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Taxonomic revision of Afrotropical Laccophilus Leach, 1815 (Coleoptera, Dytiscidae)
expand article infoOlof Biström, Anders N. Nilsson§, Johannes Bergsten|
‡ Finnish Museum of Natural History, University of Helsinki, Helsinki, Finland
§ Umeå University, Umeå, Sweden
| Swedish Museum of Natural History, Stockholm, Sweden
Open Access

Abstract

The African species of the genus Laccophilus Leach, 1815, are revised, on the basis of study of adult specimens. In all, 105 species are now recognized. A phenetic character-analysis was undertaken, which resulted in a split of the genus into 17 species groups. Diagnoses and a description of each species are given together with keys for identification of species groups and species. We also provide habitus photos, illustration of male genitalia and distribution maps for all species. New species are described as follows: L. grossus sp. n. (Angola, Namibia), L. rocchii sp. n. (Tanzania, Namibia, Botswana, Mozambique), L. ferrugo sp. n. (Mozambique), L. furthi sp. n. (Madagascar), L. isamberti sp. n. (Madagascar), L. inobservatus sp. n. (Gambia, Senegal, Mali, Niger, Sudan, Chad, Ethiopia, Burkina Faso, Ivory Coast, Ghana, Nigeria, Cameroon, Zaire and Asia: Yemen), L. cryptos sp. n. (Zaire, Mozambique, Zimbabwe, Namibia, Botswana, South Africa), L. enigmaticus sp. n. (Nigeria, Sudan), L. bellus sp. n. (Benin, Nigeria), L. guentheri sp. n. (Guinea, Ghana), L. guineensis sp. n. (Guinea), L. decorosus sp. n. (Uganda), L. empheres sp. n. (Kenya), L. inconstans sp. n. (Guinea, Ivory Coast, Ghana, Nigeria, Cameroon), L. brancuccii sp. n. (Central African Republic), L. incomptus sp. n. (Cameroon), L. australis sp. n. (Tanzania, South Africa), L. minimus sp. n. (Namibia), L. eboris sp. n. (Ivory Coast), L. insularum sp. n. (Madagascar), L. occidentalis sp. n. (Gambia, Senegal, Mali, Guinea, Sierra Leone, Ivory Coast, Ghana, Benin, Nigeria, Central African Republic, Zaire) and L. transversovittatus sp. n. (Madagascar). L. restrictus Sharp, 1882, is restored as good species; not junior synonym of L. pictipennis Sharp, 1882. New synonyms are established as follows: L. continentalis Gschwendtner, 1935 = L. perplexus Omer-Cooper, 1970, syn. n., L. taeniolatus Régimbart, 1889 = L. congener Omer-Cooper, 1957, syn. n., L. adspersus Boheman, 1848 = L. vitshumbii Guignot, 1959, syn. n. = L. adspersus nigeriensis Omer-Cooper, 1970, syn. n. = L. adspersus sudanensis Omer-Cooper, 1970, syn. n., L. modestus Régimbart, 1895 = L. espanyoli Hernando, 1990, syn. n., L. flaveolus Régimbart, 1906 = L. pampinatus Guignot, 1941, syn. n., L. trilineola Régimbart, 1889 = L. simulator Omer-Cooper, 1958, syn. n., L. mediocris Guignot, 1952 = L. meii Rocchi, 2000, syn. n., L. epinephes Guignot, 1955 = L. castaneus Guignot, 1956, syn. n., L. saegeri Guignot, 1958 = L. comoensis Pederzani & Reintjes, 2002, syn. n., L. restrictus Sharp, 1882 = L. evanescens Régimbart, 1895, syn. n., L. incrassatus Gschwendtner, 1933 = L. virgatus Guignot, 1953, syn. n., L. cyclopis Sharp, 1882 = L. shephardi Omer-Cooper, 1965, syn. n., L. burgeoni Gschwendtner, 1930 = L. wittei Guignot, 1952, syn. n., L. secundus Régimbart, 1895 = L. torquatus Guignot, 1956, syn. n., L. desintegratus Régimbart, 1895 = L. sanguinosus Régimbart, 1895, syn. n. and L. flavopictus Régimbart, 1889 = L. bergeri Guignot, 1953, syn. n. = L. segmentatus Omer-Cooper, 1957, syn. n. Lectotypes are designated for the following taxa: L. productus Régimbart, 1906, L. ruficollis Zimmermann, 1919, L. sordidus Sharp, 1882, L. alluaudi Régimbart, 1899, L. pictipennis Sharp, 1882, L. wehnckei Sharp, 1882, L. continentalis Gschwendtner, 1935, L. simplicistriatus Gschwendtner, 1932, L. complicatus Sharp, 1882, L. rivulosus Klug, 1833, L. ampliatus Régimbart, 1895, L. pilitarsis Régimbart, 1906, L. adspersus Boheman, 1848, L. livens Régimbart, 1895, L. modestus Régimbart, 1895, L. nodieri Régimbart, 1895, L. flaveolus Régimbart, 1906, L. pallescens Régimbart, 1903, L. restrictus Sharp, 1882, L. vermiculosus Gerstaecker, 1867, L. mocquerysi Régimbart, 1895, L. bizonatus Régimbart, 1895, L. tschoffeni Régimbart, 1895, L. persimilis Régimbart, 1895, L. poecilus Klug, 1834, L. lateralis Sharp, 1882, L. lateralis var. polygrammus Régimbart, 1903, L. cyclopis Sharp, 1882, L. shephardi Omer-Cooper, 1965, L. conjunctus Guignot, 1950, L. grammicus Sharp, 1882, L. flavoscriptus Régimbart, 1895, L. flavosignatus Régimbart, 1895, L. brevicollis Sharp, 1882, L. secundus Régimbart, L. desintegratus Régimbart, 1895, L. gutticollis Régimbart, 1895, L. luctuosus Sharp, 1882 and L. inornatus Zimmermann, 1926. Laccophilus remex Guignot, 1952, comprises a species complex with uncertain taxonomic delimitation; the complex includes L. concisus Guignot, 1953, L. turneri Omer-Cooper, 1957 and L. praeteritus Omer-Cooper, 1957, as tentative synonyms of L. remex Guignot, 1952.

Keywords

Coleoptera, Dytiscidae, Laccophilus, Africa, Madagascar, taxonomy, revision, description, new species

Introduction

The genus Laccophilus Leach, 1815 is by far the most species rich genus among Laccophilinae. The most recent world catalogue (Nilsson 2015) lists 263 valid species out of which a considerable number occur in Africa including Madagascar (94 species prior to this publication, 105 species after). The high species number is not surprising due to the extensive distribution on the Globe of the genus. Laccophilus species are found on all continents except for Antarctica. The last taxonomic work, which treated the whole genus on global level, was Sharp (1882). After that a number of works have been published but all of them have focused on a limited geographical area. The East-Palearctic, Oriental and Australian faunas containing 59 species have been revised by Brancucci (1983). Zimmerman (1970) revised the Laccophilus in North America, recognizing in all 27 species. The African fauna including Madagascar has been revised by Régimbart (1895) and Guignot (1959a). An additional larger work on African Laccophilus was published by Omer-Cooper (1965) who treated the fauna in southern Africa. Besides these, in their scope larger treatments, there are numerous taxonomic papers, giving valuable information on regional and country level. One of the most recent being Hájek and Brancucci (2015), dealing with a species-rich group of Laccophilus in South East Asia. A comprehensive, up-to-date revision on the whole African continent is, however, still lacking. The main aim of this work is to fill this gap. We are perfectly well aware of the fact that this work is not complete and that there are still many taxonomic questions to be solved in Africa. Anyway, the present revision, hopefully fulfils its function as a solid base for future studies.

