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Tijdschrift voor Entomologie 160 (2017) 1–23

Revision of Dobodura Darlington (Coleoptera: Carabidae: Odacanthini): Diversification on accreted terranes of northern New Guinea James K. Liebherr

The Papuan endemic genus Dobodura Darlington is taxonomically revised, with five newly described species — Dobodura alildablldooya sp. n., D. hexaspina sp. n., D. obtusa sp. n., D. svensoni sp. n., and D. toxopei sp. n. — complementing the type species, D. armata Darlington. The sympatric Dobodura alildablldooya and D. svensoni are described from Chimbu Province, Papua New Guinea. Known distributions of the other three new species are: D. hexaspina, Madang Province, P.N.G.; D. obtusa, Olsobip, Fly River, Western Province, P.N.G.; and D. toxopei, Bernhard Camp, Papua, Indonesia. Dobodura is the sole precinctive Papuan genus in an Australian-Papuan clade also including Clarencia Sloane, Dicraspeda Chaudoir, and Eudalia Laporte. Phylogenetic analysis of Dobodura places its known earliest divergence event on the northern New Guinea margin of the Australian craton. Later divergence events result in species occupying island-arc terranes progressively incorporated into present-day northern New Guinea, commencing in the Miocene. Keywords: aquatic insects; area of endemism; biogeography; cladistic analysis; speciation J. K. Liebherr*, Cornell University Insect Collection, John H. and Anna B. Comstock Hall, 129 Garden Avenue, Ithaca, NY 14853-2601 U.S.A. [email protected].

Introduction Cladistic analysis of these six Dobodura species root- The genesis of this paper started innocently enough. ed at outgroup taxa established during a global phy- During exploration of specimens determined as logenetic analysis of Odacanthini (Liebherr 2016), Dobodura armata using Darlington’s (1968) key to demonstrates that the Dobodura lineage underwent Odacanthini of New Guinea, two novel male geni- its earliest Papuan divergence events on the northern talic configurations were discovered among two male portion of the Australian cratonic portion of New specimens collected in Chimbu Province by G. J. Guinea. Within Dobodura, subsequent divergence Svenson. A careful reading of Darlington’s treatment events led to species that occupy areas subsequently of D. armata — e.g. “The specimen from Bernhard accreted to the northern margin of New Guinea dur- Camp has the strial punctures of the elytra coarser ing Miocene through Pliocene times (Pigram & Da- than in the types and the tip of the aedeagus slightly vies 1987, Polhemus 2007). Areas comprising these different (Darlington 1968: 216)” — led to suspi- accreted island arcs house substantial numbers of cions that the name D. armata cloaked undescribed precinctive taxa in other taxa; e.g. cicadas (Duffels diversity. Subsequent gathering of specimens allowed 1986, De Boer & Duffels 1996, Duffels & Turner the characterization of five undescribed species to 2002) and aquatic Hemiptera (Polhemus & Polhe- complement D. armata, all of them assignable to mus 1998, 2002). The biogeographical history of Dobodura based on synapomorphies of the genus. Dobodura is compared to hypotheses proposed for

Tijdschrift voor Entomologie 160: 1–23, Figs 1–29. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 21 July 2017. DOI 10.1163/22119434-16001001 Downloaded from Brill.com09/30/2021 12:34:38AM via free access

2 Tijdschrift voor Entomologie, volume 160, 2017 these and other groups, putting in context the diver- termed sculpticells by Allen & Ball (1980), vary sification ofDobodura during geological assembly of from isodiametric, overlapping six-sided sculpticells New Guinea. The aggregate distribution of Dobodu- arrayed as roof tiles, to transverse sculpticells arrayed ra is restricted to areas of orogenic uplift in northern as a mesh, to transverse lines not cross-connected New Guinea, with the associated geological faulting to one another (Lindroth 1974). Ball made signifi- producing the streams and cascades within which cant use of sculpticell configuration for interpreting Dobodura beetles reside. evolutionary history in the carabid tribe Galeritini (Ball & Nimmo 1983, Ball 1985a, b). This study fol- lows Lindroth and Ball’s terminology for describing Material and methods sculpticell shape. Taxonomic material. This revision is based on 18 Variation among individuals for counts and rela- Dobodura specimens borrowed from six institutional tive positions of the lateral elytral setae — i.e. those se- collections (see Acknowledgements). Type label data tae present in the broadened eight elytral interval — is are transcribed verbatim, with label line breaks indi- presented using the convention of a + (b + c + d–e) + cated by a slash (/) and subsequent labels indicted by (f + g). In this sequence, “a” represents the number of a double slash (//). setae in the anterior series of lateral elytral setae com- Laboratory techniques. Standard light microscopy, mencing just laterad the humerus, and “b, c, and d–e” dissection and staining protocols used in this study represent numbers of setae present along the lateral are described in Liebherr (2015: 18–20). In order to margin of the elytron. Variation in the number of setae examine the abdominal structures of segments VIII present is designated by numbers connected by a dash: and IX in association with the male aedeagus, both e.g., d–e. Each number in the middle parenthetical set abdominal segments VIII and IX and the associ- represents an isolated seta or group of setae. The trail- ated aedeagus were removed together from males. ing parenthetical setae (f + g) represent the two groups This procedure allowed examination of mediotergite of setae just anterad the lateroapical spine or denticle VIII and the conformation of the antecostal margin in these beetles. of segment IX (Deuve 1993), herein called the ring Standardized body length used to describe body sclerite. Male genitalia were slide mounted in glycer- size is the sum of three measurements: 1, head length ine for microscopic examination and photography. measured from the labral medioapical margin to the Descriptive conventions. Several measurements cervical ridge at the head–pronotal juncture; 2, pro- proved useful for diagnosing species. Eye develop- notal length measured along the dorsal midline; 3, ment was quantified by the ocular ratio: maximum elytral length defined as the distance from the base head width across eyes divided by the minimum of the scutellum (generally entirely exposed in these frons width between eyes. The relative diameter of beetles) to the apex of the sutural spine. As this sum the eyes in proportion to their convexity is expressed of measurements ignores the apical portion of the as the ocular lobe ratio; i.e. the depth of the eye mea- elongate mandibles (always in variable positions sured with the eye fossa vertical in the field of view, specimen to specimen) and the length of the apical over the longitudinal diameter of the eye measured spines, the standardized body length measure will be in the same orientation. Higher values are associated smaller than the body size perceived by eye. with more “popeyed” eyes. The degree of constric- Phylogenetic analysis — Taxa. In a global cladistic tion of the neck was assessed using the ratio MHW/ analysis of Odacanthini comprising 79 taxa (Lieb- NW, or maximum head width across eyes divided herr 2016), Dobodura armata was placed as adelpho- by neck width, measured in dorsal view. The relative taxon to Clarencia quadridens Darlington, with the elongation of the prothorax was quantified by the exemplar taxa Dicraspeda quadrispinosa (Chaudoir) ratio PL/MPeW, or pronotal length measured me- and Eudalia atrata Baehr as successively more in- dially, divided by the width of the prothorax across clusive adelphotaxa. In order to develop the current the bulbously expanded proepisterna, in dorsal view. phylogenetic hypothesis for relationships among The ranges of all ratios are used only for descriptive Dobodura spp., the five newly described species were purposes. added to the data matrix; those taxa scored for rel- The body surface of carabid beetles is covered evant characters. The six Dobodura ingroup taxa plus with a network of raised cuticular ridges; these ridges the three generic outgroups were rooted at a fourth are collectively described as cuticular microsculpture. outgroup; Porocara nigricollis Baehr. In the global This microsculpture “probably correspond[s] to the analysis (Liebherr 2016), Porocara nigricollis was shapes and dimension of the epidermal cells that se- the earliest divergent representative Australian taxon creted the cuticle (Hinton & Gibbs 1969).” A stan- within the subtribe Odacanthina, to which the other dard terminology of the various shapes of the cells taxa in this analysis also belong. Through this array defined by these ridges was first proposed for cara- of outgroups, relationships within Dobodura can be bid beetles by Lindroth (1974). The individual cells, hypothesized while retaining the same intergeneric Downloaded from Brill.com09/30/2021 12:34:38AM via free access

