taxonomy, ecology, and phylogeny of species of lophyra motschulsky 1859, subgenus

Tropical Zoology 22: 57-70, 2009

Taxonomy, ecology, and phylogeny of species of Lophyra Motschulsky 1859, subgenus Eriolophyra Rivalier 1948 (Coleoptera Cicindelidae)

Jonathan R. Mawdsley

Department of Entomology, National Museum of Natural History, Smithsonian Institution, P.O. Box 37012, Washington, DC, 20013-7012, USA(E-mail: [email protected])

Received 30 December 2008, accepted 19 March 2009

A key is provided for the identification of the five species of the subgenus Eriolophyra Rivalier 1948 of the genus Lophyra Motschulsky 1859 (Coleoptera Cicindelidae): L. (E.) alba (Horn 1894); L. (E.) albens (Horn 1895); L. (E.) arnoldi (Horn 1904); L. (E.) barbifrons (Boheman 1848); and L. (E.) somalia (Fairmaire 1882). The taxonomy of this sub-genus is reviewed and Lophyra (Eriolophyra) barbifrons marqueza (Péri-nguey 1896) is placed as a synonym of L. (E.) barbifrons. Information about the ecology of species of Lophyra (Eriolophyra) is summarized: Lophyra (Eriolophyra) alba, L. (E.) albens, and L. (E.) arnoldi are associ-ated with sandy riverine areas in southern, central, and west-central Afri-ca, respectively, while L. (E.) barbifrons and L. (E.) somalia are associated with coastal sand beaches in southern and eastern Africa, respectively. Descriptions of adult foraging, thermoregulatory, reproductive, and defen-sive behaviors are provided for L. (E.) alba, based on recent observations at sites along the Letaba River in Kruger National Park, South Africa. A hypothesis of phylogenetic relationships is proposed for the five species of the subgenus Eriolophyra on the basis of adult morphological characteris-tics. Morphological characters suggest that this group forms a monophy-letic clade within the broader complex of species allied to Lophyra.

key woRds: Eriolophyra, Cicindelidae, ecology, phylogeny, morphology, distribution.

Introduction . . . . . . . . . . . . . . . 58Genus Lophyra Motschulsky 1859 Subgenus Eriolophyra Rivalier 1948 59Key to Species . . . . . . . . . . . . . . 60

Lophyra (Eriolophyra) alba (Horn 1894) . . . . . . . 60Lophyra (Eriolophyra) albens (Horn 1895) . . . . . . . 61Lophyra (Eriolophyra) arnoldi (Horn 1904) . . . . . . . 61Lophyra (Eriolophyra) barbifrons (Boheman 1848) . . . . . 62

58 J.R. Mawdsley

Lophyra (Eriolophyra) somalia (Fairmaire 1882) . . . . . 63Ecology . . . . . . . . . . . . . . . . 64Phylogenetic relationships . . . . . . . . . . . 66Acknowledgments . . . . . . . . . . . . . 68References . . . . . . . . . . . . . . . 68

INTRODUCTION

Tiger beetles (Coleoptera Cicindelidae) have long been one of the most popular insect groups among amateur and professional entomologists (acoRn 2001, PeaRson & VogleR 2001). In recent years, these beetles have also come to the attention of conservation biologists, due to the fact that population declines have been noted in many species. Many of these declines can be attributed to human activities, including urbanization, dam construction, and recreational use of sensitive habitat areas (gRaVes & BRzoska 1991, knisley & schultz 1997). As a result, species of tiger beetles are protected under the biodiversity laws of many countries (PeaRson & cassola 1992, cassola & PeaRson 2000, PeaRson & VogleR 2001, PeaRson et al. 2006).

Coastal and riverine tiger beetle species appear to be especially sensi-tive to anthropogenic disturbances (knisley & schultz 1997). Human recre-ational use of coastal and riverine beaches has been directly responsible for declines in several North American tiger beetle species, including Cicindela dorsalis dorsalis Say 1817 and Cicindela puritana Horn 1871, whose larvae are easily trampled by vehicular and human foot traffic on beaches (hill & knisley 1993, 1994).

