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Downloaded from Brill.Com09/30/2021 12:34:38AM Via Free Access 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 <UN> 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 <UN> 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
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