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 subgenus is reviewed and Lophyra (Eriolophyra) barbifrons marqueza (Péringuey 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 associated with sandy riverine areas in southern, central, and west-central Africa, 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 defensive 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 characteristics. Morphological characters suggest that this group forms a monophyletic clade within the broader complex of species allied to Lophyra.
key words:
Eriolophyra, Cicindelidae, ecology, phylogeny, morphology,
distribution.
Introduction
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Genus Lophyra Motschulsky 1859 Subgenus Eriolophyra Rivalier 1948
Key to Species
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Lophyra (Eriolophyra) alba (Horn 1894)
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Lophyra (Eriolophyra) albens (Horn 1895)
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Lophyra (Eriolophyra) arnoldi (Horn 1904) .
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Lophyra (Eriolophyra) barbifrons (Boheman 1848)
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Lophyra (Eriolophyra) somalia (Fairmaire 1882)
Ecology
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Phylogenetic relationships
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Acknowledgments
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References
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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 sensitive to anthropogenic disturbances (Knisley & Schultz 1997). Human recreational 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 riverine 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 prepared a key to species and a provisional cladistic analysis of the species of
this subgenus.
Taxonomy of Lophyra (Eriolophyra)
59
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:
•
Elytra predominantly ivory-white; dark elytral ground color reduced to
narrow dark linear markings or entirely absent; one species with four
dark quadrate areas on each elytron;
•
Frons, pronotum, antennomeres 1-4, venter, and legs metallic coppery
(occasionally metallic green);
•
Frons, pronotum, scape, antennomeres 3-4, venter, and legs with dense
suberect and erect white setae;
•
Pretarsal claws enlarged and elongate, nearly as long as tarsomeres;
•
Antennomeres 1-4 metallic coppery-black; antennomeres 5-11 yellow;
•
Antennomere 4 in male often with erect sensory setae, which may take
the form of a “penicillum” (sensu Cassola 1983);
•
Associated with coastal beaches or sandy riverine microhabitats in subSaharan 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 Eriolophyra 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 followed 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, similar groups of species within the broader Lophyra complex are now recognized at the generic level, including Habrodera Motschulsky 1862 and Chaetodera 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 Lophyra (Lophyra) is far from clear, and Lophyra (Lophyra) may well be paraphyletic 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
—
2
—
3
—
4
—
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, eastern Africa, or southern Africa.........................................................................
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...................................................
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..........................
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 species, length 14-16 mm; Mozambique, Namibia, South Africa, Tanzania,
Zimbabwe............................................................... L. (E.) alba (Horn 1894)
2
3
4
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).
Taxonomy of Lophyra (Eriolophyra)
61
Type locality. “Africa orient. (Quilimane)” (Horn 1894).
Diagnosis. A large tiger beetle (length 14-16 mm) with predominantly 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 apices (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 1.31.4 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 Province, 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), Rivalier (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 predominantly 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 predominantly 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 Quissico; 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 Province: Durban (Péringuey 1896); mouth of Tugela River (Cassola 1975, Werner 2000, Brzoska & Cassola 2005); Manzengwenya (Werner 2000); Sipongo [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,
Taxonomy of Lophyra (Eriolophyra)
63
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: KwaZulu-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 Cicindela chrysographa is not conspecific with L. (E.) barbifrons and is in fact now
considered a doubtful subspecies or synonym of Habrodera capensis (Linnaeus 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 morphologically from the nominate form in having its elytra entirely white, without any
dark ground color present (Péringuey 1896). Cicindela chrysographa var. marqueza was subsequently placed as a synonym of Cicindela barbifrons Boheman 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 KwaZulu-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 specimens 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 morphologically distinct, the form with entirely white elytra is not localized to a
specific geographic area and is therefore not worthy of recognition on a subspecific 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 introduction 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 distinct 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 coastal 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.) barbifrons 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.” Brzoska & 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) presented 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
Taxonomy of Lophyra (Eriolophyra)
65
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 vegetation. Over 50 adult L. (E.) alba were observed on or near this dune. Activity 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 behavior 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 (Eriolophyra) 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 oviposition 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, shuttling, 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 detected except when disturbed. Adult L. (E.) alba have several defensive behaviors. If approached closely, their first response is to stop moving and remain
motionless for a short period. This behavior renders them quite inconspicuous 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 beetles observed on this sandbar included Chaetodera regalis (over 100 individuals) 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 following 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 subgenera 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).
Taxonomy of Lophyra (Eriolophyra)
67
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, broader 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 elytral 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 (Lophyra) and is widespread throughout sub-Saharan Africa (Wiesner 1992, Werner 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.
Species
Character number
1 2 3 4 5 6 7 8 9 10
L. (L.) neglecta
0000000000
L. (E.) alba
1111111111
L. (E.) albens
1111111011
L. (E.) arnoldi
1111100011
L. (E.) barbifrons
0110100010
L. (E.) somalia
1111100010
68
J.R. Mawdsley
Fig. 2. — Most parsimonious cladogram from cladistic analysis of
the matrix in Table 1, with 10 steps.
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 species, which are united in a clade by one synapomorphy (the derived state
of character 10). These three species also share a similar elytral color pattern (Werner 2000). Finally, L. (E.) alba and L. (E.) albens are clearly sisterspecies, 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 Kruger 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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