Resolution of taxonomic problems in Australian Harpalini, Abacetini, Pterostichini, and Oodini (Coleoptera, Carabidae)

Abstract Taxonomic changes are made for several problematic Australian Carabidae in the tribes Harpalini, Abacetini, Pterostichini, and Oodini. Examination of types resulted in the synonymy of Veradia Castelnau, 1867 with Leconomerus Chaudoir, 1850; Nelidus Chaudoir, 1878, Feronista Moore, 1965, and Australomasoreus Baehr, 2007 with Cerabilia Castelnau, 1867; and newly combining Fouquetius variabilis Straneo, 1960 in the genus Pediomorphus Chaudoir, 1878; Australomasoreus monteithi Baehr, 2007 in the genus Cerabilia Castelnau, 1867; and Anatrichis lilliputana W.J. Macleay, 1888 in the genus Nanodiodes Bousquet, 1996. Cuneipectus Sloane, 1907 is placed in Pterostichini Bonelli, 1810, which is a senior synonym of Cuneipectini Sloane, 1907.


Introduction
In a continuing effort to make the faunal list of Australian carabid beetles as accurate as possible, I sought out and studied type specimens for a number of historically problematic taxa. Results of my study require a number of adjustments to recognized taxa.
Notes. Originally this species was placed by Castelnau (1867) near Moriodema Castelnau, 1867, a Moriomorphini taxon, which was then considered to be within Pterostichini. Subsequently it was moved to Harpalini by Chaudoir (1880) and according to Chaudoir it did not differ from Hypharpax W.S. Macleay, 1825. Sloane (1898) agreed with the placement in Harpalini, but deferred on the generic assignment and its possible similarity to Hypharpax. Straneo (1941) thoroughly reviewed the pertinent literature and studied the type specimen of Veradia brisbanensis. He concurred with the placement in Harpalini and suggested there were similarities with Nemaglossa Sloane, 1920(=Lecanomerus Chaudoir, 1850, not Nemaglossa Solier, 1849, Euthenarus Bates, 1874 and Diaphoromerus Chaudoir, 1843 (= Notiobia (Anisotarsus) Chaudoir, 1843). These taxa fall in three different tribes of Harpalini and Straneo pointed out that without access to Australian material for comparison that he could not make a decision regarding the status or relationships of the genus and species. Moore et al. (1987) maintained the genus in Harpalitae incertae sedis, accurately reflecting the uncertainty of the placement of the taxon at that time.
I examined the holotype and confirm that it has typical Harpalini character states, e.g. single supraorbital seta and no elytral plica, and does not have any character states that would place it in any other tribe. Additionally the male has the front and middle tarsomeres expanded with spongy ventral pads, the penultimate labial palpomere is bisetose, the posterior lateromarginal seta of pronotum is absent and the angular base of stria 1 is absent. This combination of character states is consistent with placement of this taxon in subtribe Pelmatellina and is identical to the state combination found in many Australia Lecanomerus species. Based on this evidence, Veradia is considered a junior synonym of Lecanomerus.
A search in the holdings of the ANIC and QM did not yield any additional specimens of this species beyond the single female, but at least six very similar looking Lecanomerus species were found. Each was distinctly different, but all are very likely closely related based on their general similarity. How many of these are currently named species cannot be assessed without recourse to the types.

= Fouquetius variabilis
Notes. Straneo (1960) discusses at length his sense that Holconotus Schmidt-Goebel, 1846 (= Fouquetius Maindron, 1906) and Pediomoprhus Chaudoir, 1878 are closely related and that Pediomorphus macleayi Sloane, 1900 could be a species of Holconotus. Moore (1965) confirmed that P. macleayi is a true Pediomorphus. Straneo's conclusions are based on very limited material and he did not discuss characters that allow for clear placement of species in these two genera. Among other characteristics, Pediomorphus has distinctly expanded penultimate labial palpomeres not found in Holconotus, while the elytral lateral bead is distinctly, finely serrate in Holconotus and smooth in Pediomorphus. The type specimen of Pediomorphus variabilis has clearly expanded penultimate labial palpomeres and smooth elytral lateral beads. Given the new combination, Holconotus is removed from the Australian faunal list.  (2011) includes Australian species placed in Feronista by Moore et al (1987) and Cerabilia species from New Zealand and New Caledonia. Baehr (2007) described Australomasoreus monteithi as a Masoreini, but he clearly noted that this placement was both anomalous for the species' characteristics and biogeography. Study of the type and additional material for both morphology and DNA data (Will unpubl.)

