Three new genera of Indo-Australian Dermestidae (Coleoptera) and their phylogenetic significance

We are grateful to T. Kiselova and J. McHugh for allowing us to examine an unpublished version of their manuscript and to J. Háva for sending a copy of his recent dermestid catalogue. The following institutions and individuals are acknowledged for allowing us to study specimens in their care: Albert Allen, Boise, Idaho, USA; D. Kavanaugh, R. Brett, California Academy of Sciences, San Francisco, California, USA; L. LeSage, Canadian National Collections of Insects, Arachnids and Nematodes, Agriculture Canada; N. Vandenberg, National Museum of Natural History, Smithsonian Institution and Systematic Entomology Laboratory, United States Department of Agriculture; R. A. B. Leschen, New Zealand Arthropod Collection, Landcare Research, Auckland, New Zealand; G. Monteith, G. Thompson, Queensland Museum, Brisbane, Qld; E. G. Matthews, South Australian Museum, Adelaide, SA. Larvae and adults of Orphilodes malleecola were collected as part of an Australian National Insect Collection insect survey of Brookfield Conservation Park and with the aid of a grant from from the Conservation and Research Committee (SEACOM) of the Chicago Zoological Society and logistics support from J. Stelman, S. Williams and P. Parker. E. Hines and the CSIRO Black Mountain Electron Microscopy Laboratory are acknowledged for their assistance in the production of scanning electron micrographs and digital camera images enhanced with automontage.

Aitken A. D. (1975). Insect Travellers. Volume I. Coleoptera. Ministry of Agriculture, Fisheries and Food, Agricultural Development and Advisory Service, Pest Infestation Control Laboratory, Technical Bulletin 31, London, UK.

Anderson W. H. (1949) Larval description and transfer of Thaumaphrastus karanisensis from Colydiidae to a new subfamily of Dermestidae (Coleoptera). Bulletin of the Brooklyn Entomological Society 44, 121–127. and Vijay Veer and Rao (2000).

Eucinetus Germar, 1818 (Eucinetidae). Based on adults and larvae of Eucinetus morio LeConte.

Hexanodes Blair, 1941 (Dermestidae : Trinodinae : Trichelodini). Based on adults and larvae of Hexanodes vulgata Blair, and notes in Peacock (1978) and Zhantiev (2000). Although Peacock (1978) synonymised this genus with Trichelodes Carter, the differences between adults of H. vulgatus and T. delicatula appear to be far greater than those separating other trinodine genera; thus, the two were coded separately for the analysis in order to test the monophyly of the combined taxon.

Mariouta Pic, 1898 (Dermestidae : Marioutinae : Marioutini). Based on adults of Mariouta stangei Reitter, and notes in Mroczkowski and Slipinski (1997) and Zhantiev (2000).

Nosodendron Latreille, 1804 (Nosodendridae). Based on adults and larvae of Nosodendron unicolor Say and Nosodendron sp. (from Queensland, Australia) and notes in Beutel (1996) and Costa et al. (1986).

Novelsis Casey, 1900 (Dermestidae : Attageninae). Based on adults of Novelsis picta Casey and notes in Rees (1943).

Orphilodes, gen. nov. Based on adults of three described species and putative larvae of O. australis, sp. nov., O. malleecola, sp. nov. and an undescribed species.

Orphilus Erichson, 1846 (Dermestidae : Orphilinae). Based on adults and larvae of Orphilus subnitidus LeConte, and notes in Paulian (1942) and Beal (1985).

Rhopalosilpha Arrow, 1929 (Dermestidae : Marioutinae : Rhopalosilphini). Based on adults of Rhopalosilpha wasmanni Arrow and notes in Mroczkowski and Slipinski (1997) and Zhantiev (2000).

Rhyzopertha Stephens, 1830 (Bostrichidae : Dinoderinae). Based on adults and larvae of Rhyzopertha dominica (Fabricius).

Thorictodes Reitter, 1875 (Dermestidae : Thorictinae : Thaumaphrastini). Based on adults of Thorictodes heydeni Reitter and notes in van Emden (1924), Anderson (1949) and Zhantiev (2000).

Thorictus Germar, 1834 (Dermestidae : Thorictinae : Thorictini). Based on adults of Thorictus castaneus Germar from Egypt and notes in Reichensperger (1926), Banck (1927) and Zhantiev (2000).

Thylodrias Motschulsky, 1839 (Dermestidae : Thylodriinae). Based on adults and larvae of Thylodrias contractus Motschulsky and notes in Rees (1943), Franciscolo (1975), Süss and Fogato (1978) and Zhantiev (2000).

Trichelodes Carter, 1935 (Dermestidae : Trinodinae : Trichelodini). Based on adults and larvae of Trichelodes delicatula Carter and notes in Peacock (1978).

Trichodryas, gen. nov. Based on adults only of T. esoterica, sp. nov. and undescribed species.

Trinodes Dejean, 1821 (Dermestidae : Trinodinae : Trinodini). Based on adults of Trinodes hirtus (Fabricius), a larva of Trinodes sp. (from Keeling, Taiwan), and notes on larvae in Korschefsky (1944) and Zhantiev (2000).