Material and methods

The study material, numbering almost 11000 adult specimens, comes from a number of institutions, museums and private collections. These are referred in the text by the following abbreviations:

AMGS Albany Museum, Grahamstown, South Africa (Ferdinand de Moor and Helen James)

BMNH The Natural History Museum, London, UK (Christine S. Taylor)

CAS California Academy of Sciences, San Francisco, USA

CCT Collection Clive Turner, Plymouth, UK

CFP Collection Fernando Pederzani, Ravenna, Italy

CGC Collection Gilbert Challet, Celonova, Foothill Ranch, California, USA

CGF Collection Garth Foster, Ayr, Scotland (UK)

CGW Collection Günther Wewalka, Vienna, Austria

CIR Collection Ignacio Ribera, Barcelona, Spain

CSR Collection Saverio Rocchi, Firenze, Italy

HNHM Hungarian Natural History Museum, Budapest, Hungary

IRSNB Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgique (Martina Peeters and Patrick Grootaert)

MHNG Museum d’Histoire Naturelle, Geneva, Switzerland (Giulio Cuccodoro)

MNHN Museum National d’Histoire Naturelle, Paris, France (Antoine Mantilleri)

MRAC Musée Royal de l’Afrique Centrale, Tervuren, Belgique (Marc De Meyer)

MSNM Museo Civico di Storia Naturale, Milan, Italy (Fabricio Rigato)

MZBS Museo de Zoologia, Barcelona, Spain (Gloria Masó, via Ignacio Ribera)

MZH Museum Zoologicum (Finnish Museum of Natural History), Helsinki, Finland

MZLU Zoological Museum, Lund, Sweden (Roy Danielsson)

MZUL Museo di Zoologia dell’Università, La Sapienza, Roma, Italy (Maurizio Mei)

NHMB Naturhistorisches Museum, Basel, Switzerland (the late Michel Brancucci and Matthias Borer)

NHRS Naturhistoriska Riksmuseet, Stockholm, Sweden (the late Bert Wiklund)

NMNW National Museum, Windhoek, Namibia [Comment: Specimens attributed to this museum are temporarily in Berlin, ZMHB]

NMPC National Museum (Natural History), Prague, Czech Republic (Jirí Hájek)

NMW Naturhistorisches Museum, Vienna, Austria (Manfred Jäch)

OLML Oberösterreichisches Landesmuseum, Linz, Austria (Fritz Gusenleitner and Claudia Reitstätter)

RMNH Nationaal Natuurhistorische Museum (Naturalis), Leiden, the Netherlands (A. van Assen)

SAMC Iziko Museum of Cape Town, South Africa (Margie Cochrane and Dawn Larsen)

SMNS Staatliches Museum für Naturkunde, Stuttgart, Germany

TAU Tel Aviv University, Israel (Netta Dorchin and Leonid Friedman)

TMSA Transvaal Museum, Pretoria, South Africa (Ruth Müller)

USNM National Museum of Natural History, Washington D.C., USA (David G. Furth)

ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany (Manfred Uhlig)

ZMSC Zoologische Staatssammlung, München, Germany (Martin Baehr)

ZMUC Zoological Museum, Copenhagen, Denmark (Alexey Solodovnikov)

Names in brackets in the list above refer to colleagues responsible for arrangements of loans.

The material studied is given for each species in a separate section, where relevant countries are arranged from west to east and north to south.

Species geographical records were provided with decimal degree latitude and longitude coordinates whenever possible (Suppl. material 1).

Preparation technique, drawings, photographs and mapping

The study material consists both of dry, pinned specimens as well as specimens preserved in ethanol. For study of the genitalia dry specimens were treated as follows. Examined specimen was softened in hot water for some minutes. After that the apical ventrite was detached and the genitalia were released from surrounding, hardened tissue. Often the hardened tissue needed to be treated in a heater-device for about 10 minutes in 10% KOH solution. The genitalia were than washed in water baths and prepared for illustrations. Drawings were made using a Wild M 11 microscope provided with a camera lucida. The cleaned male genitalia were put in a drop of glycerine on a slide for the illustration-process. After this the genitalia and the detached apical ventrite were mounted on a card together with the specimen. Wet specimens were treated in same manner as dry specimens. If the wet specimen studied was still preserved as wet, the genitalia were placed in a microvial together with the specimen. Penis and paramere were illustrated either detached in two pieces or undetached, together in one piece, depending on what was accessible.

Illustrations of external body-parts were made using a Wild M 5 -microscope provided with a camera lucida.

Habitus photos were taken with a Canon EOS 5D Mark II DSLR camera with a supermacro MPE-65 mm f/2.8 1–5× lens and mounted on a stackshot motorized rail from Cognisys. For light the macro twin-head flash MT-24EX was used with homemade light diffusors both directly on the flash heads and as a cylinder around the specimen. Extended focus was achieved with focusstacking technique with between 6 and 20 photos taken for each specimen. The motorized Stackshot rail was controlled via the software Zerene Stacker (Version 1.04 Build T201402072140, Zerene Systems, LLC). Focus range was assessed with live view images delivered by EOS Utility (Version 2.14.10, Canon INC). All species were photographed with the MPE-65 lens set at 3:1 magnification. The PMax algorithm in Zerene Stacker under default settings was used to create an extended focus image from the original stack of photos. Postediting of images was done in Adobe Photoshop CS5 Extended (Version 12.1 x64, Adobe Systems Incorporated) where also a scale was added using a calibration file.

Species geographical records were provided with decimal degree latitude and longitude coordinates whenever possible. Named geographical units were identified on the Microsoft Encarta Premium Suite (version 2003) world map when present. Other sources for the geographical position of named units include published expedition reports, vintage atlases and google search. Records providing only general information like names of regions and countries were not provided with coordinates. Other problematic cases include redundant names without discriminating information. For each species a list of records expressed as decimal degree latitude and longitude coordinates were added to a basemap layer provided by ESRI using ArcGIS 10 and WGS 1984 projection.

The species groups used in this revision are based on the phenetic analysis presented on p. 12. Within the species groups the species are ordered on the basis of morphological similarity.

Systematics

Laccophilus Leach, 1815

Type species

(by monotypy). Dytiscus minutus Linnaeus, 1758.

Laccophilus Leach, 1815: 84 (673 alternative page number) (original description); Aubé 1838: 415 (description, global distribution); Sharp 1882: 286, 287 (description, faunistics, species list, faunistics, discussion, species group delimitation); Kolbe 1883: 386, 401 (faunistics, discussion); Peschet 1917: 23 (discussion, key); Zimmermann 1919: 119 (description); Zimmermann 1920a: 16 (catalogue, faunistics); Bertrand 1928a: 184 (description, faunistics); Bertrand 1928c: 364 (larva description); Guignot 1937: 137, 138 (discussion, description, key to genera; type species of genus incorrectly given as L. hyalinus De Geer); Guignot 1946a: 116 (type species Dytiscus minutus Linnaeus); Guignot 1946c: 260, 261, 315 (description, key to genus and species groups, discussion); Guignot 1948: 15 (description, key to genera); Bertrand 1948: 12 (description larva, faunistics); Bertrand 1951: 114 (discussion, faunistics); Bertrand 1954: 284, 288, 289 (discussion larva, description, faunistics); Guignot 1955a: 37 (biology); Omer-Cooper 1956: 21, 23 (faunistics, biology); Omer-Cooper 1957: 8, 11, 90 (key, description); Omer-Cooper 1958b: 36 (key, subgroups, description); Guignot 1959a: 530 (description, discussion, faunistics, 11 species groups distinguished and keyed); Omer-Cooper 1962: 294, 295 (faunistics); Bertrand 1963: 402, 411, 448 (juvenile discussion); Omer-Cooper 1965: 61, 65 (description, discussion, faunistics, biology); Bertrand 1970: 18, 38 (description, larva); 1971: 252 (larva, faunistics); Bertrand and Legros 1971: 244 (faunistics, biology); Forge 1981: 501 (description, faunistics); Brancucci 1983: 251, 253 (description, key); Brancucci 1983b: 241–426 (description, faunistics, discussion, taxonomic revision Oriental, East-Palearctic and Australian species); Pederzani 1988: 107 (faunistics); Nilsson et al. 1989: 299 (list, type species by monotypy, Dytiscus minutus Linnaeus, 1758); Nilsson and Persson 1993: 79 (discussion, faunistics, discussion); Pederzani 1995: 43, 73 (cosmpolitan genus, key, list); Nilsson et al. 1995: 505 (faunistics); Balke et al. 1997: 295–320 (review New Guinea species, melanism, discussion); Nilsson and Roughley 1997: 4 (list); Alarie et al. 2000: 121–164 (Laccophilinae phylogeny discussion, based on larval morphology); Nilsson 2003: 76 (type species: Dytiscus minutus Linnaeus); Reintjes 2004: 66 (faunistics list, all continents); Bilardo and Rocchi 2006: 130, 133 (faunistics, discussion); Bilardo and Rocchi 2011: 226 (biology); Bilton 2015: 446 (biology); Nilsson 2015: 208 (catalogue, faunistics). [Comment: literature, associated with Africa are only included. Accordingly, the list is incomplete for non-African species.]