Liebherr: Revision of Dobodura (Coleoptera) 3 phylogenetic relationships presented in the global 12. Elytral apical spine at apex of stria 3: absent analysis of Odacanthini. (0); present (1; Figs 9–14). Phylogenetic analysis — Characters. Based upon 13. Apical spines: in same orientation as sutural characters represented in the global cladistic analy- spines (0); pointing upward relative to sutural sis (Liebherr 2016), a character × taxon matrix was spines (1; Figs 10–12). developed for the six Dobodura spp. and the four 14. Subapical sinuation: nearly obsolete, elytral outgroups, allowing characters varying among the apices obliquely truncate (0); evident (1); ten taxa to be evaluated. These characters were then distinct, angularly concave (2; Fig. 9); very scored cladistically in WinClada (Nixon 2002), with distinct, with lateroapical spine laterally (3; characters numbered 0–24 as per the WinClada de- Figs 10–14). fault. Multistate characters were treated as ordered 15. Frons microsculpture: evident, isodiametric to transformation series. The cladistic matrix and as- transverse (0); reduced, not traceable except in sociated phylogenetic analysis may be viewed using spots (1); absent (2). Winclada (Supplementary material S1). The charac- 16. Elytral disc microsculpture: isodiametric ters used are: mesh, may be in transverse rows (0; Fig. 1); transverse mesh (1); evident transverse lines 0. Mandibular length: short, ≤ to ~1.5 × distance (2); not evident, surface glossy (3; Fig. 2). antenna base–labral margin (0); moderately 17. Femora: concolorous from base to apex, or elongate, 1.5–1.75 × distance (1); elongate, apex only slightly darker (0; Fig. 9); with apex >1.8–2.25 × distance (2); very elongate, >2.3 × dark, basal 1/2–4/5 pale (1). distance (3; e.g. Fig. 12). 18. Male tergite IX ring sclerite: angulate apically, 1. Eyes: moderate (0); large, greatly projected wishbone-shaped (0); hemicircular apically, (1); hypertrophied, convexly projected (2). horseshoe shaped (1; Fig. 4). 2. Eye diameter: moderate (0); very small (1; 19. Male aedeagal median lobe shaft: straight Figs 9–14). in apical half, tip may be curved dorsally 3. Proepisternum: smooth, impunctate (0); (0; Figs 17–24); sinuously recurved in apical punctured in posterior half (1); distinctly half in addition to curved tip (1; Figs 15, 16, punctured or wrinkled over entire surface (2). 25, 26). 4. Elytral humerus: rounded laterad parascutellar 20. Aedeagal median lobe apex: gradually nar- striole (0); tightly rounded to angulate laterad rowed apicad ostium (0; Figs 17–20); parallel- base of fifth interval (1); distinctly angulate sided, elongate apicad ostium (1; Figs 15, 16, laterad base of fifth interval (2; Fig. 1). 21–26). 5. Elytral lateral margin: convex laterad anterior 21. Male aedeagal median lobe apex: of moderate series of lateral elytral setae (0); invaginated length, attenuate or spoon-shaped (0; Figs 17– laterad anterior series of lateral elytral setae (1; 24); exceedingly elongate, porrect (1; Figs 15, Figs 9–14). 16, 25, 26). 6. Parascutellar seta: single each elytron (0); dou- 22. Female apical abdominal ventrite: convex or ble each elytron (1; Fig. 1). slightly emarginate medially (0); with rounded- 7. Dorsal elytral setae: 3–4 or less (0); more than keyhole marginal invagination (1; Figs 5, 7); 3–4, 4 or 5–6 present along length of interval with cordately invaginated margin (2; Fig. 6). 3 (1). 23. Female bursa copulatrix: membranous, inner 8. Dorsal elytral setae: articulatory socket (ante- surface smooth (0); sparsely covered with spic- rior seta) coplanar with elytral disc (0); set in ules on inner surface (1). foveate articulatory socket (1; Fig. 11). 24. Apical gonocoxite 2: with two lateral ensiform 9. Elytral macrosetae: present only on interval setae (0); with three lateral ensiform setae (1; 3 (0); also present on interval 5 (1); also pres- Fig. 28); with 4–5 lateral ensiform setae (2). ent on intervals 5 and 7 (2); also present on intervals 1, 5 and 7 (3); present on all intervals Phylogenetic analysis — Analytical methods. The 1–7 (4). WinClada matrix was submitted to the cladistic par- 10. Elytral striae: impunctate to minutely punc- simony computational program NONA (Goloboff tate (0; Figs 9–14); regularly punctate (1); dis- 1999) and analyzed using the ratchet (Nixon 1999). tinctly, broadly and deeply punctate (2). An initial run using 200 iterations of the ratchet 11. Elytral apices: meeting at suture, margins even- with Winclada default settings found one tree of ly curved (0); separately rounded with space 54-step length (CI = 68, RI = 73). A subsequent between elytra (1); separately pointed, elytra run using 10,000 ratchet iterations, same settings, bispinose apically (2); separately spinose, each found the identical, single most parsimonious tree elytron with apical spine (3; Figs 9–14). (Fig. 8). Downloaded from Brill.com09/30/2021 12:34:38AM via free access

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Figs 1, 2. Elytral features of Dobodura spp. — 1, elytral base, D. obtusa; parascutellar setae doubled (anterior parascutellar seta, aps; posterior parascutellar seta, pps), elytral striae 1–7 numbered; 2, elytral apex, D. alildablldooya; posterior series of lateral elytral setae numbered 1–4, subapical seta (sas) and apical seta (as) both present, elytral spines include lateroapical spine (lasp), apical spine (asp), and sutural spine (ssp). Asterisks placed just anterad setal articulatory sockets.

Phylogenetic analysis results the spine. The sister-group relationship of the ter- The single most parsimonious cladogram (Fig. 8) minal species pair of this lineage—D. armata and illustrates robust support for the monophyly of Do- D. alildablldooya—is again supported by aedeagal bodura, with all seven characters that support this configuration, with these two species characterized node unreversed within the generic ingroup. These by the most evenly narrowed, least upturned tip to seven synapomorphies include: 1, mandibles very the median lobe (Figs 17–20). Thus aedeagal evolu- elongate (character 0; e.g. Fig. 12); 2, proepisternum tion among these six species is hypothesized to fit a smooth (character 1); 3, parascutellar setae doubled transformation series whereby the median lobe apex (character 6; Fig. 1); 4, anterior dorsal elytral seta is shortened and straightened relative to the plesio- within broad shallow depression (character 8; e.g. morphic, porrect and curved condition (Figs 15, 16, Fig. 11); 5, elytra with apical spine (character 12; 25, 26). Fig. 2); 6, antecostal margin of abdominal segment IX horseshoe-shaped (character 18; Fig. 4); and 7, Taxonomy female apical abdominal ventrite with medioapical Genus Dobodura Darlington “keyhole” invagination (character 22; Figs 5–7). Dobodura Darlington, 1968: 215. Within Dobodura, D. obtusa and D. svensoni comprise the adelphotaxon to the other four species Diagnosis. Members of this genus can be imme- based on aedeagal configuration. The former two diately diagnosed from other Odacanthini by the species are characterized by sinously recurved aedea- doubled parascutellar setae (Fig. 1) and presence of gal median lobes (Figs 15, 16, 25, 26), versus me- a long apical elytral spine apicad the third stria to dian lobes with a straight shaft and less elongate, less accompany shorter sutural and lateroapical elytral porrect apex (Figs 17–24). Among the four-species spines or denticles (Fig. 2). The beetles have glossy clade subtended by D. toxopei (Fig. 8), the species piceous body coloration, contrasted to the testaceous triplet D. hexaspina, D. armata, and D. alildabll- palps, antennae and legs (Figs 9–14). The proepi- dooya is characterized by more elongate apical elytral sternum and prosternum are smooth and glossy, spines (Figs 10–12), these spines oriented upward not punctate as in members of the related genera relative to the sutural spines that extend directly pos- Eudalia, Dicraspeda, and Clarencia. The elytral basal terad from the sutural apex. Apical spines in these groove is tightly curved anteriorly near the base species also deviate from those of the other three of the fifth stria resulting in an angulate to subangu- species (Figs 9, 13, 14) in pointing outward, away late hitch in the groove mesad the humerus (Fig. 1), from the suture, though this attribute was not coded and the anterior dorsal elytral seta is situated in independently from the more upward orientation of a broad shallow indentation of the third elytral Downloaded from Brill.com09/30/2021 12:34:38AM via free access

Liebherr: Revision of Dobodura (Coleoptera) 5

Figs 3, 4. Male aedeagus and associated abdominal segments, D. hexaspina. — 3, male aedeagal median lobe (aed) and left paramere (lp) lying in situ within ring sclerite, dorsal view; 4, male abdominal segments VIII and IX, dorsal view. Abbreviations include: ac VIII, antecostal margin of abdominal VIII; ac IX, antecostal margin of abdominal IX, i.e. the anterior portion of ring sclerite; dgo, defensive gland orifice in membrane between abdominals VIII and IX; dgr, defensive gland reservoir; hg, hindgut, ltg VIII, laterotergite of abdominal VIII; mtg IX, mediotergite of abdominal IX, i.e., the posterior portion of ring sclerite; spir VIII, spiracle of abdominal VIII associated with laterotergite VIII. Asterisk in Fig. 4 indicates that tergite of segment VIII is divided medially by membrane.

interval. Dobodura species are the only Odacanthi- Description. Head capsule elongate with distinctly ni observed wherein the male 9th tergal apodeme constricted neck; eye with small diameter, outer sur- (Deuve 1993) — comprising the apex of the ring face convex to very convex or pop-eyed; mandibles sclerite — is broadly rounded, or “horseshoe”-shaped elongate, distance from anterior condyle to apex (Fig. 4), rather than broadly angulate. And though greater than 2.3× distance from condyle to latero- females are not known for all of the six species treat- apical labral margin; antennae elongate, third anten- ed here, available females of three species exhibit a nomere 1.1× length of fourth antennomere, basal medial “keyhole” along the apical margin of the three antennomeres glabrous except for apical setae, apical ventrite (Figs 5–7). Presence of this notch fourth antennomere glabrous in basal third of length; on the apical ventrite in the three species assessable mentum with acute median tooth; submentum with for this character — D. obtusa, D. hexaspina, and two setae present each side. Pronotum elongate, lon- D. armata — optimizes the character to be present in ger than broad, completely margined laterally, the the common ancestor of Dobodura (Fig. 8, character marginal carina extended to base; median base not 22), and so it is predicted that all Dobodura spp. fe- margined, a deep transverse sulcus defining a sinu- males exhibit this derivation. ous transverse collar basally; median longitudinal Downloaded from Brill.com09/30/2021 12:34:38AM via free access