Trogoderma Berthold, 1827 (Dermestidae : Megatominae : Megatomini). Based on adults and larvae of Trogoderma granarium Everts, adults of T. apicipenne Reitter and notes in Rees (1943), Korschefsky (1944), Hinton (1945), Beal (1991) and Zhantiev (2000).

(1) Upper body surfaces: (0) not clothed with stiff, erect, dark bristles; (1) clothed with stiff, erect, dark bristles.

These bristles may also be dehiscent, with a narrowed base that is easily broken or bent; as a result they are often inclined in various directions.

(2) Head: (0) prognathous, not or only slightly declined; (1) moderately to strongly declined.

This refers only to the position of the base of the head relative to the prothorax; some prognathous heads may be strongly declined anteriorly, so that the mouthparts are ventral.

(3) Median occipital endocarina: (0) absent; (1) present.

This begins at the edge of the occipital foramen and extends anteriorly.

(4) Median ocellus: (0) absent; (1) present.

The median ocellus is coded as present in Hexanodes, although it may be very weakly developed, and absent in Trichelodes, which is certainly a secondary loss.

(5) Frontoclypeal region: (0) slightly to moderately, gradually declined; (1) strongly and abruptly declined.

(6) Mandibular base: (0) distinctly wider than antennal scape; (1) distinctly narrower than antennal scape.

(7) Mandibular apex: (0) bidentate; (1) unidentate or rounded.

(8) Mandibular mola: (0) present; (1) absent.

(9) Mandibular prostheca: (0) present; (1) absent.

The prostheca usually refers to a hyaline lobe, often lined with hairs, a brush of hairs or both, which lie immediately distal to the mandibular mola. Such a structure occurs in all mandibles with a mola (e.g. Derodontus, Nosodendron and Orphilus), but a similar structure, here considered as homologous, also occurs in several taxa that lack a basal mola (e.g. Dermestes, Derbyana, Mariouta, and Orphilodes).

(10) Mesodorsal edge of mandibular base: (0) without projecting brush; (1) with projecting brush.

(11) Galea: (0) much broader than subapical maxillary palpomere; (1) subequal to or narrower than subapical maxillary palpomere.

(12) Lacinia: (0) subequal in width to galea; (1) distinctly narrower than galea; (2) highly reduced or apparently absent.

In Thylodrias, the lacinia is a short, basal lobe that is clearly set off from the galea; in Trichodryas this lobe is fused to the base of galea. Only the latter has been coded as state 2.

(13) Inner edge of lacinia: (0) without acute, glabrous, hyaline lobe; (1) with acute, glabrous, hyaline lobe.

(14) Lacinial apex: (0) with uncus; (1) without uncus.

(15) Labial palp articulation: (0) exposed; (1) concealed by mentum.

(16) Corporotentorium: (0) present; (1) absent.

(17) Laminatentorium: (0) complete; (1) incomplete at midline; (2) absent (anterior arms not widened).

(18) Posterior pronotal angles: (0) obtuse or rounded; (1) more-or-less acute.

(19) Lateral portions of pronotal disc: (0) not tuberculate; (1) tuberculate.

(20) Base of pronotal disc: (0) without sublateral impressions or grooves; (1) with sublateral impressions or grooves.

(21) Pronotal disc: (0) without sublateral carinae; (1) with sublateral carinae.

(22) Pronotal hypomeron: (0) without impression or cavity; (1) with broad, shallow impression; (2) with discrete antennal cavity.

(23) Inner edge of pronotal hypomeron: (0) without narrow antennal groove; (1) with narrow antennal grove.

(24) Prosternum: (0) without or with narrow head rest; (1) with broad head rest.

The head rest is a shiny infolding of the anterior edge of the prosternum. It has been called the prosternal collar by Zhantiev (2000).

(25) Prosternal process: (0) complete, extending behind coxae; (1) incomplete or absent.

(26) Procoxa: (0) less than twice as long as wide, not or slightly projecting ventrally; (1) more than twice as long as wide, strongly projecting ventrally.

(27) Procoxa: (0) without distinct lamina or plate (may be carinate); (1) with distinct lamina or plate.

(28) Procoxa: (0) without secondary mesal articulation; (1) with secondary mesal articulation.

This secondary articulation involves a pair of lateral projections at the apex of the prosternal process, each of which fits into a notch near the procoxal apex.

(29) Protrochantin: (0) at least slightly exposed; (1) completely concealed.

(30) Procoxal cavities internally: (0) open; (1) closed.

(31) Base of scutellum: (0) not or only slightly or gradually elevated; (1) distinctly, abruptly elevated.

(32) Elytral epipleuron: (0) complete; (1) incomplete.

(33) Mesoventral procoxal rests: (0) absent; (1) present.

(34) Anterior edge of mesoventrite: (0) on same plane as metaventrite; (1) on different plane than metaventrite.

(35) Mesoventrite: (0) separated by sutures from mesepisterna; (1) fused to mesepisterna.

(36) Mesoventrite: (0) without cavity for reception of prosternal process; (1) with cavity for reception of prosternal process.

(37) Anterior portion of mesoventrite: (0) neither tumid nor carinate; (1) tumid or carinate.