Information about the ecology and taxonomy of tiger beetles has played an important role in the development of conservation strategies for these insects (knisley & schultz 1997). While much is known about the ecology of North American and European tiger beetles (knisley & schultz 1997, PeaRson et al. 2006), very little has been published on the ecology of most African tiger beetle species (guPPy et al. 1983, FahR 1998, Mawdsley & sithole 2008).

The present paper is an outgrowth of recent ecological studies of river-ine tiger beetles in Kruger National Park, South Africa (Mawdsley & sithole 2008). These surveys have produced the first detailed information about the ecology of the riverine species Lophyra (Eriolophyra) alba (Horn 1894), which is presented for the first time below. In preparing the notes on this species for publication, however, it became apparent that a more substantial paper was needed on the taxonomy of the subgenus Eriolophyra Rivalier 1948 of Lophyra Motschulsky 1859. Although the five species of this subgenus were recently illustrated by weRneR (2000), this work did not include a key to the species of this subgenus, nor did it include an analysis of the phylogenetic relationships among species of Lophyra (Eriolophyra) using modern cladistic approaches. I have therefore reviewed the taxonomy of this group and pre-pared a key to species and a provisional cladistic analysis of the species of this subgenus.

59Taxonomy of Lophyra (Eriolophyra)

Genus Lophyra Motschulsky 1859

Subgenus Eriolophyra Rivalier 1948

References. RiValieR 1948 (as genus); 1957 (subgenus of Lophyra); wiesneR 1992 (subgenus of Lophyra); weRneR 2000 (subgenus of Lophyra).

Type species. Cicindela barbifrons Boheman 1848 (designation by RiValieR 1957).

Diagnosis. Important diagnostic features for the genus Eriolophyra include:• Elytrapredominantly ivory-white;darkelytralgroundcolorreducedto

narrow dark linear markings or entirely absent; one species with four dark quadrate areas on each elytron;

• Frons,pronotum,antennomeres1-4, venter,and legsmetalliccoppery(occasionally metallic green);

• Frons,pronotum,scape,antennomeres3-4,venter,andlegswithdensesuberect and erect white setae;

• Pretarsalclawsenlargedandelongate,nearlyaslongastarsomeres;• Antennomeres1-4metalliccoppery-black;antennomeres5-11yellow;• Antennomere4inmaleoftenwitherectsensorysetae,whichmaytake

the form of a “penicillum” (sensu cassola 1983);• Associatedwithcoastalbeachesorsandyriverinemicrohabitatsinsub-

Saharan Africa.

Separation from allied genera and subgenera. Species belonging to two other tiger beetle taxa, Habrodera Motschulsky 1862 and Lophyra (Lophyra) Motschulsky 1859, have elytra with expanded white or pale yellow markings and are found in riverine or coastal beach habitats in sub-Saharan Africa where they might be confused with species of the subgenus Eriolophyra (weRneR 2000). Species of Lophyra (Lophyra) can be readily separated from species of subgenus Eriolophyra by the lack of dense white setae, especially on the frons, scape, pronotal disc, and legs, and by the less extensive white or yellow areas on the elytra. The few species of Lophyra (Lophyra) with expanded white or yellow elytral markings (such as L. (L.) candida Dejean 1825) can be readily distinguished from the species of subgenus Eriolophy-ra by the lack of dense white setae on the frons, scape, pronotal disc, and legs. Species of the genus Habrodera have expanded lobes on either side of the pronotal base which give the pronotum a trapezoidal shape when viewed from above. In addition, the species of Habrodera have the dark areas of the elytra distinctly impressed, whereas the dark areas of the elytra of species of subgenus Eriolophyra are not impressed.