Diagnosis

According to Miller and Bergsten (2014) the tribe Laccophilini, including the genus Laccophilus, is characterized by not visible scutellum when elytra closed, a single metatarsal claw, and prominent lobes at the anteroapical apices of the metatarsomeres. All African species of Laccophilus have bifid metatibial spines (Fig. 9), which separate them from the other Laccophilini genera in Africa.

Description

Body parameters: Length of body 2.8–6.0 mm, width 1.5–3.4 mm. Shape somewhat variable, elongate to oval, rarely sub-cylindrical (Fig. 382). Often, posteriorly flattened, with various colour pattern (Figs 393, 401, 451, 457, 471, 489, 515, 526).

Microsculpture and reticulation of two different kinds: Simple (meshes equally large, almost uniform, no size categories of meshes distinguished) and double (meshes of two kinds; size categories distinguished). When distinctly double, body covered with large meshes which generally contain a various number (2–8) of fine (less pronounced) meshes. Commonly, lines of large meshes in part reduced and weakly developed; sometimes almost absent and only discerned as fragments/rudiments. Less commonly, lines of finer meshes are reduced and difficult to discern within larger meshes. Sometimes mesh-categories in part mixed and microsculpture appears indistinct or absent while distinct in another location of same specimen. Rarely meshes of microsculpture elongated, being comparatively long in relation to breadth. Dorsal surface of body shiny to mat. Large parts of body in ventral-aspect with very fine, simple and slightly undulate linear microsculpture, which can be reduced, in part absent. Punctures on dorsal surface of body generally sparse and concentrated to various regions. Head at eyes with fine and irregular punctures. Punctures at area of head often enlarged narrowly towards head-centre, forming a sparse, transverse row of punctures connecting ocular punctuate areas. On pronotum fine punctures often discernible, especially at pronotal margins. Elytra with fine, irregular, longitudinal rows of punctures often discernible on disc, dorsoventrally and laterally. Ventral surface largely lacking punctures. Apical ventrite, however, generally with scattered, fine punctures. Lateral, pre-apical furrow of elytra generally distinct and pubescent.

Ventral aspect: Prosternal process slender, often strongly extended posteriorly and apically pointed (Figs 1–5). Metacoxal plates often provided with transverse, slightly obscure and shallow furrows, which can be rather indistinct. Stridulatory apparatus, when present, is located posteriorly on metacoxal plates, quite close to midline of body. Apparatus consists of dense ridges forming a semicircular file (Fig. 6). All African Laccophilus species have curved, fine striae on basal ventrites of abdomen (Fig. 6). Apical ventrite variable in shape, often modified and asymmetric, provided with a fine knob-like process on one side (Figs 110, 118). Apical ventrite with posterior edge modified with medial part posteriorly to a variable degree extended (Figs 47, 112). Some species groups lack modifications on apical ventrite (Figs 26, 43). Metacoxal process posteriorly rarely expanded (Fig. 7).

Legs: Male pro-and mesotarsus slightly enlarged and provided with suckers, length of which is variable (Fig. 10) – female lacks suckers. Metatibial spurs bifid (Fig. 9).

Sexes: Similar but males provided with pro- and mesotarsal suckers. Male apical ventrites in many species groups more strongly modified than in female; often asymmetric with one-side lateral knob on apical ventrite. Rarely female epipleuron with intraspecific, partial enlargement (Fig. 8).

Distribution

Global distribution covering all continents but Antarctica. According to the world catalogue 263 species recognized (Nilsson 2015).

Ecology and collecting circumstances

In Africa the genus occurs in all kinds of freshwater habitats. Often collected in quite shallow water with sparse vegetation on sandy-clay-bottom, e.g. in drinking pools for domestic animals. No comprehensive work on ecology of Laccophilus exists. Scattered information can be obtained by scrolling through faunistic literature, here listed. Additional sparse information on ecology is documented on many collecting labels. Experiences from Madagascar by the last author gives Laccophilus as one of the most ubiquitously occurring dytiscid genera. Different species have been found from sea level up to an altitude of over 2000m. Laccophilus inhabits many types of both lotic and lentic waters with different species and species groups more specialized. The L. alluaudi species group for example contains typical lotic species. The group is characteristic of small to medium-size canopy-covered rainforest streams with sandy or gravel bottoms lacking vegetation but collecting dead leaves at margins. Species from other groups like the L. taeniolatus group are often very abundant in red-clayish ponds visited by zebu cattle. The L. leonensis group can be found in vegetation-rich forest swamps and marshes or at margins of slow flowing vegetated sections of open landscape meandering rivers. When taken out of the water and put on dry land and when disturbed they can jump distances at least 20x their own body length. The behavior has not been studied in detail and could be both an anti-predatory escape behavior or used when semipermanent streams or side pools gradually dry out and the beetle can without flying move sideways or downstream to new habitats.

Laccophilus: immature stages

As larval morphology is known only for six of the 13 genera of Laccophilinae (Miller et al. 2005), it is hard to give a valid diagnosis for the entire subfamily. Larvae of the genus Laccophilus were described in detail by Alarie et al. (2000) based on the study of seven species from Europe and the New World including L. poecilus Klug known also from Africa. In the same work, characters were presented for the separation of especially instar III Laccophilus larvae from those of the genera Africophilus Guignot, 1948, Australphilus Watts, 1978 and Neptosternus Sharp, 1882. Mature Laccophilus larvae are good swimmers with legs provided with dorsal rows of natatory setae on tibiae and tarsi. The coxae and femora are provided with rows of comb-like spinulae or pectens. The larvae have long urogomphi provided with many secondary setae. A specific feature of the instar I larva is the strong submedian constriction of the frontoclypeus that bears only two marginal spatulate setae. Miller et al. (2005) provided a larval description as well as molecular characters used to link larvae and adults of African Philodytes umbrinus. They discuss characters shared between Philodytes and known Laccophilus larvae as well as diagnostic differences.

Larvae of only a few Afrotropical Laccophilus species have been described so far, and most of the descriptions are not very detailed. As already mentioned, Alarie et al. (2000) described the instar III larva of L. poecilus, and all three instars of this species had previously been described in much detail by De Marzo (1976; as L. variegatus Dejean). As no other larvae have been reared from eggs laid in captivity, identifications remain slightly uncertain, and some of the descriptions refer only to Laccophilus spp. Bertrand (1928) described the larva of L. complicatus Sharp based on material from Madagascar, and later he provided descriptions of unidentified species collected in the Democratic Republic of the Congo, Guinea and the Ivory Coast (Bertrand 1935, 1948 & 1954). More unidentified Laccophilus larvae were reported by Bertrand (1966a & 1968) from various wetlands in the Democratic Republic of the Congo, Kenya, Rwanda, Uganda, and Tanzania. Based on differences in the pigmentation of the head capsule and association with adults, Bertrand (1966b, 1969) gave records of supposed larvae of L. adspersus Boheman, L. cyclopis Sharp and L. lineatus Aubé from various ponds and streams in Lesotho, South Africa and Zimbabwe. Larvae of L. adspersus were also reported from Kenya (Bertrand 1963). These and a few more records of Laccophilus larvae from Africa were reviewed by Bertrand (1972).