This refers to the so-called body of the ventrite, as opposed to the mesoventral process, which extends between the mesocoxae.

(38) Mesocoxa: (0) without secondary mesal articulation of dermestine type; (1) with secondary mesal articulation of dermestine type.

This type of secondary articulation, which was first recognised by Lawrence and Newton (1995), involves a mesal impression near the apex of the mesocoxa and a lateral projection near the apex of the metaventral process.

(39) Mesocoxa: (0) without secondary mesal articulation of orphiline type; (1) with secondary mesal articulation of orphiline type.

This type of secondary articulation involves a mesal projection near the apex of the mesocoxa, which fits into the side of the mesoventral process near its apex.

(40) Mesocoxal cavities: (0) contiguous or separated by less than half the shortest diameter of a cavity; (1) separated by more than half the shortest diameter of a cavity.

(41) Metathoracic discrimen: (0) present; (1) absent.

(42) Exposed portion of metanepisternum: (0) moderately narrow (more than 3 times as long as wide); (1) very broad (more than 3 times as long as wide).

(43) Metacoxa: (0) strongly transverse; (1) slightly transverse; (2) more-or-less globular.

(44) Metacoxae: (0) separated by less than 0.25 times coxal width; (1) separated by more than 0.25 times coxal width.

(45) Lateral metacoxal articulation: (0) exposed or barely concealed by elytral epipleuron; (1) concealed by metanepisternum.

(46) Metacoxa: (0) meeting elytral epipleuron; (1) separated from elytral epipleuron.

(47) Metacoxal plates: (0) well developed and complete to lateral edge of coxa; (1) well developed mesally but absent laterally; (2) very weak or absent.

(48) Radial cell of hind wing: (0) well developed; (1) reduced and almost obliterated by pigment; (2) absent.

(49) Cross-vein r3: (0) strongly oblique, almost perpendicular to long axis of wing; (1) not or slightly oblique, more-or-less parallel to long axis of wing; (2) absent.

(50) Posterior radial vein: (0) not extending basally beyond mid wing; (1) extending at least to basal third of wing.

(51) Wedge cell of hind wing: (0) present; (1) absent.

(52) Number of free veins in medial field of hind wing: (0) five; (1) four; (2) three or fewer.

The presence of five free veins in the medial field (MP3, MP4+ CuA1, CuA2, AA3 and AA4) appears to be the primitive condition in this group, and possibly in all Coleoptera, owing to the fusion of the medial and cubital systems. Species of Attagenus and related genera may have an additional apical forking, so that there are six free veins. Since these are rarely found in both wings and may involve either MP3 or CuA2, they are considered anomalous.

(53) Anal notch of hind wing: (0) present; (1) absent.

(54) Mesotibia: (0) not or slightly expanded; (1) strongly expanded at or near apex.

(55) Outer edge of mesotibia: (0) simple; (1) crenulate; (2) armed with socketed spines and sometimes teeth.

(56) Tibial spurs: (0) well developed; (1) absent or reduced and not apparent.

(57) First meso- and metatarsomere: (0) not or only slightly shorter than second; (1) much shorter than second.

(58) First and second ventrite: (0) free; (1) more-or-less connate.

(59) Abdominal intercoxal process: (0) subacute to narrowly rounded; (1) broadly rounded to truncate; (2) absent.

(60) Sternite III: (0) not as long as IV and V combined; (1) at least as long as IV and V combined.

(61) Sternite III: (0) without pair of oblique grooves and pits; (1) with paired oblique grooves arising from deep pits.

(62) Sternite III in male: (0) without median patch of setae and glandular pores; (1) with median match of setae and glandular pores.

(63) Sternites V and VI in male: (0) without median patches of setae and glandular pores; (1) with median patches of setae and glandular pores.

(64) Apex of tergite IX in male: (0) not or shallowly emarginate; (1) deeply emarginate, tergite X distinct; (2) deeply emarginate, partly fused to tergite X; (3) completely fused to and indistinguishable from tergite X.

(65) Phallobase: (0) articulating with bases of parameres; (1) overlapping bases of parameres.

(66) Parameres: (0) not joined together dorsally; (1) joined together dorsally.

(67) Parameres: (0) not joined together ventrally; (1) joined together ventrally.

(68) Malpighian tubules: (0) free; (1) of modified cryptonephridial type.

The modified type of cryptonephridism is one in which all Malpighian tubules are reunited with the gut in a single bundle (Saini 1964). This is usually indicated in cleared dry specimens of bostrichoid adults by the presence of a rectal bar, which supports the perinephral sac. In adult dissections of Orphilus and Orphilodes species, the typical rectal bar could not be seen, although a light sclerotisation was present that could have been homologous to that structure. Orphilus was coded as having the modified type of cryptonephridism based on the illustration of the larval gut in Kiselova and McHugh (2005), whereas Orphilodes was coded as unknown for this character.

(69) Larva: body: (0) with sclerotised tergites; (1) without sclerotised tergites.

(70) Larva: body: (0) more-or-less fusiform; (1) C-shaped and grub-like.

(71) Larva: dorsal surfaces: (0) with simple setae only; (1) with at least some setae of a modified type.

