult Deretaphrus , however, are collected under the bark of fallen trees. Although little is known about the adult feeding habits, it is assumed adults feed within the galleries of wood-boring insects. In The Biology of the Coleoptera, Crowson (1981) suggested that the ectoparasitic nature of the colydiid subfamilies Deretaphrinae and Bothriderinae (in part) developed from close associations between wood-boring beetle larvae and bothriderids inside the fungus-filled galleries where they lived.

The North American D. oregonensis is most often collected under the loose, dry bark of trees. The literature and specimen label data indicate that larvae of D. oregonensis are associated with a variety of hosts and are found under bark of a number of different tree species. It has been collected from under the bark of ponderosa pine ( Pinus ponderosa Douglas ex C. Lawson , Pinaceae ), Jeffrey pine ( Pinus jeffreyi Balfour ), sugar pine ( Pinus lambertiana Douglas ), lodgepole pine ( Pinus contorta subsp. murrayana (Balfour) Engelmann ), Douglas fir ( Pseudotsuga menziesii (Mirbel) Franco , Pinaceae ), red fir ( Abies magnifica Murray ), white fir ( Abies concolor (Gordon and Glendinning) Hildebrand , Pinaceae ), western larch ( Larix occidentalis Nuttall , Pinaceae ), and canyon oak ( Quercus chrysolepis Liebm. , Fagaceae ). Burke (1919) reported that cocoons had been collected “from the wood of an old fire scar on the side of a living incense cedar [ Calocedrus decurrens (Torrey) Florin , Cupressaceae ].” Label data also indicate that D. oregonensis has been recovered from under the bark of burned pines. It has been associated with the following presumed beetle hosts: Curculionidae : Dendroctonus jeffreyi Hopkins , Dendroctonus ponderosae Hopkins ; Buprestidae : Buprestis laeviventris (LeConte) , Buprestis aurulenta L., Trachykele opulenta Fall , Trachykele nimbosa Fall ; and Cerambycidae : Asemum striatum (L.) ( Burke 1919; Essig 1926; Balduf 1935; Clausen 1940; Philips and Ivie 2002). It is hypothesized ( Burke 1919; Balduf 1935) that Deretaphrus parallel the life cycles of their hosts, potentially persisting in the larval stage for over a year. Amusingly, D. oregonensis was given the nickname “The Buprestid Destroyer” by Essig (1926), who stated that the adult “lives in the burrows and destroys the larvae and pupae…” of a number of wood-boring beetles in California and Oregon.

The Australian Deretaphrus species have been collected in association with the larvae of several cerambycids, including Paroplites australis Erichson , Penthea saundersii Pascoe , and Phoracantha sp. Adults and pupal cocoon masses have also been discovered inside the pupal cells of a cetoniine scarab, Trichaulax philipsii (Schreibers) (previously unpublished). Specimens of D. viduatus have been collected from within Sarcocornia stems ( Amaranthaceae ) containing Belus sp. ( Coleoptera : Belidae ) (previously unpublished). Deretaphrus have historically been found under the bark of Eucalyptus (e.g., Eucalyptus amplifolia Naudin , E. diversicolor F. von Mueller, E. mannifera Mudie , E. tereticornis Smith , E. viminalis Labill. , Myrtaceae ), Acacia (e.g., Acacia anceps DC. , A. mearnsii De Wild. , Fabaceae ), Vachellia bidwillii (Benth.) Kodela (Mimosaceae) , Ficus macrophylla Desfontaines ex. Persoon ( Moraceae ), red cedar ( Toona ciliata M. Roem. , Meliaceae ), dogwood ( Jacksonia scoparia (Bonpl.) DC. , Fabaceae ), and Australian saltbrush ( Atriplex semibaccata R. Brown , Amaranthaceae ). Adults have also been collected from various sclerophyll plants and on the dead crowns of Kingia australis R. Brown (Dasypogonaceae) and Xanthorrhoea sp. (Xanthorrhoeaceae) .

