Rediscovery of Clivina morio Dejean with the description of Leucocara, a new subgenus of Clivina Latreille (Coleoptera, Carabidae, Clivinini)

Leucocara, a new subgenus of Clivina Latreille, is established for C. americana Dejean and its relatives, whose collective geographical range includes the Western Hemisphere Nearctic Region and the Eastern Hemisphere Palaearctic, Oriental, and Afrotropical Regions. Previously, these taxa were included in the subgenus Reichardtula Whitehead, 1977, a taxon now confi ned to the Eastern Hemisphere. Members of Leucocara diff er from those of other Nearctic Clivina by the presence of a small, apically truncate preapical protuberance on the mesotibia with its seta inserted apically. Clivina morio, previously known only from the holotype, is reported here from Louisiana and Texas, confi rming its presence on the North American continent; the species is also included in Leucocara. Th e following new synonymies are proposed: C. dilutipennis Putzeys, 1866, C. insularis Jacquelin du Val, 1857, C. klugii Putzeys, 1846, C. sculptifrons Putzeys, 1846 with C. fasciata Putzeys, 1846 and C. morula LeConte, 1857 with C. americana Dejean, 1831.


Introduction
Clivina morio was described in 1831 by Count Dejean from a single specimen reported from "Amérique septentrionale."Th e species was transferred to the genus Ardistomis Putzeys by Putzeys (1866: 205).Subsequently it was recorded from Lancaster County in Pennsylvania (Rathvon 1869: 524), Georgia (LeConte 1879: 32), the vicinity of New York city (Leng and Beutenmuller 1893: 135), Fort Myers in Florida (Leng 1915: 571), and Th omasville in Georgia (Fattig 1949: 15).Bousquet (2006) reviewed the Nearctic species of Ardistomina based on materials from 11 major North American collections, including the California Academy of Sciences, the Canadian National Collection of Insects, the Florida State Collection of Arthropods, the Museum of Comparative Zoology, and the National Museum of Natural History, and was unable to fi nd a single specimen conspecifi c with the holotype of C. morio.Th is led him to believe the species may inhabit the Neotropical Region and that the region originally mentioned by Dejean was in error.Bousquet (2006) transferred the species back to the genus Clivina Latreille but did not provide a subgeneric placement for it.
In 2008, Igor S. Sokolov found two specimens in the Louisiana State Arthropod Collection that belong to C. morio and sent them to me for confi rmation.Th is fi nding led me to investigate the systematic position of this species and its putative relatives in both the Western and Eastern Hemispheres.

The Western Hemisphere subgenera of Clivina
As discussed by Ball (2001), the Western Hemisphere species of Clivina Latreille belong to fi ve subgenera: Paraclivina Kult, Semiclivina Kult, Antroforceps Barr, Clivina s. str., and Reichardtula Whitehead.A list of the species, assigned to the respective subgenera, is includes in Table 1.Members of Paraclivina are characterized in having the mandibular scrobe distinctly laterad, the lateral bead of pronotum without posteriolateral denticle and not extended to the basal edge but extended parallel anterior to it, and the elytral interval 3 with only two discal setigerous punctures.Th is taxon corresponds to groups 21 and 22 of Putzeys (1866) and is indigenous to the New World.It contains at least 33 species in the Nearctic and Neotropical Regions.According to Nichols' (1988)  Th e subgenus Semiclivina is characterized mainly by the presence of a narrow, sculptured band [deep longitudinal or submarginal furrow of Kult (1947: 31), submarginal ridge of Nichols (1988: 154), longitudinal carina of Ball (2001: 136)] extended more or less parallel to the lateral edge.Th is state is likely synapomorphic and suggests that the taxon is monophyletic.Th e group is indigenous to the Western Hemisphere and includes at least nine species (Table 1).Probably many other Middle American and South American species, placed by Putzeys (1866) in his group 24, belong to this subgenus.
Antroforceps Barr is indigenous to northeastern Mexico (Tamaulipas) and eastern United States.It includes three species (Table 1) and is recognized by the bidentiform lateral edge of pronotum posteriorly and bidentiform or crenulate humeri.Th e remaining two subgenera are inadequately characterized.Adults of the nominotypical taxon lack the paramedian sulci on abdominal sternum III as in members of Paraclivina but contrary to those of other Western Hemisphere taxa.As presently conceived, Clivina s. str. is worldwide, markedly speciose, and in my opinion possibly paraphyletic, if not polyphyletic.Nine species inhabiting the Western Hemisphere are currently assigned to this subgenus (Table 1).
Th e only distinctive or peculiar structural character state for Reichardtula is that the two setigerous punctures on each side of abdominal sternum VII (the so-called "last visible sternum") are close to each other.Th e subgenus contains at least two distinct groups of species.One has the preapical protuberance on the mesotibia small, truncate at apex, its seta inserted apically (Fig. 2).Th e second group has that protuberance long, acuminate at apex, its seta inserted laterally (Fig. 3).In view of the current classifi cation of the genus Clivina, I believe these two groups should each be given subgeneric rank.Because Reichardtula is a replacement name for Eupalamus Schmidt-Göbel, both have the same type species (ICZN 1999, Article 67.8) which is the Oriental Clivina castanea Westwood, 1837, a senior synonym of Eupalamus clivinoides Schmidt-Göbel the sole species originally associated with Eupalamus.Th at species, based on the description and keys to Indian species by Andrewes (1929: 353-355, 374-375), has "a spur above apex" of the mesotibia (e.g., with a relatively long protuberance) contrary to some species which are "without spur above apex" (e.g., with a small protuberance).Th erefore, the species of the americana group (see Bousquet and Larochelle 1993: 103) are those in need of a new subgenus.
Clivina morio has a small, truncate protuberance on the mesotibia and a close examination of the new material at hand shows that the species is externally markedly similar to this species of the americana group.Th erefore the species is placed in this group which is given herein the subgeneric name Leucocara (see Table 2 for list of Etymology.From the Greek leukos (white) and kara (head).Th e name is proposed in memory of Donald Robert Whitehead  who had an interest in Clivina and the Clivinini in general.His family name has been used in the past to denote a scaritine genus (Whiteheadiana Perrault) and a curculionid genus (Whiteheadia Alonso-Zarazaga & Lyal).
Recognition.Members of this subgenus diff er from those of other Nearctic Clivina by the presence of a small, apically truncate preapical protuberance on the mesotibia with its seta inserted apically.
Description.Head.Supraantennal lobes not prominent, distinctly posteriad anterior edge of clypeus.Lateral wings of clypeus isolated from median portion by extension of frontal impressions; median portion of clypeus coarsely beaded.Labrum with seven long setae.Mandible with scrobe depressed, not distinctly laterad, evident from dorsal aspect.Labial mentum with prominent U-shaped ridge; glossal sclerite acutely carinate medially, carina not sinuate; apex of glossal sclerite with one long seta medially; paramedian pit organs widely separate medially.Prothorax.Pronotum with lateral bead extended to basal edge; posteriolateral angle delimited, angulate; side without accessory dentiform projection posteriad posterior angle.Proepisternum without sculptured band.Elytra.Lateral gutter clearly extended inside humerus, humeral portion clearly delineated; umbilical setae not set up in ringed depressions.Interval 3 with three or four discal setae; second discal seta not adjoining stria 3 though close to it in some individuals, in most individuals rela- tively close to stria 2 or in middle of interval 3; interval 8 carinate toward apex and briefl y toward base.Striae 4 and 5 joined at base.Legs.Profemur ventrally with a small dentiform projection toward apex.Mesotibia with preapical apophysis small, seta apicad (Fig. 2).Abdomen.Sternum III with coxal lines medially; sternum VII with preapical setae on each side proximate, distance between them less than that between medial setae.
Phylogenetic status.A small, truncate protuberance of the mesotibia (Fig. 2) is probably the plesiomorphic state among the Clivinini and does not support the idea that Leucocara is monophyletic.In fact, the genus Clivina is a large, inadequately defi ned complex and consequently the structural characters are diffi cult to polarize.Th erefore, I am unable to off er any morphological evidence that Leucocara is natural.However, the species, at least those I have seen (see "Species included" section), are extremely similar to one another in external features and this is suffi cient grounds for recognition of the group until it is subjected to phylogenetic analysis.
Geographical distribution.Th e known range of Leucocara includes the Western Hemisphere Nearctic Region, and the Eastern Hemisphere Palaearctic, Oriental, and Afrotropical Regions.
Species included.Names of species belonging to Leucocara are listed in Table 2. Taxonomic remarks about selected taxa are as follows.