Modified setae in Dermestidae include: those that are apically expanded or frayed; those that are flattened, expanded and ridged; spicisetae, which may be slender or stout and linear or apically expanded and club-like but always covered with spicules; and hastisetae, which have a spicisetal type of shaft with apex bearing recurved barbs.

(72) Larva: spicisetae: (0) absent; (1) present.

(73) Larva: major tergal spicisetae: (0) not widened subapically with narrowly acute apex; (1) slightly widened subapically with broadly acute apex; (2) strongly widened subapically with truncate apex.

(74) Larva: spicules on major tergal spicisetae: (0) not arranged in distinct vertical rows; (1) arranged in distinct vertical rows.

(75) Larva: flattened, ribbed setae: (0) absent; (1) present.

(76) Larva: abdominal apex: (0) without caudal brush of long hairs; (1) with caudal brush of long hairs.

(77) Larva: hastisetae: (0) absent; (1) present.

(78) Larva: frontal arms: (0) arising from base of head or from very short epicranial stem; (1) arising from moderately long epicranial stem; (2) absent.

(79) Larva: frontal arms: (0) lyriform; (1) V-shaped, U-shaped or absent.

(80) Larva: number of stemmata on each side: (0) five or six; (1) four; (2) three; (3) one; (4) none.

(81) Larva: sensorium on preapical antennomere: (0) less than 0.75 times as long as apical antennomere; (1) more than 0.75 times as long as apical antennomere.

(82) Larva: labroepipharyngeal margin: (0) without apically widened or spatulate setae; (1) with at least some apically widened or spatulate setae.

(83) Larva: paired epipharyngeal rods: (0) absent; (1) short, broad, parallel or forked; (2) long, slender, diverging posteriorly; (3) long, slender, parallel.

(84) Larva: basal row of epipharyngeal sensilla: (0) absent; (1) present.

(85) Larva: medial row of epipharyngeal sensilla: (0) absent; (1) present.

(86) Larva: sensilla in medial epipharyngeal row: (0) six; (1) four; (2) two.

(87) Larva: epipharyngeal sensilla in distal group: (0) none; (1) two; (2) four or more.

(88) Larva: mandible: (0) more-or-less evenly pigmented; (1) heavily pigmented apically, lightly so basally.

(89) Larva: mandibular apex: (0) tridentate; (1) bidentate; (2) unidentate, acute; (3) unidentate, rounded.

(90) Larva: mandibular mola: (0) large, complex; (1) reduced, simple; (2) absent.

(91) Larva: mandibular prostheca: (0) present; (1) absent.

This hyaline lobe is distad of the mola, when present; however, in some mandibles lacking a mola, a similar lobe occurs just distad of the penicillus. This is considered to be homologous to the prostheca.

(92) Larva: mandibular penicillus: (0) present; (1) absent.

This structure is always near the base of the mesal edge.

(93) Larva: accessory ventral process of mandible: (0) present; (1) absent.

(94) Larva: lacinial uncus: (0) at least tridentate; (1) bidentate; (2) unidentate.

(95) Larva: number of maxillary palpomeres: (0) four; (1) three.

(96) Larva: abdominal segments I–VIII: (0) without distinct laterotergites; (1) with distinct laterotergites.

Laterotergites are lateral or ventral, spiracle-bearing sclerites, separated by a carina or suture from the tergites and by a suture from the sternites (when the latter are also sclerotised).

(97) Larva: tergum IX: (0) with urogomphi; (1) without urogomphi. The paired, posteriorly-oriented processes on tergite IX in Orphilus are not considered to be homologous to urogomphi in other taxa.

(98) Larva: abdominal segment X: (0) without pair of longitudinally oval anal pads; (1) with pair of longitudinally oval anal pads.

(99) Larva: spiracles: (0) annular-biforous; (1) annular.

(100) Larva: first abdominal spiracles: (0) not placed at ends of spiracular tubes; (1) placed at ends of spiracular tubes.

(101) Larva: spiracles on abdominal segment VIII: (0) not at end of terminal siphon; (1) at end of terminal siphon.

(102) Larva: spiracles on abdominal segment VIII: (0) about the same size as anterior abdominal spiracles; (1) much larger than anterior abdominal spiracles.

(103) Pupa: (0) free from larval skin, which is shed during pupation; (1) lying partly within larval skin, which is not shed at pupation. If pupation is known to occur in soil, the taxon is coded as having state 0.

(104) Pupa: urogomphi: (0) present; (1) absent.

U = unambiguous changes only; A and D = additional changes using accelerated or delayed transformation, respectively

Thylodrias + Trichodryas (U8): 6,1 – mandible narrower than scape; 7,1 – mandible unidentate; 16,1 – corporotentorium absent; 17,2 – laminatentorium absent; 31,0 – scutellum not elevated; 33,0 – procoxal rests absent; 47,2 – metacoxal plates weak or absent; 59,2 – abdominal process absent; (A6): 12,1 – lacinia narrower than galea; 73,2 – spicisetae strongly widened and truncate; 80,2 – stemmata three; 89,3 – larval mandible unidentate, rounded; 95,1 – larval maxillary palpomeres three; 102,0 – larval eighth spiracles not enlarged.

