A synopsis of the genus Cypholoba Chaudoir (Coleoptera, Carabidae, Anthiini) known to occur in the Republic of South Africa

Abstract Nearly one third of the described species of Cypholoba Chaudoir (Coleoptera: Carabidae) are known to inhabit the Republic of South Africa. A key and diagnostic notes are provided for their identification, as well as notes about way of life for some of the species based on observations in the Kruger National Park. Fifteen species and subspecies of the genus are recorded from the Republic of South Africa; adult specimens of each species and subspecies are illustrated and information about the distribution of each species in the Republic of South Africa is summarized and mapped: Cypholoba alstoni (Péringuey), Cypholoba alveolata (Brême), Cypholoba amatonga Péringuey, Cypholoba fritschi (Chaudoir), Cypholoba gracilis gracilis (Dejean), Cypholoba gracilis scrobiculata (Bertoloni), Cypholoba gracilis zuluana Basilewsky, Cypholoba graphipteroides graphipteroides (Guérin-Méneville), Cypholoba leucospilota semilaevis (Chaudoir), Cypholoba macilenta (Olivier), Cypholoba notata (Perroud), Cypholoba oberthueri seruana Strohmeyer, Cypholoba opulenta (Boheman), Cypholoba rutata (Péringuey), and Cypholoba tenuicollis aenigma (Dohrn).


introduction
The genus Cypholoba Chaudoir is one of the most diverse lineages within the tribe Anthiini of the beetle family Carabidae, with 156 described species and subspecies (Lorenz 2005) distributed throughout southern and eastern Africa. These beetles are conspicuous elements of savanna and woodland ecosystems, where they are typically found running in bright sunshine over bare ground, or in short grasses ( Fig. 1; Marshall and Poulton 1902). Like most other members of the tribe Anthiini, species of Cypholoba have the ability to excrete formic acid from their pygidial glands as a defensive behavior (Péringuey 1896). Most species in this genus are black and many species have white setal patches or setal tufts (Fig. 1) that are thought to have evolved through mimicry of Mutillidae, Formicidae, and other stinging Hymenoptera (Marshall and Poulton 1902). These beetles are of potential interest to entomologists and evolutionary biologists studying phenomena such as mimicry, aposematic coloration, and the evolution of chemical defenses. Species of Cypholoba, like many other Anthiini, also show close associations with particular ecosystems or vegetation communities and their activity patterns are closely tied with environmental variables such as temperature and rainfall, and overall climate patterns such as seasonal monsoons (Schmidt 2001;Mawdsley et al. 2011). Given the relatively large adult body size of most Cypholoba species (length 15-33 mm), their diagnostic color and setal patterns (Strohmeyer 1928) and their conspicuous activity patterns and behaviors (Schmidt 2001), these beetles could easily be incorporated into environmental monitoring programs which track overall ecosystem condition, status, and trends.
As with other Anthiini, the taxonomic history of Cypholoba is rather convoluted. Chaudoir (1850) initially established two genera, Cypholoba on p. 43 for the single species Anthia alveolata Brême and a second, Polyhirma, on p. 44 for the group of species that included Anthia macilenta Olivier, A. gracilis Dejean, A. intermedia Boheman, A. ferretti Reiche, A. tetrastigma Chaudoir, A. leucospilota Bertoloni, A. caillaudi Gory, and A. polioloma Chaudoir. Péringuey (1896) combined these two genera in his revision of southern African Carabidae, treating Cypholoba as a synonym of Polyhirma and recognizing 27 species-level taxa, ten of which were recorded from what is now the Republic of South Africa. Further descriptive work by other workers, primarily in the east African fauna, led to the recognition of 102 species in the genus by the time of Csiki's (1929) comprehensive catalogue. In the "Coleopterorum Catalogus," Csiki (1929) followed Péringuey in placing Cypholoba in synonymy with Polyhirma. Strohmeyer (1928) was the first to recognize the page priority of Cypholoba and also the first to attempt a comprehensive revision of this group. His revision radically altered the taxonomy of the genus, recognizing just 16 species and relegating most of the former species to subspecies status. This approach came under criticism from Basilewsky (1948Basilewsky ( , 1955 who pointed out a number of errors in Strohmeyer's revision and argued that many of the so-called "subspecies" of Strohmeyer were separated by characteristics that suggested they were in fact perfectly good species. Basilewsky published a series of studies in which he argued against Strohmeyer's reductionist approach while at the same time continuing to describe new species and subspecies of Cypholoba (Basilewsky 1948;1955;1963;1967;1980;. This paper is intended to provide an overview of the species of Cypholoba currently known from the Republic of South Africa (RSA). This is a fauna very much in need of good diagnostic materials, particularly keys and illustrations that can be used by nonspecialists. Most of the Cypholoba species in RSA have never been illustrated and none of the published keys include all of the taxa now known to occur in RSA. The government of RSA has taken recent positive steps towards protecting certain carabid beetles under the South African Biodiversity Act of 2004 (Harrison and Müller, pers. comm.) and carabid beetles are increasingly being incorporated into ecosystem and agricultural monitoring programs in southern Africa (Kotze 2000;Magagula 2003). Both types of conservation approaches (carabid beetles as the subject of direct conservation efforts, and carabid beetles as environmental monitoring targets and ecological indicators) are clearly contingent on the availability of high-quality identification materials for the carabid fauna of interest. Fortunately, development of these identification materials is relatively straightforward. The species of Cypholoba from RSA are well represented in museum collections, as a result of large-scale survey activities that began in the 1950s and have continued more or less until the present day. The species-level taxonomy of the RSA fauna is also reasonably well known; there have been only modest changes in the taxonomy of the RSA species of Cypholoba since the first revision by Péringuey (1896), despite the major changes introduced by Strohmeyer (1928) in other parts of the genus. In the synopsis that follows, we follow the classification of this genus presented by Lorenz (2005) in the most recent catalogue of world Carabidae. It is hoped that this short communication will help to inspire further interest and field studies of these remarkable beetles.