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Figs 5-7. Apical abdominal ventrite of female Dobodura spp., ventral view, showing median “keyhole” invagination. — 5, D. obtusa; 6, D. hexaspina; 7, D. armata. impression very fine; anterior transverse impres- proepimeron transversely wrinkled to punctulate. sion absent medially, shallow and somewhat evident Elytra slightly broader apically, lateral margins subpar- toward front angles; front angles rounded to acute- allel, slightly constricted near posterior seta of anterior angulate, little projected; lateral pronotal seta lateral setal series; intervals flat, striae represented by present in lateral marginal depression before mid longitudinal series of isolated punctures in basal two- length; vertical furrow between proepisternum and thirds of length (Fig. 1),Downloaded striae difficult from Brill.com09/30/2021 to trace apically 12:34:38AM via free access

Liebherr: Revision of Dobodura (Coleoptera) 7

Fig. 8. Most-parsimonious 54-step cladogram for Dobodura spp. also including representative generic exemplars Eudalia nigra, Dicraspeda quadrispinosa, and Clarencia quadridens, rooted at the relatively more plesiotypic Austral- ian taxon Porocara nigricollis. Underlying data presented in Supplementary material S1. due to reduced punctation; subapical and apical setae sutural spine, the apical spine 0.10–0.12× ely- present. Metepisternum elongate, metathoracic flight tral length so measured; elytral margin anterad wings fully developed. Legs elongate, gracile; metatar- subapical excavation obtusely convex, without somere 4 (mt4) outer lobe 0.33–0.55× median tarso- evident tooth (Fig. 9) �� mere length; mt4 with both apical and subapical setae �� Dobodura obtusa sp. n. present (Habu 1978); mt5 with ventrolateral rows of 2. Prothorax more quadrate, median pronotal ~6 elongate setae. length to breadth across laterally extended Due to the paucity of female specimens, only a proepisterna (PL/MPeW) = 0.94–1.01 �� 3 female of D. armata was dissected, and so the type – Prothorax more elongate, PL/MPeW = 1.07– species must serve as proxy for all Dobodura with re- 1.11 �� Dobodura armata Darlington gard to female tract characters. These characters are 3. Elytral margin anterad subapical excavation presented below in the D. armata species treatment. with well-projected, acute tooth; elytral su- Type species. Dobodura armata Darlington (1968: tural and apical spines elongate, sutural spine 215) by original designation. 0.06× elytral length (base of scutellum to apex Distribution. All species assignable to this genus are of subapical spine), apical spine 0.15–0.17× precinctive to New Guinea. elytral length so measured �� 4 – Elytral margin anterad subapical sinuation with small projected tooth; elytral sutural Key to the adults of Dobodura Darlington and apical spines shorter, sutural spine 1. Discal elytral intervals glossy, any sculpticells 0.04× elytral length, apical spine 0.12× elytral that may be present over portions of surface length �� 5 transverse in orientation; sutural and apical 4. Elytral basal groove distinctly angulate on elytral spines longer, sutural spine 0.03–0.06× humerus, the basal carina bent at nearly right length of elytra from base of scutellum to apex angle basad fifth stria; prothorax elongate, me- of sutural spine, apical spine 0.12–0.17× ely- dian pronotal length to breadth across later- tral length so measured; elytral margin anterad ally extended proepisterna (PL/MPeW) = 1.01 subapical excavation bearing an acute tooth- �� Dobodura alildablldooya sp. n. like projection (Fig. 2) �� 2 – Elytral basal groove obtusely projected ante- – Discal elytral intervals covered with distinct riorly on humerus, the basal carina obtusely (female) or shallowly margined (male) iso- convex basad fifth stria; prothorax more quad- diametric sculpticells; both sutural and apical rate, median pronotal length to breadth across spines short, the sutural spine 0.02× length laterally extended proepisterna (PL/MPeW) = of elytra from base of scutellum to apex of 0.96 ��DownloadedDobodura from Brill.com09/30/2021 hexaspina sp.12:34:38AM n. via free access

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Figs 9-12. Dorsal habitus views of Dobodura spp. — 9, D. obtusa; 10, D. armata; 11, D. alildablldooya; 12, D. hexaspina.

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Liebherr: Revision of Dobodura (Coleoptera) 9

Figs 13-14. Dorsal habitus views of Dobodura spp. — 13, D. toxopei; 14, D. svensoni.

5. Ventral surfaces of femora piceous, much glossy reflection on the elytral disc. The elytra apex darker than dorsal surfaces, and antenno- near the bases of the apical spines is covered with meres 1–3 pale, testaceous, antennomere 4 a well-developed isodiametric mesh, the sculpticells with brunneous cast in apical ¾ of length to raised in their centers. The apical elytral projections match brunneous antennomeres 5–11 are also the least developed among all species, with ��Dobodura toxopei sp. n. the lateroapical projection very obtuse, rounded, – Ventral and dorsal surfaces of femora con- and apical spine length 0.10–0.12× elytral length. colorous, testaceous, only femoral apices The elytral basal groove exhibits an obtusely an- with brunneous cast to match tibial bases, gled hitch at the base of the fifth stria. The neck and antennomeres 1–4 testaceous, con- is quite narrow relative to the moderately convex trasted to brunneous antennomeres 5–11 eyes; MHW/NW = 1.77–1.83. As for D. armata �� Dobodura svensoni sp. n. and D. alildablldooya, there are three dorsal elytral setae. Finally, as in D. svensoni (Fig. 14) the femora are concolorous on their dorsal and ventral Dobodura obtusa Liebherr, sp. n. surfaces (Fig. 9), not darker ventrally as in the other Figs 1, 5, 9, 15, 16, 29 four species (Figs 10–13). The apical abdominal ventrite of the female has an ovoid-keyhole Type material invagination medially, the apical opening to the Holotype. : Papua New Guinea: Fly R. / Olsobip, ♂ circular keyhole comprising about 90° of the cir- 400 m- / 600 m. VIII.69 // J. & M. Sedlacek / Col- cumference (Fig. 5). Standardized body length lectors / BISHOP // Dobodura / armata Darlington 9.4–10.0 mm. / det. D. Kavanaugh 1989 // HOLOTYPE / Dobo- dura / obtusa / J.K. Liebherr 2016 [black-bordered red label] (BPBM). Description Eyes prominent, convex and popeyed, ocular ratio Allotype. ♀: same label data except // ALLOTYPE / Dobodura / obtusa / J.K. Liebherr 2016 [black- 1.63–1.64, ocular lobe ratio 0.41–0.44. Prothorax bordered red label] (BPBM). quadrate to slightly broader than long in dorsal view due to expanded proepisterna, PL/MPeW = 0.94– Diagnosis 0.99; pronotal lateral marginal carina low, the lateral This species is distinct among Dobodura due to the marginal furrow broad at lateral seta; transverse fur- evident isodiametric sculpticells visible through the row on median base irregularly rugose medially, with Downloaded from Brill.com09/30/2021 12:34:38AM via free access

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Figs 15-26. Male aedeagal median lobe, Dobodura spp; odd-numbered figures left-lateral view, even-numbered figures, right-lateral view. — 15, 16, D. obtusa; 17, 18, D. armata; 19, 20, D. alildablldooya; 21, 22, D. hexaspina; 23, 24, D. toxopei; 25, 26, D. svensoni.

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Liebherr: Revision of Dobodura (Coleoptera) 11

Fig. 27. Female abdominal segments VIII and IX, and female reproductive tract D. armata, ventral view. Abbrevia- tions include: bc+spph, bursa copulatrix with lumen filled by spermatophore; co, common oviduct; ga, gonocoxal apodeme; gc, gonocoxa; hg, hindgut filled with head capsules of larval Ceratopogonidae (Supplementary material S2); lo, lateral oviduct; sab, spermathecal apical bulb that is joined to spermathecal basal sclerite by narrow duct; sbb, spermathecal basal bulb with spermathecal gland duct joined to apex; sbs, spermathecal basal sclerite that supports spermathecal assembly and joins it to common oviduct; sg, spermathecal gland. deep and narrow depression each side at terminus opening at the adeagal apex through which the inter- inside lateral marginal bead; anterior transverse im- nal sac everts is oriented toward the dorsal surface of pression obsolete medially, well defined just inside the median lobe, and thus is not evident in either left front angles resulting in a narrow anterior marginal (Fig. 15) nor right (Fig. 16) views. bead at angles; front angles slightly protruded, obtuse. Elytra with short sutural spine, length 0.2× Distribution elytral length; 15 lateral elytral setae arranged as 6 + This species is known only from Olsobip in an up- (1 + 1 + 3) + (2 + 2), the orientation of the posterior per drainage of the Fly River (Fig. 29). This local- two setae of the posterior series nearly in line with ity was termed the Trans-Fly Foreland by Polhemus the anterior setal pair of that series. Mesepisternum et al. (2004), and is defined by endemic species in with nine punctures in two vertical rows; metasternal the Gerridae and Veliidae (Hemiptera), and by vari- process acute, tip unmargined though sides of pro- ous fish species (Polhemus & Allen 2007). cess are margined posteriorly. Abdominal ventrite 1 with triangular depression at mid-breadth each side. Etymology Metatarsomere 4 outer lobe elongate, length 0.57× The adjectival species epithet obtusa refers to the very median tarsomere length. obtuse lateroapical elytral projections of these beetles Male genitalia. Male aedeagal median lobe sinu- (Fig. 9), the “spine” in this species is represented only ously recurved, the apex parallel-sided and elongate by a small obtuse denticle at the lateral reach of the with a tightly rounded tip (Figs 15, 16). The ostial subapical sinuation. Downloaded from Brill.com09/30/2021 12:34:38AM via free access