The pupal stage has been described briefly for selected Laccophilus species by Wilson (1923) and Bertrand (1928). A more detailed description of the pupa of the Nearctic L. fasciatus rufus was provided by Sizer et al. (1998). No pupae of African origin have been studied so far.

Species groups of African Laccophilus

The present revision of the genus Laccophilus focuses on the species occurring in the African mainland and Madagascar with its neighbour islands. Accordingly, from this standpoint no thorough analysis of the phylogeny of the genus is therefore possible. About 60% of the recognized species-bulk is distributed outside Africa on various other continents. Plans for future, however, include a phylogenetic survey of the whole genus on a global basis in which both morphological characters and molecular data will be considered. The forthcoming study will also focus on groups of species recognized in the genus Laccophilus.

Despite problems in understanding Laccophilus systematics at a global level a division of the genus in different species groups only for Africa including Madagascar is justified and can be motivated by practical reasons. Management of a total 105 species can be quite demanding without division in practical groups. The survey here undertaken does not count on detection of synapomorphies for delimitation of monophyletic groups, but is based on simple similarity (presence of shared characters). Below, recognized and examined characters are briefly described and discussed. Three additional Laccophilinae genera are included in the survey: Philodytes J. Balfour-Browne, 1939, Neptosternus Sharp, 1882, and Philaccolus Guignot, 1937.

The recognized groups of species in African Laccophilus, introduced in this revision do not coincide well with those presented by Guignot (1959a). Within species groups recognized the species are listed in accordance with morphological similarity between the species.

In Table 1 the possession of the discussed characters in the species groups and three reference genera is presented.

Species groups and outgroups scored for characters 1-10 discussed in the text.

Species group/character numbers 1 2 3 4 5 6 7 8 9 10
sp.gr. 1 (morondavensis) 1 2 1 2 1 2 2 2 2 1
sp.gr. 2 (ruficollis) 1 1 2 2 1 2 2 2 2 1
sp.gr. 3 (hyalinus) 1 1 2 1/2 1 2 2 2 2 1
sp.gr. 4 (alluaudi) 1 1 2 2 1 2 2 2 2 1
sp.gr. 5 (isamberti) 1 1 2 2 1 2 1 2 2 1
sp.gr. 6 (pictipennis) 1 1 2 2 1 2 2 1 2 1
sp.gr. 7 (taeniolatus) 1 1 2 2 1 1 2 1 2 2
sp.gr. 8 (immundus) 1 1 2 2 1 2 2 1 2 2
sp.gr. 9 (pellucidus) 1 1 2 2 1 2 2 1 2 2
sp.gr. 10 (adspersus) 1 1 2 2 1 1 2 1 1 2
sp.gr. 11 (deceptor) 1 1 2 2 1 1 2 1 1 2
sp.gr. 12 (poecilius) 1 1 2 2 1 2 2 1 1 2
sp.gr. 13 (lineatus) 1 1 2 2 1 1 2 1 1 2
sp.gr. 14 (desintegratus) 1 1 1 2 1 1 2 1 1 2
sp.gr. 15 (luctuosus) 1 1 2 2 1 1 2 1 1 2
sp.gr. 16 (leonensis) 1 1 2 1 1 1 2 1 1 2
sp.gr. 17 (laeticulus) 1 1 2 1 1 1 2 1 1 2
Philodytes 2 1 2 2 1 1 2 1 2 2
Neptosternus 2 1 1? 2 2 2 2 2 2 1
Philaccolus 2 1 1? 1 1 2 2 2 2 2
  1. Apices of metatibial spurs bifid (Fig. 9) (1) / Not bifid (2).

    The bifid spurs have total coverage in Laccophilus, all species exhibiting the feature. In all other Laccophilinae genera the corresponding spines are pointed and most probably bifid spines are a derived character, which indicates the genus Laccophilus is monophyletic. However among species outside Africa Balke et al (1997) have reported Laccophilus species with pointed metatibial spurs, a likely reversal.

  2. Body in posterior half dorso-ventrally somewhat flattened (Fig. 418) (1) / Body subcylindrical (Figs 378–384) (2).

    The dorso-ventrally flattened body is widely distributed in Laccophilinae and also present in all but one species group of Laccophilus. Accordingly, the feature could represent a plesiomorphy while the subcylidrical body-shape is a synapomorphy, characteristic of the L. morondavensis species group.

  3. Microsculpture of dorsal body surface simple (1) / Double, mixed (2).

    Simple microsculpture indicates reticulation on body, where the meshes are similar in size and shape. No size-categories can be distinguished between meshes, neither are there differences in their qualitative feature – the meshes form a smooth coverage on body surface. Double microsculpture means that there are two kinds of reticulation mixed on same location; large meshes and small meshes. The large meshes are generally more strongly impressed in the body surface than the small ones. A large mesh encloses often a number of small meshes, which can vary in number between 2 and 8. Reduction of meshes occurs often in regard of large meshes but can also be the case for small meshes. When reduced, the meshes are either in part or totally lacking. In such cases fragments of meshes can be detected, mixed with complete meshes of different kind. Sometimes mesh-categories appear variable so that division in size-classes is impossible. Rarely the meshes are deformed and their shape is elongate. Two species groups of African Laccophilus have simple microsculpture (i.e. L. morondavensis and L. desintegratus species groups), while 15 species groups seem to have double, by the above definition.

  4. Metacoxal plates have a stridulatory file (Fig. 6) (1) / Stridulatory file absent (2).

    The stridulatory file is a semicircular device which is formed by densely located ridges on the metacoxal plate. The function of it has not been thoroughly studied and possible sound has not been heard, nor recorded. In African Laccophilus both sexes seem to have the device when present. Presumable use of it can be related to intraspecific communication but it may also be used e.g. in defence against predators. Three species groups of African Laccophilus exhibit this feature. In one group (L. hyalinus species group) with a modest number of species, only two species have it. In the two remaining species groups (L. leonensis and L. laeticulus species groups) all representatives are provided with it. In the latter species group the file, however, is very weakly developed and may be rudimentary and out of function. A similar stridulatory device is also present in other Laccophilinae genera, e.g. in genus Philaccolus while lacking in e.g. Philodytes.

  5. Abdominal ventrites provided with sparse, somewhat curved striae (Fig. 6) (1)/ Striae absent (2).

    All African Laccophilus species have a number of sparse, curved striae on abdominal ventrites. This character seems to be widely distributed in Laccophilinae as it may be recognized at least in Philaccolus and Philodytes and in a reduced state in Neptosternus.

  6. Prosternal process slender, posteriorly distinctly extended, apically pointed (Fig. 5) (1) / Prosternal process shorter; comparatively broad, posteriorly not strongly extended (Figs 1–4) (2).

    At least 10 species groups of Laccophilus have slender, extended prosternal process while in 7 species groups the process is shorter and broader, which seem to be the case in Laccophilinae outside Laccophilus as well. The slender process may be a synapomorphy of a supposed clade containing the respective species groups.

  7. Metacoxal process posteriorly expanded and modified (Fig. 7) (1) / Metacoxal process posteriorly not expanded; ends abruptly (Fig. 6) (2).

    In Laccophilus one species placed in its own species group (L. isamberti species group) exhibits this enigmatic and unique feature. It definitely represents the derived state and future studies will reveal if the species deserves a status of a separate genus within Laccophilinae.