Hexanodes clade (Hexanodes + Thylodrias + Trichodryas) (U2): 26,1 – procoxa elongate and projecting; 40,0 – mesocoxal cavities narrowly separated; (A2): 34,0 – mesoventrite on same plane as metaventrite; 73,1 – spicisetae slightly widened and broadly acute.

Trichelodes clade (Trichelodes + Hexanodes + Thylodrias + Trichodryas) (U5): 2,0 – head prognathous; 5,1 – frontoclypeal region strongly, abruptly declined; 19,1 – pronotal disc laterally tuberculate; 48,1 – radial cell reduced; 49,2 – cross-vein r3 absent; (A2): 24,0 – prosternum without head rest; 77,0 – hastisetae absent.

Trinodini (U7): 17,1 – laminatentorium incomplete; 21,1 – pronotum with sublateral carinae; 23,1 – hypomeron with narrow antennal groove; 44,1 – metacoxae widely separated; 45,1 – metacoxal articulation concealed; 59,1 – abdominal process rounded to truncate; 74,1 – spicules in vertical rows; (D2): 24,1 – prosternum with head rest; 34,1 – mesoventrite on different plane than metaventrite.

Trinodinae (U7): 1,1 – stiff, erect, dark bristles; 18,1 – posterior pronotal angles acute; 20,1 – pronotum with sublateral grooves; 25,1 – prosternal process incomplete or absent; 30,0 – procoxal cavities internally open; 42,0 – metanepisternum narrow; 102,1 – larval eighth spiracles enlarged; (A2): 14,0 – lacinia with uncus; 89,1– larval mandible bidentate; (D4): 9,1 –prostheca absent; 56,1 – tibial spurs absent or reduced; 81,1 – larval sensorium long; 95,0 – larval maxillary palpomeres four.

Megatominae (U7): 7,1 – mandible unidentate; 22,2 – hypomeron with antennal cavity; 29,1 – protrochantin concealed; 36,1 – mesoventrite with cavity; 58,1 – first two ventrites connate; 86,2 – epipharyngeal sensilla in medial row two; 87,3 – epipharyngeal sensilla in distal group six or more; (A3): 34,0 – mesoventrite on same plane as metaventrite; 81,0 – larval sensorium short; 95,1 – larval maxillary palpomeres three; (D4): 14,1 – lacinia without uncus; 24,1 – prosternum with head rest; 77,1 – hastisetae present; 89,3 – larval mandible unidentate, rounded.

Megatominae + Trinodinae (U5): 33,1 – procoxal rests present; 72,1 – spicisetae present; 91,1 – larval prostheca absent; 92,1 – larval penicillus absent; 103,1 – pupa within larval skin; (A7): 9,1– mandible bidentate; 24,1 – prosternum with head rest; 27,1 – procoxa with lamina; 37,0 – mesoventrite not carinate; 52,2 – veins in medial field three or fewer; 56,1 – tibial spurs absent or reduced; 77,1 – hastisetae present; (D2): 52,2 – veins in medial field three or fewer; 89,1 – larval mandible bidentate.

Orphilinae (U11): 12,1 – lacinia narrower than galea; 28,1 –secondary procoxal articulation; 39,1 – orphiline mesocoxal articulation; 46,0 – metacoxa long; 55,2 – mesotibia with socketed spines; 64,0 – tergite IX shallowly emarginate; 65,0 – phallobase articulated with parameres; 71,0 – larvae with simple setae; 83,1 – epipharyngeal rods short, broad; 90,1 – larval mola reduced, simple; 96,1 – distinct laterotergites; (A2): 80,3 – stemmata one; 88,0 – larval mandible evenly pigmented; (D5): 27,1 – procoxa with lamina; 34,1 – mesoventrite on different plane than metaventrite; 37,1 – mesoventrite carinate; 52,1 – medial field with four free veins; 81,1 – larval sensorium long.

Orphiline clade (Orphilinae + Megatominae + Trinodinae) (U5): 22,1 – hypomeron with shallow impression; 40,1 – mesocoxal cavities widely separated; 67,1 – parameres joined ventrally; 86,1 – epipharyngeal sensilla in medial row four; 104,1 – pupal urogomphi absent; (A3): 34,1 – mesoventrite on different plane than metaventrite; 52,1 – medial field with four free veins; 81,1 – larval sensorium long.

Attageninae (U7): 7,1 – mandible unidentate; 18,1 – posterior pronotal angles acute; 36,1 – mesoventrite with cavity; 57,1 – first meso- and metatarsomere short; 75,1 – flattened, ribbed setae; 76,1 – long caudal brush; 87,1 – epipharyngeal sensilla in distal group two; (D6): 14,1 – lacinia without uncus; 27,1 – procoxa with lamina; 37,1 – mesoventrite carinate; 88,1 – larval mandible lightly pigmented basally; 89,3 – larval mandible unidentate, rounded, rounded; 95,0 – larval maxillary palpomeres four.