The New Caledonian species, D. interruptus , has been collected in association with the larvae of Agrianome fairmairei Montrouzier ( Cerambycidae : Prioninae ) under the bark of Aleurites moluccana (L.) Willdenow ( Euphorbiaceae ) (C. Mille, personal communication).

There is one record of Deretaphrus adults sheltering inside the galls of rust fungi ( Uromycladium sp. , Pucciniomycetes) on Acacia in Australia ( D. viduatus , specimen UGCA 1NL). There is another record of an adult Deretaphrus being collected as an inquiline with ants. The tendency of specimens to occur under the bark of dead trees and logs may be a possible explanation for this association. Several Deretaphrus species were found to have small mites beneath their elytra, sometimes by the hundreds (previously unpublished). These mites are probably phoretic, a common occurrence in many wood-inhabiting beetles (Lawrence and Britton 1991).

In a study of adult Deretaphrus mandibles by utilizing Environmental Scanning Electron Microscopy (ESEM) equipped with backscatter detectors on uncoated specimens, elemental analysis via energy-dispersive spectroscopy identified the presence of zinc and chlorine in the apex of the mandibles of multiple Deretaphrus species ( Figs. 42–43 , NPL unpublished data). This discovery, along with the general shape of the mandibles, suggests that adult Deretaphrus are incorporating elements for strengthening or wear resistance. Furthermore, examination of the gut contents of dissected specimens did not reveal any traces of insect cuticle but contained fibrous material, implying that members are feeding on wood or other plant tissues as opposed to preying on soft-bodied insects as previously suspected.

Distribution. Deretaphrus has a disjunct distribution ( Figs. 173–182 ). The majority (22/25) of species are Australian, occurring in all states. Most specimens seem to occur with relative abundance in the dry sclerophyll regions, although a number have been collected in more subtropical habitats of Queensland. One species, D. interruptus , occurs in New Caledonia. One species, D. oregonensis , is the only described New World species and is moderately common in the western United States (California, Oregon, Washington, Idaho, Montana, Utah, Nevada) and British Columbia, Canada. The distribution of D. oregonensis appears to be limited to the western side of the Rocky Mountains. It is possible that D. oregonensis may occur in other western states (Wyoming, Colorado, Arizona, perhaps New Mexico), but there are currently no records from these localities. There is also a single specimen ( D. boliviensis new species) collected from the interior of Bolivia. The distribution of the genus is perhaps an old amphipolar distribution, with representatives occurring in both Northern and Southern Hemispheres (discussion of amphiopolar distributions in Crowson 1980). However, the Australasian, Nearctic, and Neotropical distribution is very odd. It is possible that D. boliviensis occurs in other countries in South America and perhaps Central America. The distributions of the Australian Deretaphrus species are by no means concrete. Over- and undercollecting in various regions of the continent have undoubtedly led to skewed distributions. It is highly probable that several species of Deretaphrus are much more widespread than current collection records indicate.

KEY TO THE SPECIES OF DERETAPHRUS NEWMAN

1. Eyes reniform, strongly excavated posteriorly by a large angular cuticular expansion of the head capsule; in dorsal view, eye and cuticular expansion appearing subtriangular and strongly projecting ( Figs. 40–41 ) (southern Australia)........ Deretaphrus ocularis Lord and McHugh , new species

1′. Eyes not reniform, not strongly excavated posteriorly, at most with a thickened strip ofcuticle at posterior margin; in dorsal view, eyes subtly to distinctly projecting, but not as above................ 2

2(1′). Pronotal disc with a weakly developed, shallow median longitudinal depression, or lacking depression; depression usually wide and lacking any defined lateral margins in central portion of disc (e.g., Figs. 62, 64 , 69 , 75, 77–78, 80 ). Note some D. carinatus ( Fig. 63 ) and D. xanthorrhoeae ( Fig. 81 ) may key here, but should go to 2′.............3

2′. Pronotal disc with a well-developed, impressed median longitudinal groove or canal; groove usually narrow, with defined lateral margins in at least part of central portion of disc (only basocentrally in some); groove often interrupted near anterior 1/3 to form smaller anterior depression, slit, or fovea (e.g., Figs. 57, 59, 65 , 71, 74 ) .................. 11