Materials and methods
We examined collections of adult Cypholoba Chaudoir and allied genera in the following institutional collections: Field Museum of Natural History, Chicago, Illinois (FMNH); Kruger National Park Museum (Scientific Services), Skukuza, South Africa (KNPC); South African National Collection of Insects, Pretoria, South Africa (SANC); National Museum of Natural History, Smithsonian Institution, Washington, D.C. (NMNH); Transvaal Museum, Pretoria, South Africa (TMSA). In the case of NMNH and TMSA, the specimens examined were authoritatively identified by the late P. Basilewsky, who studied this genus for many years (Basilewsky 1948;1955;1963;1967;1980;. Museum collections were augmented by a series of field visits in 2007, 2008, 2009, 2010, and 2011 to the Kruger National Park in northeastern South Africa, where adults of three species (C. alveolata, C. graphipteroides, and C. notata) were collected. Survey methods for Cypholoba species and other Anthiini involved systematic walking along roads, dry river washes, or sandy areas adjacent to major rivers. Surveys in the main park area were conducted from dawn until dusk and under all available weather conditions. Head-lamping surveys were conducted at night for Anthiini and other Carabidae in the N'waswitshaka Research Camp at Skukuza. Pitfall traps were also widely deployed throughout the Skukuza Ranger District of the Kruger National Park, to study the distribution of species of Carabidae across different vegetation communities and to record the responses of carabid assemblages to common landscape-scale disturbances such as mammalian grazing and ground fire. Driving surveys  were also employed as a survey technique for larger Anthiini, as well as species of Tefflus Leach (Coleoptera: Carabidae: Panagaeini); adults of larger species of Cypholoba such as C. graphipteroides can be detected during driving surveys. When encountered during diurnal surveys, the individual beetles were highly conspicuous and could be easily captured by hand. Further information about our survey methods is available in papers by Mawdsley and Sithole (2008;2009), Mawdsley (20092011), andMawdsley et al. (2011).
Methods and species concepts follow those previously described (Erwin and Kavanaugh, 1981;1991). The species validation and diagnosis format follows as closely as possible that suggested in Erwin and Johnson (2000). Measurements of length (ABL) and width (TW) follow those of Ball (1972) and Kavanaugh (1979): ABL (apparent body length), measured from apex of labrum to apex of longer elytron.
punctures with reddish-orange pubescence at the bottom of the punctures (see Fig.  3). It can be separated from C. macilenta, C. notata, and C. oberthueri seruana by the lack of a white patch of setae across both elytral apices and from C. amatonga by its larger ABL (24-28 mm length in C. amatonga) and the differences in elytral surface sculpture mentioned in the key above. Some specimens have a patch of white setae on each elytron immediately adjacent to the scutellum, while others also have a patch of white setae on the scutellum and/or a line of white setae along the midline of the pronotum.
Materials examined. 159 specimens from the following localities: RSA: Gaut- Notes on biology. During our surveys (see Materials and Methods above), adults of this species were found in riverine and upland areas of the Kruger National Park. Specific vegetation communities (Gertenbach 1983) where adults were collected included riverine gallery forest and upland Acacia nigrescens Olivier -Combretum apiculatum Sonder savanna. Adults were collected diurnally, in pitfall traps, and with headlamps at night.  (Péringuey) Strohmeyer (1928:448) Diagnosis. Apparent body length (ABL) 24-28 mm; similar to C. alveolata but separated from that species by the smaller ABL (31-33 mm in C. alveolata) and by the differences in elytral surface sculpture mentioned in the key above.