Attagenine clade (Attageninae + Orphilinae + Megatominae + Trinodinae) (U3): 4,1 – median ocellus present; 42,1 – metanepisternum broad; 47,0 – metacoxal plates complete ; (A6): 14,1 – lacinia without uncus; 25,0 – prosternal process complete; 27,1 – procoxa with lamina; 29,0 – protrochantin exposed; 88,1 – larval mandible lightly pigmented basally; 89,3 – larval mandible unidentate, rounded; (D3): 2,1 – head declined; 51,1 – wedge cell absent; 78,1 – epicranial stem long.

Thorictinae (U11): 9,1 – mandible bidentate; 32,0 – epipleuron complete; 35,1 – mesoventrite fused to mesepisterna; 41,1 – metathoracic discrimen absent; 43,2 – metacoxa globular; 44,1 – metacoxae widely separated; 45,1 – metacoxal articulation concealed; 59,1 – abdominal process rounded to truncate; 60,1 – sternite III as long as IV and V combined; 69,1 – larva without sclerotised tergites; 80,4 – stemmata none; (A5): 78,0 – epicranial stem short; 89,1 – larval mandible bidentate; 92,1 – larval penicillus absent; 95,1 – larval maxillary palpomeres three; 104,1 – pupal urogomphi absent; (D1): 29,1 – protrochantin concealed.

Marioutinae (U3): 13,1 – lacinia with acute, hyaline lobe; 43,1 – metacoxa slightly transverse; 55,2 – mesotibia with socketed spines; (A2): 37,0 – mesoventrite not carinate; 52,1 – medial field with four free veins; (D2): 29,1 – protrochantin concealed; 51,1 – wedge cell absent.

Dermestinae (U2): 22,1 – hypomeron with shallow impression; 63,1 – setal patches on sternites V and VI; (A4): 29,0 – protrochantin exposed; 47,0 – metacoxal plates complete; 51,0 – wedge cell present: 64,0 – tergite IX shallowly emarginate.

Dermestinae + Marioutinae (U2): 11,0 – galea broader than subapical palpomere; 61,1 – oblique grooves on sternite III; (A3): 3,1 – occipital endocarina present; 7,1 – mandible unidentate; 72,1 – spicisetae present; (D3): 49,1 – cross-vein r3 slightly oblique; 50,1 – posterior radial vein extending to basal third; 53,1 – anal notch absent.

Dermestine clade (Thorictinae + Marioutinae + Dermestinae) (U4): 10,1 – mesodorsal edge of mandible with brush; 31,0 – scutellum not elevated; 38,1 –dermestine mesocoxal articulation; 97,0 – urogomphi present; (A4): 2,0 – head prognathous; 49,1 – cross-vein r3 slightly oblique; 50,1 – posterior radial vein extending to basal third; 53,1 – anal notch absent; (D1): 25,1 – prosternal process incomplete or absent.

Dermestidae (U4): 8,1 – adult mola absent; 71,1 – modified larval setae; 82,1 – epipharyngeal margin with spatulate setae; 85,1 – medial epipharyngeal row present; (A2): 37,1 – mesoventrite carinate; 90,2 – larval mola absent; (D5): 46,1 – metacoxa short; 79,1 – frontal arms V- or U-shaped; 83,2 – epipharyngeal rods long, slender, diverging; 90,1 – larval mola reduced, simple; 94,1 – larval uncus bidentate.

Bostrichoidea (U9): 30,1 – procoxal cavities internally closed; 64,3 – tergites IX and X fused; 65,1 – phallobase overlapping parameres; 66,1 – parameres joined dorsally; 68,1 – Malpighian tubules of modified cryptonephridial type; 84,0 – basal epipharyngeal row absent; 92,0 – larval penicillus present; 93,1 – larval mandible without accessory ventral process; 100,0 – larval spiracles not at ends of tubes; (A11): 2,1 – head declined; 25,1 – prosternal process incomplete or absent; 29,1 – protrochantin concealed; 46,1 – metacoxa short; 51,1 – wedge cell absent; 78,1 – epicranial stem long; 79,1 – frontal arms V- or U-shaped; 83,2 – epipharyngeal rods long, slender, diverging; 89,0 – larval mandible tridentate; 90,1 – larval mola reduced, simple; 94,1 – larval uncus bidentate; 95,0 – larval maxillary palpomeres four; 99,1 – larval spiracles annular; (D2): 17,0 – laminatentorium complete; 99,1 – larval spiracles annular.

Thylodrias + Trichodryas (U8): 6,1 – mandible narrower than scape; 7,1 – mandible unidentate ; 16,1 – corporotentorium absent ; 17,2 – laminatentorium absent ; 31,0 – scutellum not elevated; 33,0 – procoxal rests absent ; 47,2 – metacoxal plates weak or absent ; 59,2 – abdominal process absent; (A6): 12,1 – lacinia narrower than galea; 73,2 – spicisetae strongly widened and truncate; 80,2 – stemmata three; 89,3 – larval mandible unidentate, rounded; 95,1 – larval maxillary palpomeres three; 102,0 – larval eighth spiracles not enlarged .

Hexanodes clade (Hexanodes + Thylodrias + Trichodryas) (U2): 26,1 – procoxa elongate and projecting; 40,0 – mesocoxal cavities narrowly separated ; (A2): 34,0 – mesoventrite on same plane as metaventrite ; 73,1 – spicisetae slightly widened and broadly acute.