clearly places this species in Cerabilia.
Cerabilia australis is known only from the holotype specimen and was reported as coming from the Paroo River area. However, this specimen is unlike any Australian species of carabid and is very similar to Cerabilia species from New Zealand. It may in fact be a synonym of one of the described New Zealand species, but until their types are studied this cannot be established. The Australian Cerabilia species are all restricted to the higher elevation rainforests in the northeastern coastal region. The Paroo River runs through the semi-arid inland region of southwestern Queensland and northwestern New South Wales and is both geographically and environmentally distant from any location where Cerabilia has been found in Australia. Likely the type locality was erroneously reported.  Sloane, 1915 [EMEC].
Notes. Sloane described a new tribe for Cuneipectus suggesting that it belonged "at the beginning of the Trigonotomid series of the subfamily Harpalinae", i.e. as sister to a group Pterostichini. Subsequent authors have placed it between Harpalini and Chlaeniini (Csiki 1931), near chaetogenyines, chlaeniines, oodines, and licinines (Callistitae sensu Erwin and Sims (1984) and Erwin (1985Erwin ( , 1991) in Licininae (Lorenz 2005) in Pterostichitae (Moore et al. 1987) or Pterostichini (Lawrence and Slipinski 2013). Moore (1965) did not include Cuneipectus in his treatment of Australian Pterostichinae. Aside from the original description, there has not been a discussion of the characteristics of Cuneipectus. Its variable placement, non-inclusion in Moore's (1965) treatment and frequent association with Chlaeniini and Licinini by various authors apparently stems from the species being described as having a single supraorbital setae in combination with the presence of an elytral plica. However, supraorbital seta number is variable, with some individuals having one and others two above each eye. Other characteristics are typical of Australian Pterostichini, including the presence of the spermathecal gland duct diverticulum (sgd) in the female (Liebherr and Will 1998). The sgd is typical in many pterostichines including Australian taxa like Prosopogmus Chaudoir, 1865(Will 2011), Paranurus Tshitshérine, 1901(Liebherr and Will 1998 and Trichosternus Chaudoir, 1865 (Will unpubl.). The sgd is not known to be present in any Chlaeniini or Licinini. Additionally, preliminary analyses of DNA data (Will unpubl.)  Notes. Nanodiodes Bousquet, 1996was proposed by Bousquet (1996 to replace Nanodes Habu, 1956 and he moved all species that where included by Moore et al. (1987) in Anatrichus LeConte, 1853 into this genus except for Anatrichis lilliputana, which Bousquet had not studied. Although some subsequent catalogs (e.g., Lorenz 2005) treated this species as Nanodiodes lilliputana, there is no indication that the character states were confirmed. I examined the syntypes and found the following: submentum with pairs of setae at the lateral edge; mesocoxa with a posterior seta and; metatrochanter without a seta. This combination is consistent with Nanodiodes, confirming that it shares the putative synapomorphic character states with species currently included in that genus. Anatrichis is therefore not found in the Australian fauna.

Acknowledgments
I thank Cate Lemann, Australian National Insect Collection, CSIRO, who contributed significantly to this effort by seeking out, examining and imaging types and other material critical to this communication. I also thank Nadine Guthrie, Department of Parks and Wildlife, Western Australia for examining specimens of Oodini and Dr. Eva Sprecher-Uebersax of the Naturhistorisches Museum, Basel for taking and sending images of the type of Fouquetius variabilis. I thank Geoff Monteith, Queensland Museum for the extremely helpful suggestions and corrections in his review of the manuscript.