  8. Posterior edge of apical ventrite modified, forming an undulate structure, with medial part distinctly extended backwards (Fig. 69) (1) / Posterior edge of apical ventrite not modified; outline of ventrite smoothly curved (Fig. 25) (2).

    In African Laccophilus 13 species groups out of 17, exhibit the modified apical ventrite. Besides Laccophilus, Philodytes has a similar modified apical ventrite while at least Philaccolus and Neptosternus lack it.

  9. Male apical ventrite strongly asymmetric, when provided with a distinct, small knob or process on one side of the midline of ventrite (Fig. 69) (1) / No asymmetric knob or process on male apical ventrite (Fig. 47)(2).

    In total 9 recognized species groups of Laccophilus in Africa, exhibit this, most probably derived character. It may turn out to be a good synapomorphy for them.

  10. Penis apex narrow, often curved and exhibits only slight modifications in anatomical shape (Figs 212, 237) (1) / Penis generally strongly modified, exhibiting various anatomical details (2).

    In all six species groups of Laccophilus here distinguished, have a slender to rather slender, often quite evenly curved penis, lacking considerable modifications.

Key to species groups of African Laccophilus

To be considered slightly tentative and mostly only applicable for male specimens.

1 Male apical ventrite symmetric, lacking lateral process/knob (Figs 12, 27, 51) 2
Male apical ventrite strongly asymmetric, provided with a minute, lateral process (located to left on ventrite, when viewed from below) (Figs 69, 96, 207) 10
2 Large species (body length 5.3–6.0 mm); body dorsoventrally flattened (Fig. 413); penis apex strongly modified (Fig. 259) group 9 (L. pellucidus) (p. 89)
Smaller species with flattened or subcylindrical body (body length 3.2–5.8 mm); penis shape different, variable 3
3 Body subcylindrical (Fig. 382); generally large species (body length 4.1–5.7 mm), except one species (3.1–3.4 mm); body microsculpture simple, of one kind group 1 (L. morondavensis) (p. 17)
Body dorsoventrally flattened; small to large species (body length 3.2–5.8 mm); body microsculpture (dorsal aspect) double (can be reduced, rarely distinctly so) 4
4 Metacoxal process posteriorly expanded (Fig. 7) group 5 (L. isamberti) (p. 47)
Metacoxal process posteriorly truncate, not expanded (Fig. 6) 5
5 Elytra provided with longitudinal, dark markings (Fig. 391) (incl. L. rivulosus (Fig. 416)) group 4 (L. alluaudi) (p. 36)
Elytral markings variable; never forming distinct, dark, longitudinal markings (excl. L. rivulosus (Fig. 416)) 6
6 Posterior margin of male apical ventrite modified, undulate with middle part posteriorly extended (Fig. 55) 7
Posterior margin of male apical ventrite not modified; non-undulate, posterior margin curved and medially not posteriorly extended (Figs 22, 24) 9
7 Elytra pale ferrugineous to pale brownish with dense, dark ferrugineous to blackish irrorations/ undulations (Figs 400, 411); one paramere (upper in illustrations) apically enlarged (Fig. 255) group 7 (L. taeniolatus) (p. 52)
Elytral colour pattern different; either uniformly, dark ferrugineous (Fig. 417) or with extensive patches (Fig. 398); parameres different, apically not enlarged (Figs 248, 258) 8
8 Elytra with distinct colour pattern (with extensive patches) (Fig. 398); penis evenly curved, narrows gradually towards apex (Fig. 247) group 6 (L. pictipennis) (p. 49)
Elytra ferrugineous to dark ferrugineous, lacking distinct paler areas (Fig. 417); penis with peculiar curvature and apical expansion (Fig. 258) group 8 (L. immundus) (p. 87)
9 Small species (body length 3.2–3.4 mm); elytra dark ferrugineous with distinct, transverse, pale ferrugineous markings (Fig. 385) group 2 (L. ruficollis) (p. 26)
Larger species (body length 4.0–5.3 mm); elytra generally pale brownish to ferrugineous, often with vague, pale ferrugineous to pale brownish areas, or colour pattern absent (Figs 386, 389) group 3 (L. hyalinus) (p. 28)
10 Metacoxal plates provided with a semicircular stridulation apparatus (Fig. 6) 11
Metacoxal plates lack stridulation apparatus 12
11 Penis in lateral view evenly curved; almost evenly broad from middle to apex; inner outline of penis close to apex uneven (provided with minute ridges) (Figs 362, 367) group 16 (L. leonensis) (p. 237)
Penis in lateral view slightly angled; narrows distinctly from middle to apex; inner outline of penis close to apex smooth (lacks minute ridges) (Figs 373, 377) group 17 (L. laeticulus) (p. 256)
12 Elytral colour pattern distinct, consists of dark longitudinal markings, which may be undulate and connected with neighbour-markings; sometimes markings merged into extensive dark areas (Figs 472, 476, 480) or reduced (Figs 478, 483) 13
Elytral colour pattern different (note that there are species with extensive dark elytra), not provided with distinct longitudinal, dark markings; often patchy (Figs 457, 467) or rather pale with dense irrorations/undulations (Figs 419, 437) 15
13 Body (dorsal aspect) microsculpture simple, of one kind (no fragments of large meshes discernible); penis apex broad, truncate (Fig. 358) group 14 (L. desintegratus) (p. 231)
Body (dorsal aspect) microsculpture double, consists of two kinds of microsculpture, mixed: often larger meshes of microsculpture reduced in part (fragments of large meshes generally discernible); shape of penis different 14
14 Penis robust, curved, apically provided with a distinct extension and inner outline provided with distinct ridges (Fig. 359); elytra extensively dark with transverse, basal, pale marking which can be reduced to separate spots (Figs 503–505); small species (body length 2.9–3.6 mm) group 15 (L. luctuosus) (p. 234)
Penis slender to robust, apex forming a distinct hook/angled enlargement (Figs 327, 335) or penis curved, sometimes also twisted (Figs 345, 350); elytral colour pattern different (variable); small to large species (body length 2.9–5.1 mm) group 13 (L. lineatus) (p. 178)
15 Penis (lateral aspect) externally close to base with a deep incision (Fig. 322) group 12 (L. poecilus) (p. 176)
Penis (lateral aspect) externally close to base without deep incision 16
16 Elytra pale ferrugineous to pale brownish, generally with extensive, often delicate, dark irrorations (Figs 419, 425), or almost unicoloured, brownish or blackish (Figs 436, 445); penis rather slender, curved or angled and provided with a distinct apex (Figs 261, 270, 289) group 10 (L. adspersus) (p. 95)
Elytral colour pattern different; consists of pale patches arranged in variable, transverse series (Figs 453 461); penis different, variable in shape (e.g. Figs 303, 307, 313, 318, 320) group 11 (L. deceptor) (p. 159)

Species group 1 (L. morondavensis group)

Diagnosis. Large species with length of body 4.1–5.7 mm, width 1.9–3.0 mm, except one small species, L. tavetensis, with length 3.1–3.4 mm and width 1.6–1.7 mm.

Shape of body subcylindrical, dorsoventrally not flattened (Fig. 382). Body dorsally, with distinct colour pattern, which is formed by rather extensive, often longitudinal, dark/pale patches especially on elytra (Fig. 381). One species with body, dorsal aspect, lacking distinct colour pattern; ferrugineous to dark ferrugineous (Fig. 384). Body microsculpture simple, of one kind.

Prosternal process moderately broad, posteriorly not distinctly extended, apically pointed. Apical ventrite modified; posteriorly on both side of midline more or less excavated and post-medially often extended to a narrow enlargement (Fig. 15). Apical ventrite lacks asymmetrical, small knob. No stridulatory apparatus on metacoxal plates. Metacoxal process not extended posteriorly (Fig. 6).