Trichelodes clade (Trichelodes + Hexanodes + Thylodrias + Trichodryas) (U5): 2,0 – head prognathous; 5,1 – frontoclypeal region strongly, abruptly declined; 19,1 – pronotal disc laterally tuberculate; 48,1 – radial cell reduced; 49,2 – cross-vein r3 absent; (A2): 24,0 – prosternum without head rest ; 77,0 – hastisetae absent.

Trinodini (U7): 17,1 – laminatentorium incomplete; 21,1 – pronotum with sublateral carinae; 23,1 – hypomeron with narrow antennal groove; 44,1 – metacoxae widely separated; 45,1 – metacoxal articulation concealed; 59,1 – abdominal process rounded to truncate; 74,1 – spicules in vertical rows; (A1): 16,1 – corporotentorium absent; (D2): 24,1 – prosternum with head rest; 34,1 – mesoventrite on different plane than metaventrite.

Trinodinae (U6): 1,1 – stiff, erect, dark bristles; 18,1 – posterior pronotal angles acute ; 20,1 – pronotum with sublateral grooves; 25,1 – prosternal process incomplete or absent; 30,0 – procoxal cavities internally open;102,1 – larval eighth spiracles enlarged; (A4): 7,0 – mandible bidentate; 34,1 – mesoventrite on different plane than metaventrite; 42,0 – metanepisternum narrow; 81,1 – larval sensorium long; (D5): 9,1 –prostheca absent; 22,1 – hypomeron with shallow impression; 56,1 – tibial spurs absent or reduced; 81,1 – larval sensorium long; 95,0 – larval maxillary palpomeres four.

Megatominae (U7): 14,1– lacinia with uncus; 29,1 – protrochantin concealed; 36,1 – mesoventrite with cavity; 58,1 – first two ventrites connate; 86,2 – epipharyngeal sensilla in medial row two; 87,3 – epipharyngeal sensilla in distal group six or more; 89,3 – larval mandible unidentate; (A2): 22,2 – hypomeron with discrete antennal cavity; 95,1 – larval maxillary palpomeres three; (D5): 7,1 – mandible unidentate or rounded; 22,2 – hypomeron with discrete antennal cavity; 24,1 – prosternum with head rest; 42,1 – metanepisternum broad; 77,1 – hastisetae present; 95,1 – larval maxillary palpomeres three.

Megatominae + Trinodinae (U5): 52,2 – veins in medial field three or fewer; 72,1 – larval spicisetae present; 91,1 – larval prostheca absent; 92,1 – larval penicillus absent; 103,1 – pupa within larval skin; (A4): 24,1 – prosternum with head rest; 33,1 – mesoventral procoxal rests present; 37,0 – mesoventrite not carinate; 56,1 – tibial spurs absent or reduced; 77,1 – hastisetae present; (D4): 33,1 – mesoventral procoxal rests present; 40,1 – mesocoxal cavities well separated; 89,1 – larval mandible bidentate .

Thorictinae (U11): 32,0 – epipleuron complete; 35,1 – mesoventrite fused to mesepisterna; 41,1 – metathoracic discrimen absent; 43,2 – metacoxa globular; 44,1 – metacoxae widely separated; 45,1 – metacoxal articulation concealed; 59,1 – abdominal process rounded to truncate; 60,1 – sternite III as long as IV and V combined; 69,1 – larva without sclerotised tergites; 78,0 –larval frontal arms arising near base of head ; 80,4 – stemmata none ; (A5): 7,0 – mandible bidentate; 89,1 – larval mandible bidentate; 92,1 – larval penicillus absent; 95,1 – larval maxillary palpomeres three; 104,1 – pupal urogomphi absent; (D3): 9,1 – prostheca absent; 29,1 – protrochantin concealed; 47,2 – metacoxal plates weak or absent.

Marioutinae (U3): 13,1 – lacinia with acute, hyaline lobe; 43,1 – metacoxa slightly transverse; 55,2 – mesotibia with socketed spines; (A2): 37,0 – mesoventrite not carinate; 52,1 – medial field with four free veins; (D2): 29,1 – protrochantin concealed; 47,2 – metacoxal plates weak or absent.

Dermestinae (U2): 51,0 – wedge cell present; 63,1 – setal patches on sternites V and VI; (A4): 22,1 – hypomeron with shallow impression; 29,0 – protrochantin exposed; 47,0 – metacoxal plates complete; 64,0 – tergite IX shallowly emarginate.

Dermestinae + Marioutinae (U2): 11,0 – galea broader than subapical palpomere ; 61,1 – oblique grooves on sternite III; (A4): 3,1 – occipital endocarina present; 9,0 –prostheca present ; 72,1 – spicisetae present; 89,0 – larval mandible tridentate; (D3): 49,1 – cross-vein r3 slightly oblique; 50,1 – posterior radial vein extending to basal third; 53,1 – anal notch absent.