Notes on status. Eriolophyra was described by RiValieR (1948) as a separate genus, distinguishable from Lophyra on the basis of the significant differences in vestiture and elytral markings. RiValieR (1957) subsequently placed Eriolophyra as a subgenus of Lophyra, based on similarities in the structure of the internal sac of the male genitalia. This classification was fol-lowed by subsequent authors, including wiesneR (1992) and weRneR (2000)

60 J.R. Mawdsley

in their treatments of global and African Cicindelidae, respectively. Although the exact relationships of the species currently classified within Lophyra and allied genera are unclear, the group of species now classified in subgenus Eriolophyra forms a distinct and probably monophyletic group. Other, simi-lar groups of species within the broader Lophyra complex are now recog-nized at the generic level, including Habrodera Motschulsky 1862 and Chae-todera Jeannel 1946 (wiesneR 1992, weRneR 2000). It is possible that similar treatment will eventually be proposed for subgenus Eriolophyra. However, the status of the group of species currently classified in the subgenus Lophy-ra (Lophyra) is far from clear, and Lophyra (Lophyra) may well be paraphy-letic relative to Lophyra (Eriolophyra). Any nomenclatural change would need to be based on a more comprehensive revisionary study of the Lophyra generic group that identifies clearly monophyletic lineages within the group of species now classified in the subgenus Lophyra (Lophyra).

KEy TO SPECIES

1 Elytra elongate, slender, more than 1.7 times longer than wide; associated with sandy beaches and other riverine microhabitats in Cameroon, Chad, Mali, Niger, and Senegal, possibly also the Central African Republic ......... ............................................................................ L. (E.) arnoldi (Horn 1904)

— Elytra stouter, 1.7 times longer than wide or less; Congo River basin, east-ern Africa, or southern Africa......................................................................... 2

2 Frons with two clusters of dense white setae along anterior margin; sea beaches of eastern South Africa and southern Mozambique ....................... ............................................................... L. (E.) barbifrons (Boheman 1848)

— Frons with two linear bands of dense white setae, extending from anterior margin to vertex; distribution not as above................................................... 3

3 Each elytron white with four quadrate brownish-black markings; sea beaches of northern Kenya and southern Somalia ...................................... ................................................................... L. (E.) somalia (Fairmaire 1882)

— Elytra white with narrow linear markings; riverine species ......................... 44 Elytra broad and robust in both sexes, 1.4-1.5 times longer than wide;

lateral margin of elytra strongly expanded at basal third in both sexes; smaller species, length 10-13 mm; Congo River basin................................. ............................................................................. L. (E.) albens (Horn 1895)

— Elytral form sexually dimorphic: broad and robust in female only, elytra 1.3 times longer than wide in female, 1.6 times longer than wide in male; lateral margin of elytra expanded at basal third in female only; larger spe-cies, length 14-16 mm; Mozambique, Namibia, South Africa, Tanzania, Zimbabwe .............................................................. L. (E.) alba (Horn 1894)

Lophyra (Eriolophyra) alba (Horn 1894)

References. hoRn (1894: 112; 1915: 268; 1926: 144; 1938: pl. 39 f. 8), PéRinguey (1896: 104), schildeR (1953: 556), RiValieR (1957: 329), seRRano (1987: 7), wiesneR (1992: 163), weRneR (2000: 136).


Type locality. “Africa orient. (Quilimane)” (hoRn 1894).

Diagnosis. A large tiger beetle (length 14-16 mm) with predominant-ly white elytra associated with riverine sandbars and other unconsolidated sand deposits along major river systems in southern Africa. The elytral shape is sexually dimorphic, with males having elongate elytra with acuminate api-ces (length 1.6-1.7 times longer than width) and females having very broad, trapezoidal-shaped elytra which are broadly rounded at the apex (length 1.3-1.4 times longer than width). The elytra are predominantly ivory-white, with narrow linear coppery-brown markings.

Known distribution. Mozambique: Zambezia Province, Quelimane [formerly Quilimane] (hoRn 1894). Namibia: Katima Mulilo (weRneR & wiesneR 1994, uhlig & JaegeR 1995); Kombat (weRneR & wiesneR 1994, weRneR 2000). Tanzania: Rufiji Region, Utete (weRneR 2000); “süden von Deutsch Ostafrika” (hoRn 1915). Zimbabwe (weRneR 2000).