Paramere simple, elongate, apically not distinctly enlarged or modified (Fig. 218). Apical half of penis slender to quite slender, almost straight to distinctly curved, apically not distinctly modified (Fig. 214). One species with tip of penis slightly enlarged (Figs 222–223).

Species composition and distribution. Seven African species are included to the group (see the identification key). None of them occurs outside Africa south of Sahara or Madagascar.

Note. Male of L. mirabilis is unknown. Large body (length 5.1–5.7 mm). Elytral colour pattern consisting of separate, longitudinal dark markings (Fig. 383). Only known from Madagascar.

Key to species (males)

1 Small species, length of body less than 3.4 mm L. tavetensis (p. 18)
Larger species, length of body between 4.1–5.7 mm 2
2 Body, dorsal aspect, lacks distinct colour pattern (Fig. 384); penis tip with slight enlargement (Figs 222–223) L. ferrugo (p. 25)
Body, dorsal aspect, with distinct colour pattern (Fig. 381); penis tip not enlarged 3
3 Penis, lateral aspect, comparatively broad; apex of penis distinctly curved backwards (Fig. 216) L. morondavensis (p. 22)
Penis, lateral aspect, narrower; apex of penis not curved backwards 4
4 Body broad, oval, large (length 4.9–5.2 mm) (Fig. 379) L. grossus (p. 19)
Body more elongate, smaller (length 4.1–4.9 mm) (Fig. 380) 5
5 Body elongate, slender; pale areas on elytra open with no closed cells or one inner cell formed by dark, narrow, longitudinal marking (Fig. 382) L. productus (p. 23)
Body slightly broader; pale areas on elytra with two closed cells, formed by dark, narrow, longitudinal markings (Fig. 380) L. rocchii (p. 20)

Laccophilus tavetensis Guignot, 1941

Figs 11, 209–211, 378, 527

Laccophilus tavetensis Guignot 1941: 36 (original description, discussion, faunistics); Guignot 1946c: 283, 285, 313 (redescription, faunistics); Guignot 1959a: 585, 587 (redescription, faunistics); Nilsson 2001: 251 (catalogue, faunistics); Nilsson 2015: 218 (catalogue, faunistics).

Type locality

Kenya: Taveta.

Type material studied

(1 ex.). Holotype: male: “Afrique Orient. Anglaise Taveta Alluaud & Jeannel mars 1912 – 750 m St. 65 / male symbol / Type / Det. Dr. Guignot Laccophilus tavetensis Guign. Type” (MNHN).

Additional material studied

(11 exs.). Sudan: “Prov. N Darfur El Geneina / ad lucem Ibrahim M. Abuzinid 20.8. 1979” (1 ex. CGW). – Kenya: “S, Voi 11. 1997 leg. Snizek” (4 exs. CFP, 4 exs. CSR); “Kenya eastern Sosoma ca. 200 km E of Thika 27.11. 2011, light trap” (1 ex. NMPC). – Botswana: “Chobe NP Savuti-Camp 18°33'55"S-24°03'53"E, 11.3. 1993 lux leg. Uhlig” (1 ex. ZMHB; habitus in Fig. 378).

Diagnosis

A deviate species, separated from the other species in this species group by having small body size in combination with peculiar shape of penis; somewhat sinuate and distinctly enlarged posterior to narrow apex. Note also differently shaped male apical ventrite in comparison with other species in the species group (Fig. 11).

Description

Body length 3.1–3.4 mm, width 1.6–1.7 mm. Pale ferrugineous, dorsal colour pattern ferrugineous and sometimes vague and slightly variable (Fig. 378).

Head: Pale ferrugineous, no colour pattern. Submat, with fine, dense microsculpture. Reticulation simple; only with small, uniform meshes. Impunctate, except at eyes; with fine, irregularly located punctures.

Pronotum: Pale ferrugineous, no distinct colour pattern. Submat, with fine, dense microsculpture. Reticulation simple; only with small, uniform meshes. Impunctate, at margins with fine to very fine, somewhat irregular punctures. Mediobasally punctures absent.

Elytra: Pale ferrugineous, sometimes with vague, ferrugineous, longitudinal markings (Fig. 378). Elytral colour pattern sometimes rather indistinct. Submat, with fine, dense microsculpture. Reticulation simple; only with small, uniform meshes. Fine, somewhat irregular punctures form a discal row. Dorsolateral and lateral rows of punctures indistinct; indicated by scattered, fine punctures. Laterally with a quite long, sparsely pubescent, pre-apical furrow.

Ventral aspect: Abdomen dark ferrugineous to ferrugineous, metathorax and –coxal plates ferrugineous, and prothorax pale ferrugineous. Submat, finely microsculptured. Abdomen with fine, curved striae. Impunctate, except a few fine punctures on apical ventrite; symmetric (Fig. 11). Prosternal process quite narrow, apex somewhat enlarged, short, apically pointed (arrow-shaped). Transverse shallow furrows reduced; 2–3 indistinct, reduced furrows discernible.

Legs: Pro- and mesotarsus slightly enlarged, extended, provided with suckers.

Male genitalia: Penis in dorsal aspect clearly sinuate with narrow tip; in lateral aspect almost evenly curved Figs 209–211.

Female: Unknown.

Distribution

Sudan, Kenya, Botswana (Fig. 527).

Collecting circumstances

In Botswana collected with light.

Laccophilus grossus sp. n.

Figs 12–13, 212–213, 379, 527

Type locality

Namibia: Damaraland, Oshikango (15.55E, 17.25S).

Type material

(5 exs.). Holotype, male: “South Africa Damaraland Oshikango, v. 1948 15.55E, 17.25S, C. Koch / B. Malkin Coll. BMNH (E) 1956–234” (BMNH). – Paratypes: Same data as holotype (1 ex. MZH; habitus in Fig. 379); “Angola Rocadas R. Cunene 19–22.2. 1972/at light“ (1 ex. CFP); “Angola Rocadas 30.3. 1972” (1 ex. CFP); “Namibia 23.2. 1994 17°26'S/14°21'E, Kunene, Ruacana Dorp, lux, leg. M. Uhlig” (1 ex. ZMHB).

Diagnosis

Laccophilus grossus belongs to a group of species, characterized by large body-size, by uniform microsculpture, with one kind of meshes (small) and by slender, slightly sinuate penis. The new species is probably closest related to L. rocchii, another so far undescribed species. The two species are distinguished by difference in body size, by deviating dorsal, colour pattern of body and by details in shape of penis apex (curved in different directions).

Description

Body: Length 4.9–5.2 mm, width 2.7–2.8 mm. Dorsal colour pattern exhibits only slight variation (Fig. 379).

Head: Pale ferrugineous to ferrugineous to brownish; posteriorly at pronotum slightly darker than anteriorly; however, change of colour gradual and no colour pattern formed. Submat, entire head finely microsculptured; meshes small and only of one kind. Impunctate, except at eyes, with some fine, irregular punctures. Anteriorly, close to edge of head with a few transverse impressions formed by elongated punctures.

Pronotum: Pale ferrugineous to ferrugineous, medially broadly, distinctly darker; basal area blackish. Submat, finely and densely microsculptured. Meshes of microsculpture small, uniform and of one kind only. Impunctate, except at margins, finely and somewhat irregularly punctate. Broad area basally in middle lacking punctures.

Elytra: Pale ferrugineous, with blackish to dark ferrugineous, slightly variable marking (Fig. 379). Submat, with fine, uniform, evenly distributed microsculpture. Meshes of microsculpture quite small, of one kind. Fine, irregular punctures form a discal row of punctures, which spread out and disappears posteriorly. Scattered, fine punctures indicate presence of a vague, dorsolateral and lateral row of punctures. Pre-apical, lateral row of punctures comparatively long, forms a, in part, distinct furrow with some setae.