Dermestine clade (Thorictinae + Marioutinae + Dermestinae) (U7): 2,0 – head prognathous; 4,0 – median ocellus absent; 10,1 – mesodorsal edge of mandible with brush; 25,1 – prosternal process incomplete or absent; 31,0 – scutellum not elevated; 38,1 –dermestine mesocoxal articulation; 97,0 – urogomphi present; (A7): 29,1 – protrochantin concealed; 42,0 – metanepisternum narrow; 47,2 – metacoxal plates weak or absent; 49,1 – cross-vein r3 slightly oblique; 50,1 – posterior radial vein extending to basal third; 53,1 – anal notch absent; 88,0 – larval mandible evenly pigmented .

Attageninae (U9): 14,1 – lacinia without uncus; 18,1 – posterior pronotal angles acute; 27,1 – procoxa with distinct lamina; 36,1 – mesoventrite with cavity; 57,1 – first meso- and metatarsomere short; 75,1 – flattened, ribbed setae; 76,1 – long caudal brush; 87,1 – epipharyngeal sensilla in distal group two; 89,3 – mandibular apex unidentate, rounded; (D5): 7,1 – mandible unidentate or rounded; 37,1 – mesoventrite carinate; 42,1 – metanepisternum broad; 88,1 – larval mandible lightly pigmented basally; 95,0 – larval maxillary palpomeres four.

Attagenine-dermestine clade (U2): 67,0 – parameres not joined together ventrally ; 86,0 – larval epipharyngeal sensilla in medial row six. (A3): 22,0 – hypomeron without impression; 40,0 – mesocoxal cavities contiguous; 104,0 – pupal urogomphi absent .

Non-orphiline dermestid clade (U5): 46,1 – metacoxa separated from elytral epipleuron; 55,0 – outer edge of mesotibia simple; 71,1 – larvae with at least some modified setae; 83,2 – larval epipharyngeal rods long, slender, diverging; 90,2– larval mola absent; (A5): 7,1 – mandible unidentate or rounded; 9,1 – prostheca absent; 80,0 – larval stemmata five or six ; 81,0 – larval sensorium short ; 88,1 – larval mandible lightly pigmented basally; (D3): 8,1 – mola absent; 64,3 – tergite IX in male fused to X; 65,1 – phallobase overlapping parameres.

Orphilinae (U7): 12,1 – lacinia narrower than galea; 27,1 – procoxa with distinct lamina; 28,1 –secondary procoxal articulation; 34,1 – mesoventrite on different plane than metaventrite; 39,1 – orphiline mesocoxal articulation; 52,1 –medial field with four free veins; 96,1 – distinct laterotergites; (A2): 64,0 – tergite X in male not emarginate; 65,0 – phallobase articulating with parameres; (D5): 22,1 – hypomeron with shallow impression; 37,1 – mesoventrite carinate; 40,1 – mesocoxal cavities widely separated; 42,1 – exposed portion of metanepisternum broad; 81,1 – larval sensorium long.

Dermestidae (U3): 4,1 – median ocellus present; 82,1 – epipharyngeal margin with spatulate setae; 85,1 – medial epipharyngeal row present; (A4): 8,1 – mola absent; 37,1 – mesoventrite carinate; 42,1 – metanepisternum very broad; 98,0 – larval segment X without longitudinal pads; (D5): 51,1 – wedge cell absent; 78,1 – larval frontal arms arising from long stem; 79,1 – frontal arms V- or U-shaped; 94,1 – larval uncus bidentate.

Endecatomus + Rhyzopertha (U8): 3,1 – median occipital endocarina present ; 25,1 – prosternal process incomplete or absent ; 29,1 – protrochantin concealed ; 47,2 – metacoxal plates weak or absent ; 69,1 – larva without sclerotised tergites ; 70,1 – larva C-shaped and grub-like; 83,3– larval epipharyngeal rods long, slender, parallel; 89,0 – larval mandible tridentate ; (A4): 22,1 – hypomeron with shallow impression ; 40,0 – mesocoxal cavities contiguous ; 78,2 – larval frontal arms absent; 94,2 – larval uncus unidentate; (D6): 64,3 – tergite IX in male fused to tergite X ; 65,1 – phallobase overlapping parameres ; 78,2 – larval frontal arms absent; 81,1 – larval sensorium long ; 94,2 – larval uncus unidentate; 98,1 – larval segment X with oval pads .

Bostrichoidea (U10): 17,0 – laminatentorium complete; 30,1 – procoxal cavities internally closed; 66,1 – parameres joined dorsally; 67,1 – parameres joined together ventrally; 68,1 – Malpighian tubules of modified cryptonephridial type; 84,0 – basal epipharyngeal row absent; 90,1 – larval mola reduced, simple; 92,0 – larval penicillus present; 93,1 – larval mandible without accessory ventral process; 100,0 – larval spiracles not at ends of tubes; (A12): 2,1 – head declined; 51,1 – wedge cell absent; 64,3 – tergite X in male fused to IX; 65,1 – phallobase overlapping parameres; 78,1 – epicranial stem long; 79,1 – frontal arms V- or U-shaped; 80,3 – larval stemmata one; 81,1 – larval sensorium more than 0.75 × apical antennomere; 94,1 – larval uncus bidentate; 95,0 – larval maxillary palpomeres four; 99,1 – larval spiracles annular; 104,1 – pupal urogomphi absent; (D2): 2,1 – head declined; 99,1 – larval spiracles annular.