Specimens examined. South Africa: Kruger National Park, Letaba River, 18.XI.2007 (1 male, NMNH), 19.XI.2007 (4 males, 4 females, KNPC, NMNH, SANC, TMSA). Zimbabwe: Kariba, 6.I.1973 (2 males, TMSA), 14.I.1973 (1 female, TMSA), 12.II.1973 (1 female, TMSA).

Lophyra (Eriolophyra) albens (Horn 1895)

References. hoRn (1895: 361; 1915: 268; 1926: 144; 1938: pl. 39 f. 9), BuRgeon (1937: 32), schildeR (1953: 556), RiValieR (1957: 329), wiesneR (1992: 163), weRneR (2000: 136).

Type locality. “Brazzaville (Afr. occid.)” (hoRn 1895).

Diagnosis. A moderately sized (length 10-13 mm) tiger beetle with white elytra associated with riverine sand bars in the Congo River basin. The elytra are broadly expanded in both sexes, with length 1.4-1.5 times the width. The elytra are predominantly ivory-white with narrow linear brown or black markings.

Known distribution. République Democratique du Congo: Kasai Prov-ince, Djeka (weRneR 2000); Kondue (BuRgeon 1937); Lusambo (weRneR 2000); Mobayi-Mbongo [formerly Banzyville]; Pool Malebo [formerly Stanley Pool] (BuRgeon 1937). République du Congo: Brazzaville (hoRn 1895).

Lophyra (Eriolophyra) arnoldi (Horn 1904)

References. hoRn (1904: 403; 1915: 268, pl. 16, f. 1; 1926: 144; 1931: 42; 1938: pl. 39, f. 10), hoRn et al. (1914: 426), RiValieR (1952: 213), schildeR (1953: 556), RiVa-lieR (1957:329), wiesneR (1992: 163), cassola (1998: 70), weRneR (2000: 137).

62 J.R. Mawdsley

Type locality. “Benue sup.” (hoRn 1904).

Diagnosis. A smaller (length 9-11 mm) riverine tiger beetle with pre-dominantly white elytra from West Africa. The elytra are more slender than other species of Lophyra (Eriolophyra), with length 1.8-1.9 times the width. Elytral markings are similar to those of L. (E.) alba and L. (E.) albens, with narrow greenish or coppery markings on otherwise ivory-white elytra.

Known distribution. Cameroon (locality not specified) (weRneR 2000). Chad: Bakare (hoRn 1931). Mali: Koulikoro (hoRn et al. 1914). Nigeria: Lau (weRneR 2000). Senegal (locality not specified) (weRneR 2000). Sierra Leone: yoka (weRneR 2000). “Benue sup[erior]” (hoRn 1904). “Oberen Senegal-Niger” (hoRn 1915). weRneR (2000) also recorded this species from the Central African Republic, with a question mark “?” as to the validity of the record. According to F. cassola (pers. comm.), the locality in Sierra Leone also needs to be checked.

Specimen examined. Mali: Koulikoro (1 male, NMNH).

Lophyra (Eriolophyra) barbifrons (Boheman 1848)

References. BoheMan (1848: 12-13), hoRn (1915: 267; 1926: 144; 1938: pl. 39, f. 5), schildeR (1953: 556), RiValieR (1957: 328-329), cassola (1975: 212), wiesneR (1992: 163), weRneR (2000: 134), BRzoska & cassola (2005: 73).

Type locality. “in Caffraria interiore” (BoheMan 1848).

Synonyms. Cicindela chrysographa PéRinguey (1896: 105), not deJean (1831: 254). Cicindela chrysographa var. marqueza PéRinguey (1896: 105).

Diagnosis. A smaller (length 9-11 mm) tiger beetle with white or pre-dominantly white elytra, occurring on coastal sea beaches in South Africa and Mozambique. The elytra are either entirely white or have a series of small vittate and/or maculate coppery-brown markings. When present, the dark markings of the elytra are not impressed as in species of Habrodera. The presence of dense white pubescence on the pronotum, antennal scape, antennomeres 3-4, and legs will readily separate it from sympatric species of Lophyra (Lophyra) and Habrodera.