Cypholoba amatonga
Materials examined. 2 specimens from the following locality: RSA: Limpopo Province: Soutpansberg.  (Bertoloni) Strohmeyer (1928:315) Diagnosis. Apparent body length (ABL) 19-20 mm; similar in general body form and appearance to C. g. gracilis but differing from that subspecies in having the median line of the pronotum not markedly impressed and in having the elytral punctures oval rather than round.
Notes on taxonomy. Based on the rather significant differences in the pronotal and elytral surface sculpture, C. g. scrobiculata may ultimately prove to represent a species distinct from C. gracilis. A thorough review of the C. gracilis species complex with a rigorous evaluation of the 41 subspecific taxa proposed by Strohmeyer (1928) is needed. Basilewsky, 1948 http://species-id.net/wiki/Cypholoba_gracilis_zuluana Figs 10, 30

Cypholoba gracilis zuluana
Cypholoba gracilis zuluana Basilewsky (1948:111, type locality "Zululand," syntype series in Naturhistoriska Riksmuseet, Stockholm) Diagnosis. Apparent body length (ABL) 19-20 mm; similar in general body form and appearance to C. g. gracilis and C. g. scrobiculata but differing from those two subspecies in the pronotal surface sculpturing, as noted in the key. There are also subtle differences in the arrangement and size of the elytral punctures and elytral costae between all three subspecies (see Figs 6-10). Materials examined. 6 specimens from the following localities: RSA: KwaZulu/ Natal Province: "E. Zululand," "Zululand."