3(2). Antennae minutely setose, setae extremely short and nearly indiscernible, appearing glabrous unless at high magnification ( Fig. 44 ).................4

3′. Antennae with clearly visible setae ( Figs. 45–47 )....................................10

4(3). Elytron with only odd intervals (some or all) distinctly raised, carinate or tuberculate (5 or less intervals raised, not including sutural interval).........5

4′. Elytron with even and odd intervals (some or all) distinctly raised, carinate or tuberculate (6 or more intervals raised, not including sutural interval) ............. 6

5(4). Pronotum evenly convex, without median longitudinal depression/groove (if with slight depression at posterior margin, not distinctly visible in central portion of disc) ( Fig. 69 ); punctures on pronotal disc more or less uniform in size; supra-ocular ridges of head welldeveloped, laterally expanded over eyes, angulate, mostly concealing eyes from dorsal view; eyes slightly protuberant, evenly arcuate; raised elytral interstitial intervals separate and straight for entire length ( Figs. 106–107 ) (western Australia)....... Deretaphrus incultus Carter and Zeck

5′. Pronotum with shallow median longitudinal depression, visible in central portion of disc; punctures on pronotal disc deeper and slightly larger within median longitudinal depression; supraocular ridges of head weakly-developed, not or only slightly concealing eyes from dorsal view; eyes strongly protuberant, approaching pointed ( Fig. 38 ); elytral interstitial intervals 3 and 5 curved towards midline at apex, merging or nearly merging (western Australia) ............... Deretaphrus parviceps Lea

6(4′). Raised elytral interstitial intervals not continuous for entire length, repeatedly interrupted to form intermittent raised portions or tubercles (especially near apex) ( Figs. 96–97 , 128–129 )............7

6′. Raised elytral interstitial intervals continuous for entire length, not frequently interrupted...........................................8

7(6). Interrupted portions of elytral interstitial intervals forming numerous denticles near elytral apex ( Figs. 96–97 ); pronotal median longitudinal depression extremely weak, only distinct at base; pronotal width at mid-length nearly equal to pronotal length ( Fig. 64 ); orange to light red in color (eastern Australia) ............................................ ........ Deretaphrus erichsoni Newman

7′. Interrupted portions of elytral interstitial intervals not forming numerous denticles near elytral apex, instead forming raised or elevated segments (short carinae, Figs. 128–129 ); pronotal median longitudinal depression extending onto anterior 1/3 of pronotum; pronotal width at mid-length distinctly less than pronotal length ( Fig. 80 ); dark red to brown in color (eastern Australia) ........ Deretaphrus wollastoni Newman

8(6′). Pronotal median longitudinal depres-

sion shallow and wide (sometimes very

faint), visible at least in posterior half,

not distinctly tapering towards anterior

margin; punctures within pronotal

median longitudinal depression ran-

domly spaced along midline and

distinctly larger than on rest of disc

( Fig. 78 ) (western Australia) ................

....... Deretaphrus rodmani Lord and

McHugh, new species

8′. Pronotal median longitudinal depression narrower (sometimes very faint), parallel-sided or distinctly tapering towards anterior margin; punctures within median longitudinal depression present or absent, if present evenly spaced and not larger than on rest of disc....... 9

9(8′). Punctures on pronotal disc round, very shallow, closely set, giving pronotal disc an alveolate, or netlike appearance ( Fig. 58 ); dorsal surface of the head extremely flat, supra-ocular ridges not laterally expanded, eyes entirely visible from above; distal margins of antennal club segments straight, leading edge of club segments inflated, but rounded proximally; abdominal ventrite V simple (western Australia) ....... Deretaphrus alveolatus Carter and Zeck