Th e Western Hemisphere species
Based on Bousquet and Larochelle (1993: 103), the americana group contains fi ve species in North America: C. americana Dejean, C. analis Putzeys, C. californica Van Dyke, C. morula LeConte, and C. rufa LeConte.However, in his unpublished thesis, Nichols (1988: 148) revalidated C. acuducta Haldeman (previously in synonymy with C. americana), synonymized C. morula with C. americana [new synonymy] and reinstated C. analis as a synonym of C. americana.Th is leaves the americana group with fi ve valid North American species: acuducta, americana, californica, morio, and rufa.All these species, except C. californica which is known only from the type locality in Lake County, California, are found in the eastern part of the continent.Th e group, as far as known, as no representatives in the Neotropical Region.

Th e Eastern Hemisphere taxa
Despite having seen but few species, there is little doubt that Leucocara is well represented in the Eastern Hemisphere.Th e tranquebarica group, of which I have seen three species only, includes 15 species (Kult 1951: 18-24) and the natalensis group, of which I have studied but one species, consists of 43 species in Africa (Kult 1959: 179-206).One Asian species, C. zebi Kult, reaches the Australian Region where it is found in New Guinea, New Britain, and Australia (Darlington 1962: 362).Th e three Afrotropical species of the rugiceps group probably also belong to Leucocara.According to Kult (1959: 176), adults of the rugiceps group have also a small mesotibial protuberance but contrary to those of the natalensis-group have only one pair of preapical setae (instead of two) on abdominal sternum VII.On the other hand, the species of the attenuata group of the Oriental region (fi ve species), despite having a small mesotibial protuberance, have the preapical setae equidistant on abdominal sternum VII (Kult 1951: 18), no discal setae on interval 3 (Kult 1951: 18), the labrum with six setae (Andrewes 1929: 353), and the glossal sclerite with two apical setae (checked on C. striata Putzeys only).In my opinion, this group of species is probably not closely related to Leucocara and would need a new subgeneric name.All remaining groups of species previously included in the subgenus Reichardtula have a long, apically acuminate mesotibial protuberance (Fig. 3).As now restricted, this subgenus is probably, in my opinion, more closely related to Clivina s. str.and Semiclivina than to Leucocara.Th e only known signifi cant character state shared between Reichardtula and Leucocara is the condition of the preapical setae on each side of sternum VII being proximate.Even if this character is eventually proven to be synapomorphic for these taxa, the clear, unambiguous diff erence in the mesotibial protuberance would justify the recognition of two distinct subgenera, considering the current classifi cation schema of the genus Clivina.

Table 1 .
List of Western Hemisphere Clivina with their subgeneric assignment

Table 2 .
List of species belonging or assumed to belong to Leucocara.Species marked with an asterix (*) have been studied.