12 Tijdschrift voor Entomologie, volume 160, 2017

/ Darl. [red label] // M.C.Z. / Holotype / 31512 [red label] (MCZ, Type No. 31,512). Paratypes. Same label data (BMNH, 1; MCZ, 4). Darlington’s (1968) two paratypes from Mt. Han- semann, Astrolabe Bay represent D. hexaspina sp. n. (see below). Diagnosis This large-bodied species, standardized body length 10.4–10.7 mm, is characterized by an elongate prothorax, PL/MPeW = 1.07–1.11 (versus ratios of 0.93–1.03 observed among individuals of all other species). The apical elytral spines are long, apical spine length 0.15–0.17× elytral length, with the spines pointed upward and outward relative to the orientation of the shorter sutural spines (Fig. 10). The eyes are among the least convex of all Dobodura, with the ocular ratio = 1.57–1.61, though the neck is quite constricted; MHW/NW = 1.78–1.81. There are three dorsal elytral setae in the third interval, and there are 15 lateral elytral setae arranged as 6 + (1 + 4) + (2 + 2), with the posterior pair of setae in the posterior series oriented to define a line that passes through the apex of the lateroapical spine. As with the four species treated below, the microsculpture is reduced, the elytral disc glossy without evident sculpticells, and the apex near the base of the apical spine with a shallow, though evident, transverse mesh. The apical margin of the female abdomen bears a round-keyhole invagination, the opening of the keyhole comprising about 50° of the circumfer- ence (Fig. 7). Male genitalia. The shaft of the aedeagal median lobe is straight in this species, and the apex is gradually narrowed alongside the ostial opening, that narrow- ing continued to the pointed tip (Figs 17, 18). The ostial opening is oriented toward the right side of the median lobe (Fig. 18). Female reproductive tract. The bursa copulatrix is elongate (holding a large spermatophore in the dissected female) (Fig. 27); common oviduct extended beyond bursa for a distance equal to the bursal Fig. 28. Female left gonocoxa D. armata, ventral view. length; spermathecal assembly joined to common Abbreviations include: afs, apical fringe setae of basal oviduct by spermathecal basal sclerite (Liebherr & gonocoxite 1; ans, apical nematiform setae of apical Will 1998); spermatheca consisting of basal bulb gonocoxite 2; ga, gonocoxal apodeme; gc1, basal gono- that extends from apex of spermathecal basal sclerite, coxite 1; gc2, apical gonocoxite 2; les, lateral ensiform and a spermathecal apical bulb that is connected to setae of gonocoxite 2. the basal sclerite by a very narrow duct; gonocoxa bi- partite, the basal gonocoxite with heavily sclerotized laterobasal apodeme (Figs 27, 28); basal gonocox- Dobodura armata Darlington ite with apical fringe of seven setae (Fig. 28), apical Figs 7, 10, 17, 18, 27–29 gonocoxite broad basally, broadly rounded apically, and with three lateral ensiform setae; a dorsal en- Type material siform seta not present; two apical nematiform se- Holotype. ♂: Dobodura / Papua N.G. / Mar–July, tae in fossate pit on medioventral surface of apical 1944 / Darlington // Holotype / Dobodura / armata gonocoxite.

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Liebherr: Revision of Dobodura (Coleoptera) 13

Fig. 29. Known geographical distributions of Dobodura spp. Dotted line indicates geosyncline associated with accretion of Solomon Arc Terranes onto northern margin of New Guinea during the Miocene (Polhemus & Polhemus 1998, 2002). Cladistic relationships of Dobodura spp. (Fig. 8) indicated in upper right.

Distribution Blackburnia mandibularis Liebherr and B. riparia Li- Dobodura armata is restricted to the vicinity of Do- ebherr & Short (2007), and the trechine Scotoplanetes bodura Airfield complex (Darlington 1968), now aquacultor Lakota et al. (2010). named Girua Airport (Fig. 29). This locality is in the Popondetta Foreland area of endemism (area 32 of Polhemus & Allen 2007) that is defined by five spe- Dobodura alildablldooya Liebherr, sp. n. cies of Gerridae, Naucoridae, and Veliidae (Hemip- Figs 2, 11, 19, 20, 29 tera) plus ten species of fish. Type material Habitat Holotype. ♂: Papua New Guinea / Chimbu Prov- Darlington (1968: 216) states “My specimens were ince, Wara / Sera Research Station / Crater Moun- taken among spray-drenched stones beside small tor- tain / Research Area // VII–8 ~ 13–2001 / Bradler, rents in rain forest.” The dissected female was found to Jarvis, / and Svenson // HOLOTYPE / Dobodura / have hundreds of head capsules of Ceratopogonidae alildablldooya / J.K. Liebherr 2016 [black-bordered (Diptera; Foote 1991) packing the hindgut (Fig. 27, red label] (CUIC). Supplementary material S2). That this female was able Paratype. ♂: NEW GUINEA: / E. Highland Dist. / to sufficiently engorge on fly larvae filling her hindgut Wanatabe Valley / Nr. Okapa, c. 5000 ft. / 5.II.1965 to repletion suggests that the fly larvae were both dis- // Stn. No. / 174 // M. E. Bacchus / B.M. 1965–120 tributed at high density within the environment, and // Dobodura / armata / det. Darl. // PARATYPE also distributed in a situation that allowed rapid forag- / Dobodura / alildablldooya / J.K. Liebherr 2016 ing. Given that Darlington found these beetles along [black-bordered yellow label] (BMNH). a torrential stream, it would appear that the dissected female entered the splash zone—i.e., a semiaquatic Diagnosis or aquatic microhabitat—where it encountered and This species (Fig. 11) shares the elongate, upwardly ingested the ceratopogonid larvae. Thus D. armata oriented elytral apical spine, and bicolored femora joins the increasing number of carabid species across with D. armata (Fig. 10) and D. hexaspina (Fig. 12). numerous tribes that have been recorded foraging However it is diagnosable from D. armata by the in hygropetric or fully aquatic situations. These in- broader neck (MHW/NW = 1.71–1.73 versus 1.78– clude the broscine Rawlinsius papillatus Davidson & 1.81 for D. armata) and longer sutural spine (sutural Ball (1998), the odacanthine Quammenis spectabilis spine length 0.04–0.06× versus a value of 0.03 for (Erwin 2000, Liebherr 2016), two platynine species, D. armata). It is diagnosable from D. hexaspina by