Known distribution. Mozambique: Inhambane Province, Praia de Quis-sico; Joao Belo Province, Praia de Chidenguela (cassola 1975); Maputo Province, Maputo Bay [formerly Delagoa Bay] (hoRn 1915). South Africa: Eastern Cape Province: Port Elizabeth (hoRn 1915); KwaZulu-Natal Prov-ince: Durban (PéRinguey 1896); mouth of Tugela River (cassola 1975, weR-neR 2000, BRzoska & cassola 2005); Manzengwenya (weRneR 2000); Sipon-go [formerly Tsipongo] (PéRinguey 1896).

Specimens examined. Mozambique: Gaza Province: Chonguene, XII.1972 (1 male, NMNH). Maputo Province: Beach E of Rikatla, XII (4 males, 1 female, NMNH); Inhaca,


I.1973 (1 male, TMSA, 2 females, NMNH, 1 female, SANC); Lourenco Marques, XII.1908 (1 male, 1 female, TMSA), II.1957 (1 male, 1 female, NMNH). South Africa: KwaZu-lu-Natal Province: Cosy Bay mouth, sandy sea-shore, 5.XII.1992 (10 males, 2 females, TMSA); Doonside, 10.XII.1937 (1 female, SANC); Durban, 1896 (1 female SANC), III (3 males, 1 female, NMNH), 27.III.1920 (1 male, TMSA); Gingindhlovu, 30.XII.1977 (1 female, SANC); Illovo Beach, 25.I.1984 (1 male, 1 female, TMSA); Isipingo, XII.1895 (1 female, TMSA); Manzengwenya, 30.XII.1979 (1 female, SANC); Mtuzini, II.1953 (13 males, 8 females, TMSA); N. Lemare, II.1953 (1 male, TMSA); Sordwana Bay, I.1939 (1 male, 1 female, SANC), I.1949 (2 males, 1 female, SANC); Tongaat, 1909 (6 males, TMSA); Turton, 30.I.1971 (1 male, 3 females, TMSA), 15.II.1971 (3 males, 4 females, TMSA); Umdoni Park, 18.III.1971 (1 male, TMSA); Umgeni (1 male, TMSA). Western Cape Province: Jackalswater, between Springbok and Uranoep, II.1953 (1 female, TMSA).

Discussion of synonymies. PéRinguey (1896) treated the species now known as L. (E.) barbifrons under the name Cicindela chrysographa Dejean 1831. However, the taxon originally described by deJean (1831) as Cicinde-la chrysographa is not conspecific with L. (E.) barbifrons and is in fact now considered a doubtful subspecies or synonym of Habrodera capensis (Linnae-us 1764) (wiesneR 1992, weRneR 2000). Cicindela chrysographa Péringuey 1896 must therefore be considered a primary junior homonym of Cicindela chrysographa Dejean, as well as a synonym of Cicindela barbifrons Boheman.

PéRinguey (1896) also described a form Cicindela chrysographa var. marqueza from Delagoa Bay, Mozambique. This form differed morphologi-cally from the nominate form in having its elytra entirely white, without any dark ground color present (PéRinguey 1896). Cicindela chrysographa var. mar-queza was subsequently placed as a synonym of Cicindela barbifrons Bohe-man by hoRn (1915, 1926). It was not treated as a valid taxon by RiValieR in his revision of Lophyra (RiValieR 1948) or his more comprehensive treatment of African cicindelines (RiValieR 1957). The name was treated by wiesneR (1992) as a subspecific epithet and it was considered a doubtful subspecies of L. (E.) barbifrons by weRneR (2000).

As noted by PéRinguey (1896), adults which have the elytra entirely white are found in populations of L. (E.) barbifrons in both Mozambique and Kwa-Zulu-Natal. In the NMNH collection, I examined a small series from Durban, KwaZulu-Natal, which contained one entirely white specimen, two specimens with the dark elytral markings as described by BoheMan (1848), and two speci-mens with reduced dark markings. I also examined an entirely white specimen from Rikatla, Mozambique, as well as two specimens from Delagoa Bay which had the dark elytral markings as described by BoheMan (1848). Although mor-phologically distinct, the form with entirely white elytra is not localized to a specific geographic area and is therefore not worthy of recognition on a sub-specific level. The name Cicindela chrysographa var. marqueza Péringuey should therefore be treated as a synonym of L. (E.) barbifrons (Boheman).