9′. Punctures on pronotal disc oval, deeply impressed; punctures around apex of median pronotal longitudinal depression slightly angled outwards, punctures on rest of disc arranged longitudinally ( Fig. 60 ); dorsal surface of the head convex, supra-ocular ridges slightly

expanded laterally, evenly curved, at

least partially concealing eyes from above (see Figs. 37, 39 ); distal margins of antennal club segments slightly concave, leading edge of club segments inflated, angulate proximally and forming a distinct point; abdominal ventrite V with apex and posterior face impressed, slightly upturned (western Australia).... ..... Deretaphrus antennatus Lord and

McHugh, new species 10(3′). At elytral apex, interstitial intervals 3

and 5 curved inwardly, merging or nearly merging ( Figs. 122–123 ); elytral apex sinuate; with weak supra-ocular ridges, eye mostly visible from above (eastern Australia) ................................ ............. Deretaphrus puncticollis Lea

10′. Elytral interstitial intervals 3 and 5 separate for entire length; elytral apex evenly rounded, not sinuate; with well-developed, angulate supra-ocular ridges, partially or completely concealing the eye from above ( Fig. 35 ) (eastern Australia) ................................ ........ Deretaphrus bucculentus Elston

11(2′) Pronotal median longitudinal canal appearing as an oval depression at base and extending as a narrow groove to about middle of pronotum ( Fig. 72 ); canal never interrupted to form distinct anterior portion; central portion of disc evenly convex; oval depression at base of pronotum bordered on either side by a smaller dimple sub-medially; most of disc with evenly-sized, small punctures except at lateral margin, where punctures are distinctly larger and sparser; body orange.......................................... .............. Deretaphrus lateropunctatus Lord and McHugh , new species

11′ Pronotal median longitudinal canal lacking oval depression as above; canal generally narrow, clearly interrupted in anterior portion forming distinctly separate anterior and posterior portions, anterior portion always much smaller than posterior portion; central portion of disc impressed; punctures near lateral margin not distinctly larger and sparser than on remainder of pronotal disc; body red to black ............................................... 12

12(11′). Anterior portion of pronotal median longitudinal canal distinct, usually round or oval, forming a deep fovea, area around fovea depressed ( Figs. 59, 61 , 67, 70, 74 , 76 )...........................13

12′. Anterior portion of pronotal median longitudinal canal weakly developed, represented at most by a small slit or shallowly depressed area, never a deep fovea........................................18

13(12). Submentum more strongly produced anteriorly, entirely concealing mentum and base of palpi in ventral view (see Figs. 48–49, 52–53 ).........................14

13′. Submentum less strongly produced anteriorly, at least part of mentum and base of palpi visible in ventral view (see Figs. 50–51 )............................ 15

14(13). Posterior portion of pronotal median longitudinal canal gradually narrowing and diminishing posteriorly, closed at base ( Fig. 70 ); base of pronotum without distinct admedian depressions, lateral margin of canal at base not appearing raised and thickened; anterior portion of pronotal median longitudinal canal small, round to oval; denticles at basolateral angles of pronotal disc small; anterior margin of submentum sinuate ( Fig. 49 ); submentum with 2 pairs of setose pits, each pit bearing a long seta; abdominal ventrite V with numerous small punctures, each bearing a small, indistinct seta ( New Caledonia) ................ .... Deretaphrus interruptus Grouvelle

14′. Posterior portion of pronotal median longitudinal canal not narrowing or diminishing posteriorly, open at base; base of pronotum with a pair of admedian depressions, causing lateral margin of canal at base to appear raised and thickened; anterior portion of pronotal median longitudinal canal deep, elongate-oval; denticles at basolateral angles of pronotal disc large ( Fig. 67 ); anterior margin of submentum arcuate to subtly sinuate (see Fig. 53 ); submentum lacking paired setose pits; abdominal ventrite V with few large punctures, each bearing a distinct seta ( Fig. 136 ) (eastern Australia) ............. ........ Deretaphrus hoplites Lord and McHugh , new species

15(13′). Abdominal ventrite V with two depressions, one on each side of midline, resulting in a median V-shaped area near posterior margin (see Figs. 132–133 ), present in both sexes.......................... 16 15′. Abdominal ventrite V simple, without a depressions on each side of midline, not forming a median V-shaped area near posterior margin.......................17