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14 Tijdschrift voor Entomologie, volume 160, 2017 the presence of three dorsal elytral setae in the third At Dobodura, Darlington collected over 4,000 cara- interval instead of the four to six dorsal elytral setae bid beetle specimens representing 217 species while observed among individuals of that species. D. alild- having one arm incapacitated (Darlington 1952: ablldooya is also characterized by the presence of a 325–326; 1971: 142). By choosing the airfield’s pit-like depression at the lateral termini of the basal locality name as the generic name for this particu- transverse sulcus of the pronotum; such terminal pits lar genus, he commemorated the locality where he otherwise observed only in individuals of D. obtusa, collected the greatest diversity of New Guinean with that species amply diagnosed by elytral spina- Carabidae (Darlington 1971). The species epithet tion and microsculpture. Standardized body length chosen for this species—alildablldooya—was cho- 9.6–10.0 mm. sen to honor the aviators who flew out of Dobodura through use of a truncated and concatenated version Description of the subsequently written television jingle used to Head foreshortened, eye convexity slightly greater advertise Brylcreem®, a favored pomade of WWII than that of gena behind eye, ocular ratio 1.53–1.58. airmen (Dick 1993). The name should be treated as Prothorax appearing elongate, but dimensions quad- a random assortment of letters even though the latter rate in dorsal view due to expanded proepisterna, portion of the species epithet alliterates the generic PL/MPeW = 1.00–1.01; pronotal lateral marginal name. carina fine, the lateral marginal depression narrow at lateral seta; basal transverse furrow distinctly punctate medially, 3–4 punctures each side of midline, Dobodura hexaspina Liebherr, sp. n. furrow broadly meeting lateral marginal bead; ante- Figs 3, 4, 6, 12, 21, 22, 29 rior transverse impression visible medially, the anterior callosity depressed across front of pronotum; Type material front angles finely protruded, acute. Elytral apical Holotype. ♂: Mt. Hansemann / Astrolabe B. (on ob- spine length 0.17–0.18× elytral length; lateroapical verse) / III.30 (on reverse) // N. Guinea / Biró 1901 spine prominent, acute, elytral subapical sinuation // Paratype / Dobodura / armata / Darl. [red label] distinctly concave mesad spine; elytral basal groove // Paratype [red label] // RMNH / Museum Leiden with angulate hitch at base of fifth stria, but groove // HOLOTYPE / Dobodura / hexaspina / J.K. Lieb- only moderately displaced anteriorly; 15 lateral ely- herr 2016 [black-bordered red label] (RMNH). tral setae arranged as 6 + (1 + 4) + (2 + 2), with Allotype. ♀: Same label data except // ALLOTYPE / the posterior pair of setae in posterior series oriented Dobodura / hexaspina / J.K. Liebherr 2016 [black- so that they define a line passing through apex of bordered red label] (RMNH). Additional paratypes: lateroapical spine. Mesepisternum with 9–10 punc- PAPUA NEW GUINEA / Madang Province: / Sim- tures in 2–3 rows; metasternal process right apically, bai / 5°17´S 144°26´E / 1 June 1988 / W.J. Palawski apex margined. Abdominal ventrite 1 with ovoid de- cllr // Dobodura / armata / Darlington / det. D.H. pression laterally and punctate linear depression ap- Kavanaugh / 1989 (CAS, 1); PAPUA NEW GUIN- proaching coxal articulation. Metatarsomere 4 with EA, / Madang Province, Tapo / Creek at North Coast outer lobe short, length 0.25× median tarsomere / Road, 75 m, 5 March 1989 / Stop #89-12 // D.H. length. Kavanugh, / G.E. Ball, N.D. Penny, / & P.A. Meyer, Male genitalia. The aedeagal median lobe shaft is collectors / CAL. ACAD. SCI. COLL. // PAPUA straight, the apex slightly upturned beyond the ostial NEW GUINEA / EXPEDITION--1989 (CAS, opening, moderately elongate, and parallel-sided be- 1); PAPUA NEW GUINEA, Madang / Province, fore the rounded tip (Figs 19, 20). The ostial open- Adelbert Range, / south slope of Mount / Men- ing is oriented toward the right side of the median gam, headwaters of / Omasa River, 1300 m, / 12 lobe (Fig. 20). Apr. 1989, Stop #89-71 // D.H. Kavanaugh and / N.D. Penny collectors // CAL. ACAD. SCI. COLL. Distribution // PAPUA NEW GUINEA / EXPEDITION--1989 This species is known from Chimbu and the Eastern (CAS, 3). Highlands Provinces of Papua New Guinea (Fig. 29). Diagnosis Etymology This species can be diagnosed by the combination: The Dobodura Airfield east of Oro Bay was used (1) third elytral interval with 4–6 dorsal elytral se- as front-line post of the U.S. Army Air Forces and tae; (2) elytral apical spines elongate, 0.12–0.18× Royal Australian Air Force during the Allied advance elytral length with spines pointed upward relative in 1942–1943 (Null 1995). During his service as an to the sutural spines (Fig. 12); (3) eyes relatively army entomologist, Philip Darlington convalesced at large in diameter and only moderately convex, ocu- Dobodura from March to July 1944 after suffering lar ratio 1.52–1.59; (4) neck broad, MHW/NW = a crocodile attack (Carpenter 1985, Wilson 1991). 1.71–1.73; and (5) elytral basal groove only broadly Downloaded from Brill.com09/30/2021 12:34:38AM via free access

Liebherr: Revision of Dobodura (Coleoptera) 15 subangulate near base of fifth stria, not angulate. The Hydraena beetles (Coleoptera: Hydraenidae). In this apical abdominal ventrite of the female has a cordate genus of water beetles 16 of 36, or nearly half of the keyhole invagation medially (Fig. 6). Standardized species, found in area 16 are endemic to that area body length 8.9–10.5 mm. (Perkins 2011). Description Habitat Pronotum quadrate, length and breadth across ex- Information on the habitat at two of the collect- panded proepisterna subequal, PL/MPeW = 0.95– ing sites for this species has been provided by D. 1.03; pronotal lateral marginal bead low, fine, lateral H. Kavanaugh (pers. comm.). On 5.iii.1989, one marginal depression moderately broad at lateral seta; specimen was collected “under rock at edge of small basal transverse furrow irregularly rugose medially, cascade in forest on north side of river”, that river up to five irregular punctures along length, furrow being Tapo Creek. On 12.iv.1989 a series of three broadly meeting little-elevated marginal bead; an- specimens was found “under rocks on moist sand terior transverse impression evident across prono- well back from water’s edge, ± shaded by overhang- tum, narrow and shallow medially, broader inside ing bank”. In both situations, D. hexaspina was the front angles; front angles slightly protruded, broadly only species collected in that microhabitat. rounded. Elytra with parascutellar striole obsolete basally due to reduction of strial punctures; sutural Etymology The adjectival epithet hexaspina refers to the well- spine elongate, 0.4–0.6× elytral length; lateroapical spine evident, apex right, and spine consisting developed sutural, apical, and lateroapical spines on of expanded elytral lateral marginal bead; 15 lateral the elytra of these beetles (Fig. 12). elytral setae arranged as 6 (1 + 1 + 3) + (2 + 2) with posterior pair of posterior series oriented so that they define a line passing through apex of lateroapi- Dobodura toxopei Liebherr, sp. n. cal spine. Mesepisternum with nine punctures in Figs 13, 23, 24, 29 two rows; metasternal process slightly acute apically, Type material tip rounded with an indistinct margin. Abdominal Holotype. ♂: [Indonesia], Neth. Ind.–American ventrite 1 with broadly triangular depression laterally / New Guinea Exped. / Mountain slope above / and linear depression approaching coxal articulation. Bernhard Camp 100 m / 8.IV.1939 L.J. Toxopeus Metatarsomere 4 with outer lobe moderate, length // Dobodura / armata / Darl. // RMNH / Museum 0.37× median tarsomere length. Leiden // HOLOTYPE / Dobodura / toxopei / J.K. Male genitalia. The aedeagal median lobe shaft is Liebherr 2016 [black-bordered red label] (RMNH). straight, the apex distinctly upturned beyond the ostial opening, and the apex is parallel-sided with the tip Diagnosis rounded (Figs 21, 22). The ostial opening is oriented Beetles of this species exhibit short elytral apical toward the right side of the median lobe (Fig. 22). spines oriented in the same plane as the sutural spines (Fig. 13); this configuration is also observed Distribution in D. obtusa and D. svensoni (Figs 9, 14). However, This species is known from Madang Province, from unlike D. obtusa, the elytral basal groove is only gen- Mt. Hansemann near Madang City, the Adelbert tly rounded anteriorly at the base of the fifth elytral Range, and Simbai in the Ramu River drainage stria, and the elytral surface is glossy without any evi- (Fig. 29). These localities span two areas of fresh- dence of sculpticells on the disc. Relative to D. sven- water endemism in the Polhemus & Allen (2007) soni, beetles of this species are larger — standardized system: 1, the northern portion of the Adelbert- body length 9.3 mm versus 8.7 mm — and the neck Finisterre-Saruwaged Mountains (Area 14); and 2, is broader relative to the protruded eyes; MHW/ the adjoining inland Sepik-Ramu-Markham Basins NW = 1.77 versus 1.88 for D. svensoni. The lone (Area 16). Area 16 houses only one endemic insect specimen has four dorsal elytral setae on the left ely- species — Ciliometra sepik Polhemus & Polhemus tron and three on the right. (Hemiptera: Gerridae) — whereas Area 14 is defined by 12 precinctive insect species across three orders Description and eight families (Polhemus & Allen 2007). Con- Head with prominent, convex eyes, the outer surface versely, Area 16 houses 16 precinctive fish whereas much more convex than genal surface behind eye, Area 14 none, demonstrating the value of combining ocular ratio 1.61. Prothorax quadrate in dorsal view, a disparate array of aquatic taxa in any classification PL/MPeW = 1.0; pronotal lateral marginal bead of freshwater areas of endemism. This taxonomic little elevated, very fine, lateral marginal depression disparity, and the necessity of including as many taxa shallow and broad inside bead, widest at lateral seta; as possible when assessing areas of endemism, is re- basal transverse furrow deep and narrow medially, inforced by geographic distributions among P.N.G. lined with laterally elongate punctulae, furrow deeply Downloaded from Brill.com09/30/2021 12:34:38AM via free access