Lophyra (Eriolophyra) somalia (Fairmaire 1882)

References. FaiRMaiRe (1882: 5-6), Fleutiaux (1892: 81), hoRn (1915: 268; 1926: 144; 1938: pl. 39, f. 7), schildeR (1953: 556), RiValieR (1957: 329), naViaux (1980: 220), cassola & Miskell (1990: 194); wiesneR (1992: 163); weRneR (2000: 135).

64 J.R. Mawdsley

Type locality. Not specified in the original description, but the introduc-tion to FaiRMaiRe (1882: 1) indicates that it should be “le pays des Çomalis.”

Diagnosis. A small (length 10-12 mm) white tiger beetle of coastal sea beaches in northern Kenya and southern Somalia. Each elytron has four quadrate dark brownish-black maculae which are subequal in size and arranged in a line on the elytral disc. The maculae may be more or less dis-tinct in individual specimens.

Known distribution. Kenya: Malindi (weRneR 2000). Somalia: Benadir, Gesira; Gesira Beach, 20 kilometers SW of Mogadishu (cassola & Miskell 1990, weRneR 2000).

Specimens examined. Somalia: Gesira, 20 km SW Mogadishu, 29.IV.1987 (1 male, 1 female, SANC).

ECOLOGy

Species of the subgenus Eriolophyra are inhabitants of riverine or coast-al sand beaches in sub-Saharan Africa. weRneR (2000) provided an overview of the biology of the species of this genus. Two species, L. (E.) barbifrons and L. (E.) somalia, are associated with coastal sand beaches in southern and eastern Africa, respectively (PéRinguey 1896, cassola 1975, weRneR 2000). The remaining species are associated with sandy riverine areas in southern Africa (L. (E.) alba), central Africa (L. (E.) albens), and west and north-central Africa (L. (E.) arnoldi) (weRneR 2000). Lophyra (Eriolophyra) alba has also been collected at lights at night (weRneR & wiesneR 1994).

cassola (1975) presented information about the ecology of L. (E.) bar-bifrons at sites in Natal and Mozambique. He characterized this species as a “halophilous, seashore-inhabiting species” and noted that “large ocean beaches” were its “typical habitat” (cassola 1975). Earlier, PéRinguey (1896) had noted that this species “do[es] not occur far from the seashore.” BRzos-ka & cassola (2005) reported recent collections of this species at the mouth of the Tugela River, where adults were collected “on the sea-facing slope of the coastal sand dunes”.

cassola & Miskell (1990), naViaux (1980), and weRneR (2000) present-ed information about the ecology of L. (E.) somalia, which appears to have similar habits to L. (E.) barbifrons, except that it is found on sea beaches in Somalia and northern Kenya (“sur les grande plage de sable” according to naViaux 1980: 220).

In November, 2007, I studied the ecology of L. (E.) alba at sites along the Letaba River in Kruger National Park, South Africa. A single male of this species was collected at 2 PM on 18 November, 2007, during routine surveys for riverine tiger beetle species (described by Mawdsley & sithole 2008). This individual was collected away from the main river channel on a dry, sandy riverbank overlooking the riverine floodplain. This section of


riverbank was actively eroding and the single individual of L. (E.) alba was collected in an area of dry, eroding, unconsolidated sand. Other tiger beetle species present at this site included Lophyra (Lophyra) differens (Horn 1892) and Lophyra (Lophyra) neglecta intermediola (Horn 1921).