16(15). Pronotum nearly smooth, with minute, scarcely impressed punctures; posterior portion of pronotal median longitudinal canal rounded ( Fig. 59 ); elytra appearing smooth, punctures scarcely impressed (although striae may be visible beneath surface of cuticle) (eastern Australia)..... .......................... Deretaphrus analis Lea

16′. Pronotum with small, impressed punctures; posterior portion of pronotal median longitudinal canal narrowing posteriorly ( Fig. 76 ); elytra with 9 rows of small, clearly impressed punctures (eastern Australia) ................................ ........... Deretaphrus piceus (Germar)

17(15′). Posterior portion of pronotal median longitudinal canal very deep, narrowing posteriorly; elytral punctures deep, very large; diameter of pronotal punctures variable; elytral interstitial intervals 3+5 merge and 7+9 merge before meeting apical margin ( Figs. 116–117 ) (western USA)...................................... .......... Deretaphrus oregonensis Horn

17′. Posterior portion of pronotal median longitudinal canal not very deep, narrowed in middle, swollen and rounded at each end; elytral punctures shallow, small; diameter of pronotal punctures uniform; elytral interstitial interval 3 ends before apex and 5+7+9 merge before meeting apical margin ( Figs. 90–91 ) ( Bolivia)........................... ....... Deretaphrus boliviensis Lord and McHugh , new species

18(12′). Antennal club segments prominent, fairly dense setation (see Figs. 45, 47 ); pronotal disc with small, shallowly impressed punctures; ( Figs. 66, 71 ); medial portion of elytra nearly smooth, with intervals flat to very subtly raised, (weakly raised only near base and apex); submentum as in Fig. 51 ................... 19

18′. Antennal club segments with or without prominent setae, but setation never dense; pronotal disc with medium to large, moderately to deeply impressed punctures; medial portion of elytra with distinctly raised interstitial intervals; submentum not as in Fig. 51..............20

19(18). Posterior portion of pronotal median longitudinal canal sharply impressed; anterior portion of canal distinct, surrounding area depressed; lateral margins of pronotal disc strongly deflexed; dorsal surface of pronotal disc medially impressed ( Fig. 66 ) (western Australia) ......... Deretaphrus gracilis Blackburn

19′. Posterior portion of pronotal median longitudinal canal shallow, not strongly impressed; anterior portion of canal feebly marked, surrounding area weakly depressed; lateral margins of pronotal disc not strongly deflexed; dorsal surface of pronotal disc very flat ( Fig. 71 ) (eastern Australia) ................. .... Deretaphrus iridescens Blackburn

20(18′). Head with well-developed supraocular ridges, eyes at least partially concealed from above (see Figs. 35–37, 39–40 )................................................... 21

20′. Head without well-developed supraocular ridges; eyes entirely visible from above .................................................. 22

21(20). At elytral midlength, elytral interstitial intervals 3–9 distinctly carinate, degree of carination subequal; eyes strongly protuberant, distinctly convex ( Fig. 36 ) ........ Deretaphrus carinatus Lord and McHugh , new species

21′. At elytral midlength, elytral interstitial intervals 3–9 carinate, but intervals 4, 6, and 8 less prominent than 3, 5, 7, 9 (sometimes only subtly so); interval 8 nearly flat; eyes weakly protuberant, only slightly convex (eastern Australia) ............... Deretaphrus ignarus Pascoe

22(20′). Anterior angles of pronotum rounded, not well-developed; elytral interstitial interval 6 not raised, or raised for nearly entire length........................................23

22′. Anterior angles of pronotum welldeveloped, with a distinct angle ( Fig. 65 ); elytral interstitial interval 6 distinctly raised for only a short distance in posterior half of elytra ( Figs. 98–99 ) (eastern Australia) ........ ............. Deretaphrus fossus Newman