16 Tijdschrift voor Entomologie, volume 160, 2017 depressed laterally just inside marginal bead; anterior Type material transverse impression very broadly and shallowly Holotype. ♂: Papua New Guinea / Chimbu Prov- depressed medially, traceable nearly to front angle, ince, Wara / Sera Research Station / Crater Moun- but surface of pronotum convex inside angle; front tain / Research Area // VII–8 ~ 13–2001 / Bradler, angles not protruded, broadly rounded. Elytra with Jarvis, / and Svenson // HOLOTYPE / Dobodura / apical spine short, length 0.12× elytral length, and svensoni / J.K. Liebherr 2016 [black-bordered red sutural spine short, length 0.04× elytral length; para- label] (CUIC). scutellar striole evident nearly to basal elytral groove, parascutellar setae in line with isolated punctulae Diagnosis of striole; 15 lateral elytral setae arranged as 6 (1 This small-bodied species — standardized body + length 8.7 mm — shares pale leg coloration (Fig. 14) 1 + 3) + (2 + 2) with posterior pair of posterior series oriented so that they define a line passing through with D. obtusa (Fig. 9). However, the neck is even apex of lateroapical spine. Mesepisternum with seven more constricted than in beetles of that species shallow punctures in three rows; metasternal process (MHW/NW = 1.88 versus 1.77–1.83 in D. obtusa), with sides meeting at slightly acute angle, tip round- and the elytral disc is glossy without evidence of iso- ed with indistinct margin. Abdominal ventrite 1 diametric sculpticells. Like all other species in the with broad shallow, rounded depression laterally and genus save D. obtusa, the elytra have well-developed linear, punctate depression approaching coxal articu- lateroapical spines. lation. Metatarsomere 4 with outer lobe moderate, Description length 0.40× median tarsomere length. Head with eyes very prominent, convex, popeyed, Male genitalia. The aedeagal median lobe shaft is their outer surface much more convex than genal straight and the broad apical extension is curved surface behind eye, ocular ratio 1.64. Prothorax upward distad the ostial opening (Figs 23, 24). The broad in dorsal view, PL/MPeW = 0.98; prono- apex is the broadest and shortest among all six spe- tal lateral marginal bead little elevated, narrowly cies (Fig. 24). The ostial opening is oriented toward rounded, lateral marginal depression shallow and the right side of the median lobe. broad inside bead, widest at lateral seta; basal transverse furrow broad and intensely strigose medially, Distribution crossed by 4–5 longitudinal punctures on each side, This species is known only from Bernhard Camp of traceable as fine punctulae laterally to shallow linear the 1938–39 Archbold Expedition to New Guinea depression inside marginal bead; anterior transverse (Toxopeus 1940). Bernhard Camp was situated on impression absent medially, evident as short, linear the Idenburg, now Taritatu River in the Mambera- depression just inside front angles; front angles not mo Raya Regency of Papua Province, Indonesia. The protruded, subangulate. Elytra with apical spine Taritatu-Tariku-Mamberamo drainage constitutes short, length 0.12× elytral length, and sutural spine area of endemism 15 in the Polhemus and Allen short, length 0.04× elytral length; parascutellar stri- (2007) system, that was established based on two ole very finely punctate, difficult to trace; 15–16 precinctive Teinobasis damselflies (Odonata: Coe- lateral elytral setae arranged as 6 + (1 + 1 + 3–4) + nagrionidae) and two fish species. (2 + 2) with posterior pair of posterior series very close together and oriented so that they define a Habitat line passing through apex of lateroapical spine. Of the type locality, Toxopeus (1940: 279) writes: Mesepisternum with 11 shallow punctures in 2–3 “Bernhard Camp, on a dead arm of the Idenburg rows. Abdominal ventrite 1 with broad, triangular R., about 10 m above the river bed in dry season, depression laterally and linear, punctate depression but flooded in the height of the west monsoon. approaching coxal articulation. Metatarsomere 4 Dense virgin jungle, locally swampy, traversed by with outer lobe moderate, length 0.40 median tar- many creeks.” This habitat description foretells Dar- × somere length. lington’s description of the habitat of D. armata at Male genitalia. The aedeagal median lobe is sinu- Dobodura. ously recurved — as in males of D. obtusa — and the Etymology. The patronym toxopei, to be treated as a long apex is broadly upturned and slightly broader noun in the genitive case, honors Lambertus J. Toxo- apically (Figs 25, 26). Also as in D. obtusa, the ostial peus, who collected the type specimen during the opening is oriented toward the dorsal surface of the Netherlands Indian–American Expedition to Neth- median lobe, the partially inflated sac visible in both erlands New Guinea (Toxopeus 1940). left and right lateral views. Distribution Dobodura svensoni Liebherr, sp. n. This species was collected sympatrically with D. Figs 14, 25, 26, 29 alildablldooya, though D. svensoni is known only Downloaded from Brill.com09/30/2021 12:34:38AM via free access

Liebherr: Revision of Dobodura (Coleoptera) 17 from this single locality (Fig. 29). The Crater Moun- outgroup, Eudalia, comprises species that occur in tain Research Area lies within the East Papuan Cen- riparian situations: on sandy river banks, in flood de- tral Highland area of endemism (Polhemus & Allen bris, or among cobble along stream margins (Baehr 2007, Area 21). 2005, 2006). The suite of ecological situations that Etymology. The species epithet svensoni, a noun in characterize habitats of species in these three genera the genitive case, honors Gavin J. Swenson, collector are all rather generalized. Available moisture is a pre- of the holotypes of both this species and D. alild- requisite for beetle occupation of these habitats, but a ablldooya. His perspicacious collecting and generous specific stream flow is not required. As such, lowland donation to the Cornell University Insect Collection riparian habitats in New Guinea and Australia would led to this revision. differ little, or at least little enough so that the species classified as these three genera can have diversified across New Guinea and eastern Australia. Within Discussion this context, Dobodura’s geographical distribution The distribution of Dobodura carabid beetles is ex- is highly derived, being limited to areas of geologi- plainable on both a biogeographical and an ecological relief sufficient to produce cascades and torrents cal scale. On the biogeographical scale, the distribu- within which these beetles have been collected. The tion of the genus can be explained relative to those vast southern half of New Guinea does not offer this of Dobodura’s outgroups; Clarencia, Dicraspeda, and sort of microhabitat, and thus Dobodura presently Eudalia. On the ecological scale, the occurrence of occupies an island of montane stream habitats caused Dobodura beetles within their particular habitat of by the faulting and mountain building due to the torrents and cascading brooks and streams can be ex- various geological terranes impinging on the north- plained in the context of the orogenic development ern cratonic margin of New Guinea + Australia. of northern New Guinea. It is noteworthy that for This montane island of northern New Guinea has Dobodura, these two spatially and temporally dispa- been derived by the successive amalgamation of at rate scales of explanation are integrally linked. For in least two island arcs — the Papuan Arc and Solo- order to have the torrents and cascades apparently mons Arc — starting in Middle Eocene, 40–50 Mya necessary for Dobodura to persist in a particular lo- (Polhemus 2007). The remnants of the Papuan Arc cale, orogenic activity must have taken place to cre- currently comprise the East and West Central High- ate those microhabitats. Given that such orogenic lands of New Guinea; areas of endemism definable activities are concentrated in the northern half of by biotic distributions (Polhemus et al. 2004, Polhe- New Guinea, the ecological restriction to cascades, mus & Allen 2007). The subsequent suturing of the and the biogeographical restriction to northern New Solomons Arc fragments to the northern margin of Guinea become one and the same. We can examine the Central Highlands commenced during Middle these two scales of phenomena, starting at the eco- Miocene (15 Mya) in the west, resulting first in up- logical scale and them moving up to a biogeographi- lift of the mountains of northern Irian Jaya, then the cal perspective. Adelbert and Finisterre Terrane of the Huon Penin- Of all genera of Odacanthini in the Australian re- sula, followed by uplift in the Papuan Peninsula, i.e. gion, Dobodura is the only taxon restricted to New the Owen Stanley Range and Popondetta Foreland Guinea (Darlington 1968). Six other odacanthine on the east. These latter emplacements of terranes genera have species that are widespread in New onto northern New Guinea occurred during the Guinea and Australia, or contain representative spe- Pliocene, 2–5 Mya. cies in those two areas; Archicolliuris Liebke, Basis- Dobodura spp. occupy several areas of endemism tichus Sloane, Clarencia, Dicraspeda, Eudalia, and within northern New Guinea (Fig. 29). The initial Ophionea Klug (Baehr 2005). Of these, Clarencia, divergence event among the six species involved Dicraspeda, and Eudalia represent successively more isolation of the common ancestor of D. obtusa and inclusive adelphotaxa along the lineage leading to D. svensoni from the common ancestor of the other Dobodura (Fig. 8). Member taxa of all three gen- four taxa that occur north of the geosyncline asso- era occur in moist microhabitats. Clarencia beetles ciated with suturing of the Miocene Solomons Arc “are usually found in wet places, often by standing to the Australian cratonic portion of New Guinea water, either among wet leaves or in or under low (Polhemus & Polhemus 1998). Subsequent diver- vegetation (Darlington 1968: 209)”. Dicraspeda gence events north of the geosyncline progress from beetles “inhabit understory foliage of rain forest west to east, first with speciation ofD. toxopei in the (Darlington 1968: 211)”. Unlike the allopatrically Mamberamo River Valley, then with divergence of distributed Dobodura spp., many New Guinean Di- D. hexaspina in Madang Province, i.e. the Adelbert craspeda can occur sympatrically, with five of the six Mountains and Ramu River Valley. The terminal New Guinean species recorded from the vicinity of sister-species pair of this quartet comprises D. alild- the Dobodura Airfield (Darlington 1968). The third ablldooya sympatric at one locality with D. svensoni, Downloaded from Brill.com09/30/2021 12:34:38AM via free access