Ventral aspect: Blackish to dark ferrugineous; no distinct colour pattern formed. Submat, finely to very finely microsculptured. Abdominal ventrites with dense, curved striae. Metacoxal plates with some transverse furrows, which posteriorly fade away. Apical ventrite lacks asymmetric knob/process (Fig. 12). Prosternal process rather slender, apex moderately, posteriorly extended, apically pointed. Almost impunctate, apical ventrite with some scattered punctures.

Legs: Pale ferrugineous, hindlegs slightly darker, ferrugineous to brownish. Pro- and mesotarsus slightly enlarged, with fine suckers.

Male genitalia: Apical half of penis slightly sinuate and when viewed from above; tip of penis slightly curved right (Figs 212–213).

Female: Pro- and mesotarsus rather slender. Apical ventrite as in Fig. 13.

Etymology

The species name grossus is a Latin adjective meaning “big”. It here associates with the body size of the new species.

Distribution

Angola, Namibia (Fig. 527).

Collecting circumstances

Almost unknown. In Angola collected at light.

Laccophilus rocchii sp. n.

Figs 14–15, 214–215, 380, 527

Type locality

Mozambique: Manica Province, 60 km W Chitobe.

Type material

(15 exs.). Holotype: male: “Mozambique Manica Province 60 km W Chitobe, 16.12. 2005 P. Schüle leg.” (SMNS). – Paratypes: “Tanzania Dodoma Pr. 40 km N Dodoma 14–16.12. 2006, 1100 m A. Kudrna Jr. lgt.” (1 ex. CFP); “Botswana: Chobe Dist., Savute Drift Camp site, 18°34'S, 24°04'E, 29. Dec. 1988 R.D. Ward / Robert D. Ward Collection / Laccophilus productus Rég. det. S. Rocchi 92” (1 ex. CSR; habitus photogr. Fig. 380); Similar label data as holotype (6 exs. SMNS, 1 ex. MZH); “Mocambique Prov. Inhambane 15 km SE Save, 18–21.12. 2005, A. Kurdna Jr. lgt.” (3 exs. CFP, 1 ex. MZH); “Namibia Exp. ZMB 1992 East Caprivi: Katima Mulilo, lux, 17°29'S/24°17'E, 3–8.3. 1992 leg. M. Uhlig” (1 ex. ZMHB).

Diagnosis

See diagnosis of L. grossus (p. 19).

Description

Body: Length 4.3–4.9 mm, width 2.3–2.6 mm. Body dorsally pale ferrugineous, with quite distinct and uniform blackish ferrugineous to dark ferrugineous marking (Fig. 380).

Head: Pale ferrugineous. Submat, finely microsculptured; reticulation simple. Meshes small, of same size and shape. Impunctate, except at eyes where head is provided with fine, somewhat irregularly distributed punctures. Anteriorly, close to frontal edge with some punctures, forming slightly irregular transverse impression.

Pronotum: Pale ferrugineous, basally in middle with distinct blackish ferrugineous spot. Submat, finely microsculptured; reticulation simple. Meshes small, of same size and shape. Impunctate, except along margins, with irregular, fine punctures, however, punctures lacking basally in middle.

Elytra: Pale ferrugineous, with quite distinct, quite uniform blackish ferrugineous to dark ferrugineous marking (Fig. 380). Submat, finely microsculptured; reticulation simple, of one size-category. Meshes small, size and appearance uniform. Discal and dorsolateral row of punctures consist of irregular, fine punctures. Rows are diffuse and mixed posterior to middle of elytra. Lateral row indicated by some scattered, fine punctures. Preapical, lateral row of punctures located in a distinct furrow provided with some hairs.

Ventral aspect: Pale ferrugineous to ferrugineous, no distinct colour pattern formed. Submat, very finely microsculptured, except abdomen basally, rather shiny, microsculpture indistinct. Apical ventrite of male (Fig. 14). Ventrites with fine, curved, and quite dense striae. Metacoxal plates with shallow, transverse furrows, which posteriorly, gradually become weaker. Prosternal process rather slender, posteriorly slightly extended, apex pointed.

Legs: Pro- and mesotarsus somewhat enlarged, with suckers.

Male genitalia: Apical half of penis in dorsal aspect only slightly sinuate, almost straight; extreme tip slightly curved to left (Figs 214–215).

Female: Pro- and mesotarsus slender, not enlarged. Apical ventrite as in Fig. 15.

Etymology

The name is a noun in its genitive form based on the name of Mr. Saverio Rocchi, Florence, Italy, who kindly provided us with a part of the type material of the new species besides various other interesting materials.

Distribution

Tanzania, Namibia, Botswana, Mozambique (Fig. 527).

Collecting circumstances

In Namibia collected at light.

Laccophilus morondavensis Guignot, 1957

Figs 16, 216–218, 381, 527

Laccophilus morondavensis Guignot 1957b: 72 (original description, faunistics); Rocchi 1991: 86 (faunistics, list); Nilsson 2001: 247 (catalogue, faunistics); Nilsson 2015: 214 (catalogue, faunistics).

Type locality

Madagascar: Foret sud de Befasy.

Type material studied

(1 ex.). Holotype: male: “Morondava foret sud de Befasy I-56 R.P. / Institut Scientifique Madagascar / F. Guignot det., 1956 Laccophilus morondavensis sp. n. Type, male symbol” (MNHN; habitus in Fig. 381).

Additional material studied

(1 ex.): Madagascar: “W Madag. 60 km NE of Morondava, Foret de Kirindi, 30 m Bednarik leg. 28.1.1996 / L. morondavensis Guignot 1957 Jiri Hájek det. 2006” (1 ex. NMPC).

Diagnosis

Laccophilus morondavensis is characterized by its distinct, elytral colour pattern and by peculiarly shaped penis apex. The species resembles externally most of L. productus but body is somewhat larger and broader. Additionally, pronotum is extensively dark while in L. productus almost entirely pale ferrugineous. Tip of penis is slightly upwards curved in L. morondavensis while it is almost straight in L. productus.

Description

Body length 5.3 mm, width 2.9 mm. Dorsal, aspect of body with rather distinct colour pattern (Fig. 381).

Head: Pale ferrugineous. Posteriorly, head becomes gradually slightly darker but lacks distinct colour pattern. Rather shiny, although finely and densely microsculptured; reticulation simple, of one kind. Impunctate, except at eyes, with fine, slightly irregular punctures. Medially, areas with punctures extend slightly towards centre of head.

Pronotum: Blackish to ferrugineous, laterally pale ferrugineous. Colour change gradual; colour pattern vague. Rather shiny, although finely microsculptured. Reticulation of one kind; consists of small meshes. Entire disc with fine, sparse punctures. At margins, except mediobasally, with slightly irregular, coarse punctures.

Elytra: Pale ferrugineous, with blackish to dark ferrugineous markings (Fig. 377). Rather shiny, although finely microsculptured. Reticulation dense, of one kind; meshes moderately sized. Discal row of punctures consists of fine to very fine, scattered, punctures. Dorsolateral and lateral row of punctures as discal row but sparser and more irregular. Laterally, elytra with a rather shallow pre-apical, finely pubescent and quite extensive furrow.

Ventral aspect: Blackish ferrugineous to dark ferrugineous, prothorax paler; pale ferrugineous to ferrugineous. Rather shiny, although finely microsculptured. Abdominal reticulation reduced, in part absent. Abdomen, with fine, curved striae. Metacoxal plates with reduced, transverse furrows, which are only discernible in anterior half. Almost impunctate. Apical ventrite, with punctures, symmetric, lacks lateral knob (Fig. 16). Prosternal process rather slender, apex arrow-shaped, quite short, pointed.

Legs: Pro- and mesotarsus rather slender, somewhat extended, with suckers.

Male genitalia: Penis both in lateral and dorsal aspect broader than related species; apical tip curved upwards (Figs 216–218).

Female: Unknown.

Distribution

Madagascar (Fig. 527).

Collecting circumstances

Unknown.

