Phylogeny of the Gondwanan beetle family Ulodidae (Tenebrionoidea) Author Leschen, Richard A. B. Author Escalona, Hermes E. Author Elgueta, Mario text Zootaxa 2016 4138 3 441 473 journal article 10.11646/zootaxa.4138.3.2 c785c9ac-b71c-4487-b770-35ebfe797c8a 1175-5326 258464 DDF68118-EA96-49A5-B316-47814E19ED3D Ulodidae Pascoe, 1869 Adult diagnosis . Antennal insertions visible in dorsal view ( Fig. 9A ). Prosternal process strongly and abruptly expanded at apex and meeting the hypomeron, procoxal cavities externally closed ( Fig.10B ). Abdominal ventrites 1 and 2 connate ( Fig. 7K ). Tarsal formula 5-5-4 or 4-4-4 ( Meryx ). Aedeagus tenebrionoid ( Figs 8J–L ); tegmen without alae and anterior edge of penis without struts. Adult description . Length 3.8–24.5 mm . Body about 1.3–2.5 times as long as wide, slightly flattened to strongly convex; sides oval to parallel. Surfaces glabrous ( Arthopus ) or subglabrous ( Ulocyphaleus ), setose ( Meryx and Trachyderastes ) or clothed with distinct bristles or scale-like setae that are erect or adpressed to the cuticle; wax-debris encrustations present on specimens of Brouniphylax , Exohadrus , some Meryx , Notocerastes , Pteroderes , some Syrphetodes , and Ulobostrichus . Color mottled to unicolorous and sometimes metallic. Head prognathous, often deeply imbedded into prothorax, not abruptly constricted posteriorly to form a neck, but a ridge behind the eye may be present; temples short or absent; frons impressed or not. Eyes ( Figs 6A–F ) moderately to strongly ( Phaennis and Pteroderes ) protuberant, finely to coarsely facetted, usually entire, sometimes emarginate, rarely by a canthus or antennal insertion ( Trachyderastes ), posteriorly emarginate in Trachyderas , narrow in Ganyme and also strongly c-shaped in Waitomophylax ; interfacetal setae usually absent (present in Phaennis ). Antennal insertions exposed from above, or slightly concealed and inserted sublaterally and hidden by a marginal deflexed ridge or cephalic horns ( Notocerastes and Phaennis ); two anterolateral horns (male Phaennis , Fig. 6G ) or one short median ( Notocerastes ) horn present in the male. Frontoclypeal suture present or absent. Antennae usually extending posteriorly beyond basal margin of elytra (barely so in Pteroderes and not at all in Ulobostrichus ); 11-segmented, usually capitate with 3-segmented club (4-segmented in Trachyderastes ), that may be weakly clubbed, rarely filiform ( Trachyderastes , Ulocyphaleus ), a 10-segmented antenna with a weak 2- segmented clubbed is present in Ulobostrichus ; sensory areas of antennomeres (usually 9–11) demarcated by a distinct rim in some taxa. Labrum visible and free, transverse to subquadrate and not concealed beneath clypeus. Mandible short and broad, bidentate; incisor edge simple or with a single tooth with well-developed mola and prostheca. Maxilla ( Figs 7B–D ) with galea setose and lacinia with or without ( Meryx and Trachyderastes ) apical spines or uncus; apical palpomere cylindrical to fusiform, slightly expanded and truncate to subtriangulate and subcylindrical-curved ( Arthopus ). Submentum without setose pit; ligula undivided to moderately emarginate; prementum sclerotized or not; apical labial palpomeres cylindrical to fusiform with variable widths. Genal ridges or projections absent; ventral part of the head expanded laterally below eye and forming an ocular shelf in Brouniphylax and subocular tubercles present in Waitomophylax . Gular sutures short or absent, widely separated and convergent, or well developed ( Ulocyphaleus ). Pronotum 0.4–0.8 times as long as wide; widest anteriorly, at middle, or posteriorly with sides more or less straight to strongly curved, sometimes variously lobed (e.g., Pteroderes ) and often explanate; disc variable, sometimes with tubercles, may be vaulted, Phaennis and Notocerastes with paired basal impressions or pits; base narrower than or equal to elytral bases. Lateral pronotal carinae complete, edges sharp or rounded, simple to coarsely dentate, with or without raised margin or bead; anterior and posterior angles acute or rounded; posterior edge straight, evenly rounded or sinuate. Antero- or ventrolateral portions of prothorax often without cavities or grooves, present in Exohadrus . Prosternum in front of coxae shorter than or as long as prosternal process, flat to strongly convex. Prosternal process strongly and abruptly expanded at apex and meeting the hypomeron; flat, concave, or only slightly elevated or curved behind coxae. Notosternal sutures present, incomplete or absent. Procoxae not projecting below prosternum, lateral extensions short, rarely long and concealed ( Waitomophylax ). Procoxal cavities externally closed with postcoxal projection meeting prosternal process ( Fig. 10B ); internally open; more or less circular or slightly transverse, narrowly to widely separated; protrochanteral notch absent or weakly indicated. Scutellar shield fully developed and abruptly elevated ( Fig. 11D ), reduced ( Brouniphylax ) or absent ( Exohadrus ). Elytra 0.85–2.1 times as long as combined width and 1.7–3.2 times as long as pronotum; sides explanate or convex and rarely declivous posteriorly; punctures penetrating deeply into the elytra forming a “window” or not, serially punctate or irregularly punctate; striole present or absent; surfaces smooth, carinate, or tuberculate; apices meeting or almost meeting at suture which is not deflected at apex; epipleuron usually complete, but vaguely defined ( Brouniphylax ) and variously formed in Syrphetodes ; lateral edge simple or microtuberculate. Mesoventrite usually separated by complete sutures from mesanepisterna (not in Brouniphylax ); anterior edge on same plane as metaventrite and without paired procoxal rests; anterior edge width variable, with or without median prosternal rest or with broadly transverse prothoracic rest. Mesanepisterna almost meeting at midline ( Meryx ) or broadly separated (measured by its width to mesoventral process). Mesocoxal cavities circular to slightly transverse ( Fig. 10C ); narrowly to widely separated at middle; open laterally, partly closed by mesepimeron; mesotrochantin always exposed; mesocoxa not conical and projecting. Mesometaventral junction simple or monocondylic. Metaventrite well-developed (longer than wide or quadrate) or reduced and transverse in apterous taxa; with discrimen present (long or relatively shortened) or absent, in the form of a groove in some taxa; postcoxal lines absent; exposed portion of metanepisternum elongate, triangulate in Brouniphylax . Metacoxae usually contiguous or narrowly separated, moderately ( Arthopus , Ganyme , Syrphetodes ) to widely separated ( Brouniphylax , Exohadrus , and Pteroderes ) and extending laterally to meet elytra. Metendosternite with arms moderately to very long ( Figs 7H–I ), laminae present ( Meryx, Trachyderas , and Ulocyphaleus ) or absent; ventrolateral processes absent or weakly developed; anterior tendons present, poorly developed and moderately to widely separated, or absent. Hind wings present ( Fig. 5J ) or absent; with moderately to very long apical field; radial cell well developed, with inner posterobasal angle right; cross-vein r3 absent; medial field with four free veins, medial fleck present; wedge cell long and narrow; medial and anal embayments absent. Femoral attachment of mid trochanter usually strongly oblique with base of femur abutting coxa. Tibiae simple; two apical spurs distinct. Tarsi 5-5-4 (4-4- 4 in Meryx ); tarsomeres ( Figs 6H, I ) simple and similar in form (protarsomere 2 variable in length), tarsomeres 1–4 usually small and with their combined length equal or subequal to tarsomere 5, combined length of tarsomeres 1–4 significantly greater in length than tarsomere 5 in Trachyderastes , metatarsus 1 of Trachyderas elongate and equal to 4; tarsal pads usually absent or poorly developed; pretarsal claws simple; empodium bisetose, setal insertions often ventral. Abdomen ( Figs 7K , 10D ) with five ventrites of relatively equal lengths, with ventrite 1 equal to or slightly shorter than ventrite 2, surface flattened to weakly convex in lateral view, postcoxal lines absent and intercoxal process usually acute or subacute, broadly truncate in Brouniphylax ; ventrites 1 and 2 connate; membranes 3 and 4 externally visible in Arthropus , Exohadrus , Dipsaconia , Meryx , Notocerastes , Phaennis , and Trachyderastes ; glandular pores present in Syrphetodes and Trachyderastes ; lateral flanges present on all or some of the ventrites, well-developed or reduced (some Ganyme , some Syrphetodes , and Trachyderastes ); ventrite 5 with or without apical grooves or impressions. Male with spiculum gastrale on sternite IX present; tergite IX completely fused to tergite X. Aedeagus tenebrionoid ( Figs 8F–G, J–L ), symmetrical, not inverted; parameres partially fused together and articulated to phallobase, forming apicale, which lacks accessory lobes; anterior edge of penis without struts, apex acute to subacute (rounded in one dissected specimen of Arthopus , Figs 8H, I ). Sternite VIII in female with long spiculum ventrale. Ovipositor ( Fig. 7N ) long and slender, bursa typically saccular with a narrow spermathecal gland ( Fig. 8E ); paraprocts distinctly longer than coxites, lightly sclerotized except for baculi, with coxites subdivided, apically slender, with well developed apical or subapical styli. Comments. The family description above is modified from that in Lawrence et al. (1999b) and Leschen & Ślipiński (2010) based on the addition of genera described in this study and recent work by Leschen & Buckley (2015) . Larval characters and descriptions can be found in Beutel & Friedrich (2005) and Leschen & Ślipiński (2010) . There are currently 16 genera and 41 species and further taxonomic work is needed to describe species in Australia ( Dipsaconia , 2 n. spp.; Ganyme , 2 n. spp.; Ulodes , 1 n. sp.), New Zealand ( Brouniphylax , 3 n. spp.) and New Caledonia ( Trachyderastes , 4 n. spp.).