18 Tijdschrift voor Entomologie, volume 160, 2017 and also recorded further east in Chimbu Province Acknowledgements (Fig. 29), plus D. armata from near Dobodura Air- I thank the following institutions and curators for ac- field. This set of area relationships amongDobodura cess to Dobodura specimens (institutional codens cit- spp. is consistent with progressive colonization of the ed above): The Natural History Museum, London, northern areas from west to east. Moreover, the ances- Beulah Garner (BMNH); Bernice P. Bishop Mu- tor of the southern species pair — D. obtusa and D. seum, Honolulu, James Boone (BPBM); California svensoni — would have been isolated from the ances- Academy of Sciences, San Francisco, David H. Kava- tor of the northern quartet by the central highlands, naugh (CAS); Cornell University Insect Collection, though the ranges of D. svensoni and D. alildablldooya Ithaca (CUIC); Museum of Comparative Zoology, subsequently converged in Chimbu Province. Harvard University, Cambridge, Philip D. Perkins The south to west to east progressive colonization (MCZ); Rijksmuseum van Natuurlijke Historie— pattern of Dobodura (Fig. 29) is also observed in the Naturalis, Leiden, Hans Huijbregts (RMNH). I also cicada sister tribes Chlorocystiini and Prasiini (De thank the Australian National Insect Collection, Boer & Duffels 1996). These tribes exhibit a pat- Tom Weir collection manager, and the University tern whereby taxa in the Papuan Peninsula are sister of Alberta Strickland Museum, Danny Shpeley col- groups to those in the north coastal mountains (the lection manager, for loan of specimens used in the Bewani, Torricelli, and Prince Alexander Mountains, cladistic analysis. I thank Dave Kavanaugh for gra- i.e. areas 12 + 13 of Polhemus et al. 2004). These ciously sharing his field notes for collections of D. areas in turn exhibit a sister-group relationship to hexaspina in Madang Province, and Peter Cranston the Vogelkop of New Guinea plus Maluku, with for identifying the ceratopogonid larval head cap- these collective areas to the New Guinean Central sules packed into the D. armata hindgut. Finally, I Highlands plus Cape York, Australia. Biogeographi- commemorate this contribution to the memory of cal analysis of the cicada subtribe Cosmopsaltriina Philip J. Darlington, Jr. He transformed a difficult (Duffels & Turner 2002) employed Brooks Parsi- personal situation — an attack by a crocodile — into mony Analysis (BPA) to find a sister-area relation- a highly productive collecting expedition at Dobo- ship between the Central Highlands and northern dura, during which he completed field collections New Guinea mountains, with the Papuan Peninsula that helped him establish a classificatory framework their respective sister area. The Bismarck Archipela- for New Guinea carabid beetles. His dedication to go, and then southern New Guinea plus Cape York, our discipline remains an inspiration. Australia, successively joined these areas in the BPA. Duffels & Turner (2002) acknowledged the tension between the BPA and the original taxon-area clado- References gram, as the latter included three taxonomic groups Allen, R.T. & G.E. Ball, 1980. Synopsis of Mexican taxa that exhibited shared-area relationships between the of the Loxandrus series (Coleoptera: Carabidae: Pteros- Papuan Peninsula and northern New Guinea, and tichini). – Transactions of the American Entomological only two groups that shared the Central Highlands Society 105: 481–576. and northern New Guinea areas in their distribu- Baehr, M., 2005. A revision of the Australian odacanthine tions (Duffels & Turner 2002, fig. 13). ground beetles, including checklists for Australia and Just as with Hydraena minute moss beetles (Perkins the Papuan subregion (Insecta: Coleoptera: Carabidae). 2011), Dobodura distributions can be rectified with – Memoirs of the Queensland Museum 50: 133–194. areas of endemism based on freshwater invertebrates Baehr, M., 2006. New species and new records of the gen- (Polhemus et al. 2004, Polhemus & Allen 2007). era Dicraspeda Chaudoir and Eudalia Castelnau from One exception among currently known distributions the Papuan and Australian regions, with a nomenclato- rial note on Deipyrus Liebke (Insecta, Coleoptera, Ca- is the presence of in two adjacent areas D. hexaspina rabidae, Odacanthinae). – Spixiana 29: 51–72. of endemism (Fig. 29; areas 14 and 16, Polhemus Ball, G.E., 1985a. Characteristics and evolution of elytral et al. 2004, fig. 1). However, our present knowledge sculpture in the tribe Galeritini (Coleoptera: Carabi- of Dobodura distributions is exceedingly scanty, and dae). – Quaestiones Entomologicae 21: 349–367. future discoveries of these beetles in the cascades and Ball, G.E., 1985b. Reconstructed phylogeny and geo- torrents of New Guinean streams will test the under- graphical history of genera of the tribe Galeritini (Co- lying biogeographical hypotheses supporting those leoptera: Carabidae). – In: G.E. Ball (Ed.), Taxonomy, areas of endemism. Given the surprising elucidation Phylogeny and Zoogeography of Beetles and Ants, of five new species among specimens previously as- pp. 276–321. Dr. W. Junk Publishers, Dordrecht, sumed to represent only Dobodura armata, it seems Netherlands. highly likely that further new species will be discov- Ball, G.E. & A.P. Nimmo, 1983. Synopsis of the species of ered in areas of endemism (Polhemus & Allen 2007) subgenus Progaleritina Jeannel, including reconstruct- not presently known to house these beetles. ed phylogeny and geographical history (Coleoptera:

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Carabidae: Galerita Fabricius). – Transactions of the Lakota, J., R. Lohaj & G. Dunay, 2010. Taxonomical and American Entomological Society 109: 295–356. ecological notes on the genus Scotoplanetes Absolon, Carpenter, F.M., 1985. Philip Jackson Darlington, Jr.: with the description of a new species from Montenegro A biographical sketch. – In: G. E. Ball (Ed.), Taxono- (Coleoptera: Carabidae: Trechini). – Natura Croatica my, Phylogeny and Zoogeography of Beetles and Ants, 19: 99–110. pp. 7–10. Dr. W. Junk Publishers, Dordrecht, Liebherr, J.K., 2015. The Mecyclothorax beetles (Coleop- Netherlands. tera, Carabidae, Moriomorphini) of Haleakala¯ , Maui: Darlington, P.J., Jr., 1952. The carabid beetles of New Keystone of a hyperdiverse Hawaiian radiation. – Guinea, Part 1. Cicindelinae, Carabinae, Harpalinae ZooKeys 544: 1–407. through Pterostichini. — Bulletin of the Museum of Liebherr, J.K., 2016. Cyphocoleus Chaudoir (Coleoptera: Comparative Zoology 126: 321–564 + 4 pls. Carabidae: Odacanthini): Descriptive taxonomy, Darlington, P.J., Jr., 1968. The carabid beetles of New phylogenetic relationships, and the Cenozoic his- Guinea. Part III. Harpalinae (continued): Perigonini to tory of New Caledonia. – Deutsche Entomologische Pseudomorphini. – Bulletin of the Museum of Com- Zeitschrift 63: 211–270. parative Zoology 137: 1–253. Liebherr, J.K. & A.E.Z. Short, 2007 [2006]. Blackburnia Darlington, P.J., Jr., 1971. The carabid beetles of New riparia, new species (Coleoptera: Carabidae: Platy- Guinea. Part IV. General considerations; analysis and nini): A novel element in the Hawaiian riparian insect history of fauna; taxonomic supplement. – Bulletin of fauna. – Journal of the New York Entomological Soci- the Museum of Comparative Zoology 142: 129–337. ety 114: 1–16. Davidson, R.L. & G.E. Ball, 1998. The tribe Broscini in Liebherr, J.K. & K.W. Will, 1998. Inferring phylogenetic Mexico: Rawlinsius papillatus, new genus and new spe- relationships within Carabidae (Insecta, Coleoptera) cies (Insecta: Coleoptera: Carabidae), with notes on from characters of the female reproductive tract. – In: natural history and evolution. – Annals of Carnegie G.E. Ball, A. Casale & V. Vigna Taglianti (Eds), Atti Museum 67: 349–378. Museo Regionale di Scienze Naturali, pp. 107–170. De Boer, A.J. & J.P. Duffels, 1996. Historical biogeogra- Museo Regionale di Scienze Naturali, Torino, Italy. phy of the cicadas of Wallacea, New Guinea and the Lindroth, C.H., 1974. On the elytral microsculpture of West Pacific: A geotectonic explanation. – Palaeogeog- carabid beetles (Col. Carabidae). Entomologica Scan- raphy, Palaeoclimatology, Palaeoecology 24: 153–177. dinavica 5: 251–264. Deuve, T., 1993. L’abdomen et les genitalia des femelles de Nixon, K.C. 1999. The parsimony ratchet, a new method Coléoptères Adephaga. – Mémoires du Muséum Na- for rapid parsimony analysis. – Cladistics 15: 407–414. tional d’Histoire Naturelle Serie A Zoologie 155: 1–184. Nixon, K.C. 2002. WinClada. – Published by the au- Dick, G.T., 1993. Beaufighters over New Guinea, No. 30 thor, Ithaca, NY, USA.[Available at http://www Squadron RAAF 1942–1943. – Royal Australian Air .softpedia.com/get/Science-CAD/WinClada.shtml; Force Museum Occasional Series No. 5, 188 pp. accessed 16.ix.2016]. Duffels, J.P., 1986. Biogeography of Indo-Pacific Cica- Null, G. 1995. Weapons of Denial, Air Power and the doidea: A tentative recognition of areas of endemism. Battle for New Guinea. – Air Force History and Muse- – Cladistics 2: 318–336. ums Program, U. S. Printing Office, Washington, DC, Duffels, J.P. & H. Turner, 2002. Cladistic analysis and bio- USA, 32 pp. geography of the cicadas of the Indo-Pacific subtribe Perkins, P.D., 2011. New species (130) of the hyperdiverse Cosmopsaltriina (Hemiptera: Cicadoidea: Cicadadae). aquatic beetle genus Hydraena Kugelann from Papua – Systematic Entomology 27: 235–261. New Guinea, and a preliminary analysis of areas of en- Erwin, T.L., 2000. A new genus and species of Lach- demism (Coleoptera: Hydraenidae). – Zootaxa 2944: nophorini and two new species of Lebiini from Costa 417 pp. Rica (Coleoptera: Carabidae). – Coleopterists Bulletin Pigram, C.J. & H.L. Davies, 1987. Terranes and the accre- 54: 279–283. tion history of the New Guinea orogen. – BMR Journal Foote, B.A., 1991. Ceratopogonidae (Chironomoidea). of Australian Geology & Geophysics 10: 193–211. – In: F. W. Stehr (Ed.), Immature Insects, Vol. 2, Polhemus, D.A., 2007. Tectonic geology of Papua. – In: pp. 758–761. Kendall/Hunt Publishing Co., Dubuque, A.J. Marshall & B.M. Beehler (Eds), The Ecology of IA, USA. Papua, Part One. The Ecology of Indonesia Series 6, Goloboff, P.A., 1999. NONA (NO NAME). – Published pp. 137–164. Periplus Editions, Singapore. by the author, Tucumán, Argentina. [Available at: Polhemus, D.A. & G.R. Allen, 2007. Freshwater bioge- http://www.softpedia.com/get/Science-CAD/NONA ography of Papua. – In: A.J. Marshall & B.M. Beehler .shtml; accessed 16.ix.2016]. (Eds), The Ecology of Papua, Part One. The Ecology Habu, A., 1978. Carabidae: Platynini (Insecta Coleop- of Indonesia Series 6, pp. 207–245. Periplus Editions, tera). – Fauna Japonica, Keigaku Publishing Co., To- Singapore. kyo, 447 pp. + 36 pls. Polhemus, D.A. & J.T. Polhemus, 1998. Assembling Hinton, H.E. & D.F. Gibbs, 1969. An electron micro- New Guinea: 40 million years of island arc accretion scope study of the diffraction gratings of some carabid as indicated by the distributions of aquatic Heterop- beetles. – Journal of Insect Physiology 15: 959–962. tera (Insecta). – In: R. Hall & J.D. Holloway (Eds),