Cypholoba graphipteroides graphipteroides (Guérin-Méneville, 1845)
http://species-id.net/wiki/Cypholoba_graphipteroides_graphipteroides Figs 1,11,12,13,14,15,26,31,33 Anthia graphipteroides Guérin-Méneville (1845:285, type locality "in regione Massilicatzi," holotype in Museum National d'Histoire Naturelle, Paris) Polyhirma graphipteroides (Guérin-Méneville) Péringuey (1896:346) Cypholoba graphipteroides graphipteroides (Guérin-Méneville) Strohmeyer (1928:343) Diagnosis. Apparent body length (ABL) 24-27 mm; this species is easily separated from most of the other sympatric species of Cypholoba by the distinctive pattern of pubescence and setae on the elytra (Fig. 11). Adults could potentially be confused with C. leucospilota, but in that species the pubescence on the pronotum and elytral suture is yellowish and the patches of setae at apical third are brilliant white (Fig. 16). Notes on biology. This is easily the most abundant and frequently encountered species of Anthiini in the Kruger National Park, RSA. Adults (Fig. 1) emerge in the early rainy season (November-December) and can be locally abundant. Adults are active both nocturnally and diurnally and are typically found in riverine or riparian areas near flowing water. The charactistic behavior observed in this species, as with most Anthiini, is a rapid walking behavior which appears to be associated primarily with foraging but also is likely involved in searching for conspecifics and prospective larval habitats. We have observed adults of this species in several distinct vegetation communities (riverine gallery forest, open Combretum apiculatum -Acacia nigrescens savanna, Phragmites reed beds) and microhabitats (sand or dirt roads adjacent to riverine communities, dry sand wash, and riverine sand bars). This species also occasionally enters human-inhabited areas; we have found individuals in the N'waswitshaka Research Camp at Skukuza and also in the main tourist areas at the Skukuza tourist camp. Marshall and Poulton (1902) suggest that this species may be a mimic of Mutillidae and other stinging Hymenoptera. Our observations suggest that, while there are certainly similarities in color pattern and behavior between adults of C. graphipteroides and sympatric Mutillidae, adults of C. graphipteroides are actually significantly larger in size than most of the sympatric black-and-white mutillid wasps, rendering the resemblance less than exact. We infer from these observations that selection pressures may not be operating as intensely on this species as on other Carabidae (for examples of carabid beetles which are much more convincing mimics of Mutillidae, see Marshall and Poulton 1902).
Notes on taxonomy. Strohmeyer (1928) recognized 20 subspecies in C. graphipteroides, many of which were separated on the basis of differences in the setal patterns of the elytra. Many of these taxa are doubtfully distinct from the nominate form and the whole group is in need of a careful revision. Such a revision should include examination of extensive series in order to determine the extent of intrapopulational variation in the elytral setal patterns. Figures 12-15 illustrate the variation in elytral setal patterns within a single population of this species, located west of Paul Kruger Gate along the Sabie River in the Kruger National Park.
other specimens lack these bands (Figs 14, 15). We have also examined highly abraded specimens that lack all of the elytral and pronotal setae but in these specimens the elytral surface sculpturing is still diagnostic.
Notes on biology. We observed adults of this species running along dirt roads in bright sunshine through open Combretum apiculatum -Acacia nigrescens savanna in close proximity to the Sabie River in the Kruger National Park, RSA. Strohmeyer, 1928 http://species-id.net/wiki/Cypholoba_oberthueri_seruana Figs 21, 31

Cypholoba oberthueri seruana
Cypholoba semisuturata seruana Strohmeyer (1928:337, type locality "Ost-Betschuanaland," one syntype from "Sogosse" the other syntype from "Serue" in Museum für Naturkunde, Berlin) Diagnosis. Apparent body length (ABL) 21-24 mm; similar in appearance to C. macilenta and C. notata but differing from both species in the length of the elytral costae (as noted in the key) and the elytral proportions which are somewhat longer and narrower in C. o. seruana than in the other two species. This is apparently a rare form, at least in collections; it is known at present from two localities in Botswana and the nominate subspecies is known only from Zimbabwe (Strohmeyer 1928 Polyhirma opulenta Boheman (1860:9, type locality "juxta fluvium Svakop," holotype in Naturhistoriska Riksmuseet, Stockholm) Cypholoba opulenta (Boheman) Strohmeyer (1928:367) Diagnosis. Apparent body length (ABL) 18-24 mm; easily recognized by the goldenyellow pubescence on the head, pronotum, and base of elytra adjacent to the suture (Fig.  22). The elytral surface sculpturing is also diagnostic, with small narrow costae separated by flat intervals with a single row of minute punctures in each interval (Fig. 28).