23(22). At elytral midlength, intervals 3, 4, and 6 not distinctly raised; setae on antennal club segments moderate and longer, reaching middle of following segment; abdominal ventrite V with posterior margin entire, simple; usually black in color (eastern Australia)...................24

23′. At elytral midlength, intervals 3, 4, and 6 distinctly raised; setae on antennal club segments very sparse and short, not reaching middle of following segment; ventrite V with posterior margin excised in females ( Figs. 138–139 ); dark red in color (western Australia)... ........ Deretaphrus xanthorrhoeae Lea

24(23). Head with anterior portion bearing a median longitudinal impression, most prominent near frontoclypeal suture. Posterior portion of pronotal median longitudinal canal deeply impressed, nearly parallel-sided for entire length. Pronotum with anterior angles broadly rounded ( Fig. 79 ); disc depressed medially. Elytral interstitial interval 3 only feebly raised at base and apex (eastern Australia) ................................ ............. Deretaphrus viduatus Pascoe

24′. Head with anterior portion evenly convex. Posterior portion of pronotal median longitudinal canal distinctly wider at base and tapering anteriorly, (appearing as an isosceles triangle), shallow. Pronotum with anterior angles pronounced, nearly forming right angles ( Fig. 57 ); disc evenly convex. Elytral interstitial interval 3 distinctly raised for entire length, carinate (eastern Australia)...................................... ... Deretaphrus aequaliceps Blackburn

Figures

Figs. 18–24. Deretaphrus fossus, head morphology. 18) Head, dorsal view; 19) Head, lateral view; 20) Head, ventral view; 21) Labrum, ventral view; 22) Maxilla, ventral view; 23) Left mandible, dorsal view; 24) Labium, ventral view.

Figs. 1–3. Deretaphrus fossus, neotype. 1) Dorsal habitus; 2) Ventral habitus; 3) Lateral habitus.

Figs. 48–53. Deretaphrus species, head capsule, ventral view, showing setae and shape of submentum. 48) D. ignarus;

Figs. 35–41. Deretaphrus species, head. 35) D. bucculentus, dorsal view; 36) D. carinatus, dorsal view; 37) D. incultus, dorsal view; 38) D. parviceps, dorsal view; 39) D. rodmani, dorsal view; 40) D. ocularis, dorsal view; 41) D. ocularis, lateral view.

Figs. 44–47. Deretaphrus species, antenna showing setal density. 44) D. erichsoni, right antenna; 45) D. interruptus, right antenna; 46) D. puncticollis, left antennal club; 47) D. interruptus, left antennal club.

Figs. 57–65. Deretaphrus species, pronotum. 57) D. aequaliceps; 58) D. alveolatus; 59) D. analis; 60) D. antennatus;

Figs. 75–81. Deretaphrus species, pronotum. 75) D. parviceps; 76) D. piceus; 77) D. puncticollis; 78) D. rodmani;

Figs. 66–74. Deretaphrus species, pronotum. 66) D. gracilis; 67) D. hoplites; 68) D. ignarus; 69) D. incultus;

Figs. 54–56. Deretaphrus species, morphology. 54) D. puncticollis, prosternum; 55) D. erichsoni, right protibia and protarsus; 56) D. erichsoni, right mesotibial spurs.

Figs. 25–28. Deretaphrus fossus, thorax and abdomen morphology. 25) Metathoracic wing; 26) Metendosternite, ventral view; 27) Pterothorax, ventral view; 28) Abdominal ventrites I–V, ventral view.