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Biogeography and Geological Evolution of SE Asia, pp. Toxopeus, L.J., 1940. Nederlandsch-Indisch Amerikaan- 327–340. Backhuys Publishers, Leiden, Netherlands. sche Expeditie naar Nederlandsch Nieuw-Guinea Polhemus, D.A., R.A. Englund & G.R. Allen, 2004. Fresh- (3e Archbold-Expeditie naar Nieuw Guinea 1938– water biotas of New Guinea and nearby islands: Analy- ’39). Lijst van verzamelstations [Netherlands Indian– sis of endemism, richness, and threats. – Conservation American Expedition to Netherlands New Guinea (3rd International. [available at: pbs.bishopmuseum.org/ Archbold Expedition to New Guinea 1938–’39). List pdf/ci-png.pdf; accessed 24.viii.2016]. of Collecting Localities]. – Treubia 17: 271–279. Polhemus, J.T. & D.A. Polhemus, 2002. The Trepobatinae Wilson, E.O. 1991. Philip Jackson Darlington, Jr. Bio- (Gerridae) of New Guinea and surrounding regions, graphical Memoirs 60, pp. 32–45. National Acad- with a review of the world fauna. Part 6. Phylogeny, emy of Sciences, Washington, DC, USA. [Available biogeography, world checklist, bibliography and final at http://www.nap.edu/catalog/6061.html; accessed taxonomic addenda. – Insect Systematics and Evolu- 5.ix.2016]. tion 33: 253–290.

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Liebherr: Revision of Dobodura (Coleoptera) 21

Supplementary material S1 The following NONA format data file (Goloboff 1999), exported from WinClada (Nixon 2002), supports the cladistic analysis of Dobodura spp. and associated outgroups. To replicate the cladistic analysis, copy all file text from “xread” to the end of the file, paste it into Notepad, and save as a file with an “.ss” suffix. This may be imported to WinClada, making certain the Input file dialog box accepts “ALL files.” Save the file in WINC format and use the “Diagnoser” toggle to view character-state distributions. xread ''25 10 Porocara_nigricollis_Oz 3102000102210–02000010010 Clarencia_quadridens_Oz 201101010103003232001100[12] Dicraspeda_quadrispinosa_Oz 1102010000030031200011002 Dobodura_armata_NG 3010111010031132321000101 Dobodura_alildablldooya_NG 3010111010031132321000??? Dobodura_svensoni_NG 3010111110031032321111??? Dobodura_hexaspina_NG 30101111100311323210102?? Dobodura_obtusa_NG 30101110100310220011111?? Dobodura_toxopei_NG 3010111010031032221010??? Eudalia_atrata_Oz 1002000000210–11000010012 ; cc + 0.24; proc /; optcode f 0.12 14.18 20.21 23.24; optcode u 13 19 22;

# $ ; cn {0 Mandibular_length short,_<=_to_~1.5_x_distance_antenna_base- labral_margin moderately_elongate,_1.5-1.75_x_distance elongate,_>1.8- 2.25_x_distance very_elongate,_>2.3_x_distance; {1 Eyes moderate large,_greatly_projected hypertrophied,_convexly_projected; {2 Eye_diameter moderate very_small; {3 Proepisternum smooth,_impunctate punctured_in_posterior_half distinctly_punctured_or_wrinkled_over_entire_surface; {4 Elytral_humerus rounded_laterad_parascutellar_striole tightly_rounded_to_angulate_laterad_base_of_fifth_interval distinctly_angulate_laterad_base_of_fifth_interval; {5 Elytral_lateral_margin convex_laterad_anterior_series_of_lateral_elytral_setae invaginated_laterad_anterior_series_of_lateral_elytral_setae; {6 Parascutellar_seta single_each_elytron double_each_elytron; {7 Dorsal_elytral_setae 3-4_or_less more_than_3-4,_4_or_5- 6_present_along_length_of_interval_3; {8 Dorsal_elytral_setae articulatory_socket_(anterior_seta)_coplanar_with_elytral_disc set_in_foveate_articulatory_socket; {9 Elytral_macrosetae present_only_on_interval_3 also_present_on_interval_5 also_present_on_intervals_5_and_7 also_present_in_intervals_1,_5_and_7 present_on_all_intervals_1-7;

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22 Tijdschrift voor Entomologie, volume 160, 2017

{10 Elytral_striae impunctate_to_minutely_punctate regularly_punctate distinctly,_broadly_and_deeply_punctate; {11 Elytral_apices meeting_at_suture,_margins_evenly_curved separately_rounded_with_space_between_elytra separately_pointed,_elytra_bispinose_apically separately_spinose,_each_elytron_with_apical_spine; {12 Elytral_apical_spine_at_apex_of_stria_3 absent present; {13 Apical_spines in_same_orientation_as_sutural_spines pointing_upward_relative_to_sutural_spines; {14 Subapical_sinuation nearly_obsolete,_elytral_apices_obliquely_ truncate evident distinct,_angularly_concave very_distinct,_with_lateroapical_spine_laterally; {15 Frons_microsculpture evident,_isodiametric_to_transverse reduced,_not_traceable_except_in_spots absent; {16 Elytral_disc_microsculpture isodiametric_mesh,_may_be_in_transverse_rows transverse_mesh evident_transverse_lines not_evident,_surface_glossy; {17 Femora concolorous_from_base_to_apex,_or_apex_only_slightly_darker with_apex_dark,_basal_1/2_-_4/5_pale; {18 Male_tergite_IX_ring_sclerite angulate_apically,_wishbone-shaped hemicircular_apically,_horseshoe_shaped; {19 Male_aedeagal_median_lobe_shaft straight_in_apical_half,_tip_may_be_curved_dorsally sinuously_recurved_in_apical_half_in_addition_to_curved_tip; {20 Aedeagal_median_lobe_apex gradually_narrowed_apicad_ostium parallel-sided,_elongate_apicad_ostium; {21 Male_aedeagal_median_lobe_apex of_moderate_length,_attenuate_or_ spoon-shaped exceedingly_elongate,_porrect; {22 Female_apical_abdominal_ventrite convex_or_slightly_emarginate_medially with_rounded_keyhole_marginal_invagination with_cordately_invaginated_margin; {23 Female_bursa_copulatrix membranous,_inner_surface_smooth sparsely_covered_with_spicules_on_inner_surface; {24 Apical_gonocoxite_2 with_2_lateral_ensiform_setae with_3_lateral_ensiform_setae with_4-5_lateral_ensiform_setae; ; tread ‘wha’ (0(9(2(1((5 7)(8(6(3 4)))))))) ;

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Liebherr: Revision of Dobodura (Coleoptera) 23

Supplementary material S2 Macrophotograph of dipteran head capsules removed from female hindgut during female reproductive tract dissection of D. armata female (Fig. 27). Head capsules determined as Ceratopogonidae by P.S. Cranston, based upon diagnostic presence of “conspicuous pharyngeal structure … consisting of two diverging arms and a series of food-filtering combs (Foote 1991).” The prognathous condition and elongate head capsule are characteristics of larvae in subfamily Ceratopogoninae (Foote 1991).

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