On 19 November, 2007, I found a large population of L. (E.) alba on a very large sand dune located in the floodplain of the Letaba River (Fig. 1). The dune was triangular in shape, approximately 30 m wide, 50 m long, and 5 m high, with a broad, flat top and steep sides. The dune was composed entirely of fine dry unconsolidated yellow sand, and lacked associated veg-etation. Over 50 adult L. (E.) alba were observed on or near this dune. Activ-ity of the adults of L. (E.) alba was confined to the top surface and sides of the sand dune and to a small wash or gully that cut through the dune and led to the main channel of the Letaba River approximately 100 m away.

Foraging, reproductive, and thermoregulatory behaviors of L. (E.) alba were observed at this site. In general, these behaviors are similar to those described for other tiger beetles (knisley & schultz 1997). Foraging behav-ior consists of short (1-3 m) runs, after which the beetle pauses briefly, and then runs again. This behavior is repeated continuously. Adult L. (E.) alba

Fig. 1. — View of the Letaba River in Kruger National Park, photographed from the south end of the high water bridge on the H1-6 main tourist road northwest of Letaba Rest Camp. The white arrow indicates the most productive collecting site for Lophyra (Eriolo-phyra) alba, while the black arrow indicates a collecting area for Chaetodera regalis.

66 J.R. Mawdsley

were observed feeding on small ants and other small arthropods: as with other cicindelids, prey is captured using the mandibles, masticated using the mandibular bases, and held by the maxillary and labial palpi against the buccal cavity for feeding. Beetles along the base of the dune and along the course of the small wash or gully exhibited amplexus, mating, and oviposi-tion behaviors. All oviposition sites were in loose dry unconsolidated sand. Adult beetles exhibited the full range of thermoregulatory behaviors reported in tiger beetles (knisley & schultz 1997), including basking, stilting, shut-tling, and burrowing in loose sand.

The ivory-white dorsal coloration of L. (E.) alba is strongly cryptic on the loose yellow sand surface, and the adult beetles are not easily detect-ed except when disturbed. Adult L. (E.) alba have several defensive behav-iors. If approached closely, their first response is to stop moving and remain motionless for a short period. This behavior renders them quite inconspicu-ous against the sand. If the disturbance continues, the beetles will fly. Escape flights are short and straight, with most between 5-10 m. Upon landing, the beetles often remain motionless for a short period of time, after which they resume normal foraging behaviors. If captured, the beetles will attempt to bite and will also regurgitate a black liquid from the digestive tract. I did not detect a chemical odor when collecting the adult beetles.

Three other species of tiger beetles were observed in the small gully near the dune with L. (E.) alba: Lophyra (Lophyra) differens (2 individuals), Lophyra (Lophyra) neglecta intermediola (1 individual), and Chaetodera regalis (Dejean 1831) (1 individual). Continued investigation of this general area led to the discovery of large populations of tiger beetles on a very large riverine sandbar which was located approximately 100 m from the dune. Tiger bee-tles observed on this sandbar included Chaetodera regalis (over 100 individu-als) and Calomera fimbriata imperatrix (Srnka 1891) (2 individuals).

Potential predators or parasites of L. (E.) alba were not observed at this site, although large robber flies (Diptera Asilidae) and dragonflies (Odonata Anisoptera) capable of capturing adults of L. (E.) alba were observed at the large riverine sandbar nearby.

Voucher specimens from these observations are deposited in the fol-lowing collections: KNPC, NMNH, SANC, TMSA (see Acknowledgments for explanation of abbreviations).

PHyLOGENETIC RELATIONSHIPS

The phylogenetic relationships of Lophyra and several closely allied genera and subgenera (the genera Chaetodera and Habrodera, and the sub-genera Lophyra (Eriolophyra) and Lophyra (Stenolophyra Rivalier 1957)) have not been studied in detail. The following 10 morphological characters have been used by previous authors (hoRn 1915; RiValieR 1948, 1957; weRneR 2000) to diagnose and separate species within the subgenus Eriolophyra.