Figs. 82–87. Deretaphrus species, elytra, dorsal and lateral views. 82–83) D. aequaliceps; 84–85) D. alveolatus;

Figs. 88–93. Deretaphrus species, elytra, dorsal and lateral views. 88–89) D. antennatus; 90–91) D. boliviensis;

Figs. 94–99. Deretaphrus species, elytra, dorsal and lateral views. 94–95) D. carinatus; 96–97) D. erichsoni;

Figs. 100–105. Deretaphrus species, elytra, dorsal and lateral views. 100–101) D. gracilis; 102–103) D. hoplites;

Figs. 106–111. Deretaphrus species, elytra, dorsal and lateral views. 106–107) D. incultus; 108–109) D. interruptus;

Figs. 112–117. Deretaphrus species, elytra, dorsal and lateral views. 112–113) D. lateropunctatus; 114–115) D. ocularis;

Figs. 118–123. Deretaphrus species, elytra, dorsal and lateral views. 118–119) D. parviceps; 120–121) D. piceus;

Figs. 124–129. Deretaphrus species, elytra, dorsal and lateral views. 124–125) D. rodmani; 126–127) D. viduatus;

Figs. 130–131. Deretaphrus species, elytra. 130) Dorsal view; 131) Lateral view.

Figs. 132–139. Deretaphrus species, abdominal apex. 132) D. analis, ventral view (either sex); 133) D. piceus, ventral view (either sex); 134) D. erichsoni, female, ventral view; 135) D. erichsoni, female, oblique lateral view; 136) D. hoplites, female, ventral view; 137) D. xanthorrhoeae, male, ventral view; 138) D. xanthorrhoeae, female, ventral view; 139) D. xanthorrhoeae, female, posterior view.

Figs. 29–31. Generalized male terminalia and genitalia of Deretaphrus. 29) Abdominal terminalia; 30) Aedeagus;

Figs. 140–148. Deretaphrus species, phallobase. 140) D. aequaliceps; 141) D. alveolatus; 142) D. analis;

Figs. 149–157. Deretaphrus species, phallobase. 149) D. gracilis; 150) D. hoplites; 151) D. ignarus; 152) D. incultus;

Figs. 158–164. Deretaphrus species, phallobase. 158) D. parviceps; 159) D. piceus; 160) D. puncticollis;

Figs. 165–172. Deretaphrus species, penis (left: dorsal view; right: lateral view; phallobase, to scale with penis). 165) D. boliviensis; 166) D. bucculentus; 167) D. gracilis; 168) D. hoplites; 169) D. ignarus; 170) D. oregonensis; 171) D. parviceps; 172) D. puncticollis.

Figs. 32–33. Generalized female terminalia and genitalia of Deretaphrus. 32) Abdominal terminalia and ovipositor;

Fig. 34. Generalized female reproductive tract of Deretaphrus, ovaries removed.

Figs. 6–11. Deretaphrus biology. 6) Series of D. viduatus depicting size variability; 7) Pupal cell of Trichaulax philipsii; 8) D. puncticollis, pupal cocoon mass (with emerging adult) attached to larva of T. philipsii; 9) D. puncticollis, with pupal cocoon; 10) D. xanthorrhoeae, pupal cocoon; 11) D. viduatus, pupal cocoon with silken threads and smooth external surface.

Figs. 12–13. Deretaphrus xanthorrhoeae, pupal cocoons. 12) Dorsal view; 13) Ventral view.

Figs. 14–15. Deretaphrus species pupal cocoon. 14) D. puncticollis, 50X; 15) D. viduatus, 70X.

Figs. 16–17. Deretaphrus xanthorrhoeae, pupal cocoon. 16) 50X; 17) 100X.

Figs. 42–43. Deretaphrus species, head showing clypeal emargination, mandibular setae, and areas of metal incorporation. 42) D. interruptus, anterior view; 43) D. piceus, anterior view.

Figs. 173–174. Distribution of Deretaphrus species. 173) D. alveolatus, D. antennatus, D. parviceps, and D. rodmani;

Figs. 175–176. Distribution of Deretaphrus species. 175) D. fossus and D. ignarus; 176) D. piceus and D. viduatus.

Figs. 177–178. Distribution of Deretaphrus species. 177) D. aequaliceps, D. erichsoni, D. iridescens, and D. ocularis;

Figs. 179–180. Distribution of Deretaphrus species. 179) D. analis, D. bucculentus, D. hoplites, and D. lateropunctatus;

Figs. 181–182. Distribution of Deretaphrus species. 181) D. boliviensis; 182) D. oregonensis.