1. Clypeus with white setae laterally, glabrous medially (0) / clypeus setose across entire width (1).


2. Scape lacking white setae (0) / scape with white setae (1).3. Antennomeres 3-4 lacking white setae (0) / antennomeres 3-4 with

white setae (1).4. Frons with dense white setae limited to clusters along the anterior

margin (0) / frons with lines of white setae extending onto vertex of head (1)5. Pronotal disc glabrous (0) / pronotal disc with dense white setae (1).6. Elytra not broadly expanded in female, length 1.5 times width or

greater (0) / elytra broadly expanded in female, length 1.3-1.4 times the width (1).

7. Elytral epipleurae in female straight or broadly and uniformly rounded, lacking lobe-like lateral expansion at basal third (0) / with lobe-like lateral expansion at basal third (1).

8. Elytral shape subequal in both sexes (0) / sexually dimorphic, broad-er in female than in male (1).

9. Dark elytral ground color extensive, covering 50% or more of elytral surface (0) / dark elytral ground color reduced, with more than 50% of ely-tral surface white (1).

10. Abdominal sternites with white setae laterally (0) / with white setae medially as well as laterally (1).

The distribution of these characters within the subgenus Eriolophyra is shown in Table 1. I have included a species of Lophyra (Lophyra), L. (L.) neglecta Dejean 1825, as an outgroup taxon for purposes of cladistic analysis. Lophyra (Lophyra) neglecta is a fairly generalized species of Lophyra (Lophy-ra) and is widespread throughout sub-Saharan Africa (wiesneR 1992, weR-neR 2000). Character states present within L. (L.) neglecta were provisionally interpreted as plesiomorphic (0) while the character states absent in L. (L.) neglecta but present within various species of Eriolophyra were provisionally interpreted as apomorphic (1).

The matrix in Table 1 yields a single most parsimonious cladogram (Fig. 2) under standard maximum parsimony analysis using the algorithm of FaRRis (1970). This cladogram shows four clear synapomorphies for this

Table 1.

Distribution of character states in Lophyra, subgenus Eriolophyra.

SpeciesCharacter number

1 2 3 4 5 6 7 8 9 10

L. (L.) neglecta 0 0 0 0 0 0 0 0 0 0

L. (E.) alba 1 1 1 1 1 1 1 1 1 1

L. (E.) albens 1 1 1 1 1 1 1 0 1 1

L. (E.) arnoldi 1 1 1 1 1 0 0 0 1 1

L. (E.) barbifrons 0 1 1 0 1 0 0 0 1 0

L. (E.) somalia 1 1 1 1 1 0 0 0 1 0

68 J.R. Mawdsley

subgenus (the derived states of characters 2, 3, 5, and 9) relative to L. (L.) neglecta. Lophyra (Eriolophyra) barbifrons is the sister-species to the four other species of Eriolophyra, which are united by two synapomorphies (the derived states of characters 1 and 4). Within this group of four species, Lophyra (Eriolophyra) somalia is the sister-species to the three other spe-cies, which are united in a clade by one synapomorphy (the derived state of character 10). These three species also share a similar elytral color pat-tern (weRneR 2000). Finally, L. (E.) alba and L. (E.) albens are clearly sister-species, united by two synapomorphies (the derived states of characters 6 and 7). The derived state of character 8 is an autapomorphy of L. (E.) alba.

ACKNOWLEDGMENTS

I would like to thank Hendrik Sithole and Freek Venter of Kruger National Park, South Africa, for their generous support and formal approval of my studies of tiger beetles. Field assistance was provided by Velly Ndlovu and Johane Baloyi of Kru-ger National Park, as well as by Alice Mawdsley and Ralph Mawdsley of Cleveland State University. For permission to examine specimens in their care, I thank Terry Erwin, National Museum of Natural History, Smithsonian Institution, Washington, DC (NMNH), Guin Zambatis, Kruger National Park, Skukuza, South Africa (KNPC), Beth Grobbelaar and Riaan Stals, South African National Collection of Insects (SANC), and James Harrison and Ruth Müller, Transvaal Museum, Pretoria, South Africa (TMSA). For helpful comments on the manuscript, I thank Fabio Cassola and Jürgen Wiesner.

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taxonomy, ecology, and phylogeny of species of lophyra motschulsky 1859, subgenus

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