Laccophilus productus Régimbart, 1906

Figs 17–18, 219–221, 382, 527

Laccophilus productus Régimbart 1906: 249 (original description, faunistics); Zimmermann 1920a: 25 (catalogue); Peschet 1921: 6 (discussion, description, faunistics); Zimmermann 1926: 23 (faunistics); Guignot 1946c: 284, 313 (description, faunistics); Guignot 1957b: 73 (discussion, faunistics); Guignot 1959a: 585, 586 (redescription, faunistics); Nilsson 2001: 249 (catalogue, faunistics); Nilsson 2015: 216 (catalogue, faunistics).

Type locality

Kenya: Samburu.

Type material studied

(5 exs.). Lectotype (by present designation): male: “Afrique Orle Anglaise Samburu (Wa-Nyika) Dr. Alluaud IV. 1904 / Museum Paris coll. Ch. Alluaud / TYPE / Laccophilus productus Rég. sp. n. typ” (MNHN; top specimen on pin with two additional paralectotypes). – Paralectotypes: Similar data and on same pin as lectotype (2 exs. MNHN); “Samburu Wa-Nyika / Afr. Orle Angl. Alluaud / Museum Paris coll. Maurice Régimbart 1908 / productus Rég.” (2 exs. MNHN; habitus in Fig. 382).

Additional material studied

(3 exs.): Tanzania: “Kwakiyembe D.O.Afr. April 1916 Methner / L. productus Rég. det. Brancucci 1982” (1 ex. ZMHB); “Narobi b. Tanga 5. 1915 Methner” (1 ex. ZMHB); “Nord-Rabeho D.O. Afr. leg. Methner” (1 ex. ZMHB).

Diagnosis

Laccophilus productus is characterized by quite large but slender body and by peculiar dorsal colour pattern and male genitalia (penis apical half slightly twisted; extreme apex bent leftwards). The species resembles most of L. morondavensis which occurs in Madagascar; diagnostic features are given under diagnosis of L. morondavensis on p. 22.

Description

Body length 4.1–4.9, width 1.9–2.5 mm. Elytra with distinct colour pattern (Fig. 382); only minor variation exhibited.

Head: Pale ferrugineous. Submat, finely and densely microsculptured. Reticulation simple; only with small, distinct meshes. Impunctate, except at eyes with scattered, fine, punctures.

Pronotum: Pale ferrugineous, mediobasally with a vague ferrugineous to dark ferrugineous marking. Submat, finely and densely microsculptured. Reticulation simple; only with small, distinct meshes. Impunctate, except at margins; with fine, somewhat sparse and irregular punctures. Mediobasally punctures absent or indistinct.

Elytra: Dark ferrugineous, with subbasal, preapical and apical, pale ferrugineous area (Fig. 382). Colour pattern stable and exhibits only minor variation. Submat, finely and densely microsculptured. Reticulation simple; only with small, distinct meshes. Fine, sparse and somewhat irregular punctures form a discal row. Dorsolateral and lateral rows indicated by scattered, fine punctures. Laterally with a comparatively long, finely pubescent, pre-apical furrow.

Ventral aspect: Dark ferrugineous to ferrugineous; colour pattern vague, indistinct. Rather shiny, finely microsculptured. Abdomen with fine, curved striae. Almost impunctate, except for apical ventrite; with scattered irregular punctures and shape symmetric (Fig. 17). Metacoxal plates with 13–15 almost transverse, fine, shallow furrows which in part are rather indistinct. Prosternal process slightly enlarged; apex moderately extended, pointed.

Legs: Pro- and mesotarsus slightly enlarged, provided with suckers.

Male genitalia: Apical half of penis in dorsal aspect slightly sinuate; extreme apex slightly bent to left (Figs 219–221).

Female: Apical ventrite apically extended (Fig. 18). Pro- and mesotarsus slender, somewhat extended.

Distribution

Kenya, Tanzania (Fig. 527).

Collecting circumstances

Unknown.

Laccophilus mirabilis Guignot, 1956

Figs 19, 383, 527

Laccophilus mirabilis Guignot 1956d: 78 (original description, faunistics); Rocchi 1991: 86 (faunistics, list); Nilsson 2001: 247 (catalogue, faunistics); Nilsson 2015: 214 (catalogue, faunistics).

Type locality

Madagascar: Bas Mangoky.

Type material studied

(3 exs.). Holotype: female: “Type / Station Agric Bas Mangoky / Institut Scientifique Madagascar /Guignot det., 1956 Laccophilus mirabilis Type” (MNHN). – Paratypes: “Station Agric Bas Mangoky / female symbol / Paratype” (2 exs. MNHN; habitus in Fig. 383).

Diagnosis

Laccophilus mirabilis belongs to a distinct group of species characterized by body shape, being longer, thicker and relatively more slender than other African Laccophilus species. Other diagnostic features are the body microsculpture, which is simple and fine and shape of penis which is narrow and in dorsal view peculiarly, slightly twisted. Laccophilus mirabilis is thus far, however, only known from female but it can be separated from closely related, continental African species by being somewhat larger and by exhibiting different colour pattern of body.

Description

Body length 5.1–5.7 mm, width 2.8–3.0 mm. Only slight variation observed in elytral colour pattern (Fig. 383).

Head: Pale ferrugineous to pale brownish. Submat, finely and distinctly microsculptured. Reticulation simple, of one kind. Impunctate, except at eyes; with a few, fine and irregularly placed punctures. Additionally, in a small depression located a short distance from eyes towards middle with some fine punctures. Frontally along anterior edge with a faint, somewhat irregular impression.

Pronotum: Dark ferrugineous to ferrugineous. Laterally pronotum becomes gradually paler; pale ferrugineous. Submat, finely but distinctly microsculptured. Reticulation simple, of one kind. At margins except basally in middle with fine, sparse and irregularly located punctures. Extremely small, scattered punctures may be discerned on disc.

Elytra: Pale ferrugineous to ferrugineous, with fairly distinct dark ferrugineous markings (Fig. 383). Submat, finely and distinctly microsculptured. Reticulation simple, of one kind. Very fine, somewhat sparse and irregular punctures form a discal, dorsolateral and lateral row of punctures.

Ventral aspect: Blackish to dark ferrugineous. Prothorax ferrugineous to pale ferrugineous. Almost impunctate, except apical ventrite, which especially on apex is distinctly punctate (Fig. 19). Rather shiny, very finely and densely microsculptured. Abdomen (all visible ventrites) with somewhat sparse, curved striae. Metacoxal plates with some 10 shallow, transversely located, furrows. Prosternal process slightly enlarged, apex short, pointed.

Legs: Pro- and mesotarsus rather slender.

Male: Unknown.

Distribution

Madagascar (Fig. 527).

Collecting circumstances

Unknown.

Laccophilus ferrugo sp. n.

Figs 20–21, 222–223, 384, 522

Type locality

Mozambique: Prov. Inhambane, 15 km SE Save.

Type material

(5 exs.): Holotype, male: “Mocambique Prov. Inhambane 15 km SE Save, 18–21.12. 2005 A. Kurdna Jr. lgt.” (CFP; habitus in Fig. 384). – Paratypes, female: Same data as holotype (2 exs. CFP, 1 ex. MZH, 1 ex. NHRS).

Diagnosis

Absence of dorsal colour pattern (or sometimes presence of indistinct vague darker areas on body) in combination with peculiar, abrupt end of penis-apex, distinguishes L. ferrugo from the other species in this species group.

Description

Body length 4.2–4.7 mm, breadth 2.2–2.4 mm. Dorsal, colour pattern lacking or very indistinct and vaguely delimited (Fig. 384).

Head: Ferrugineous; frontally often narrowly and slightly paler; with a vague, pale ferrugineous border at foremargin. Impunctate, except at eyes; with scattered, irregular punctures. Slightly matt, reticulation, simple, meshes small and of equal size.

Pronotum: Ferrugineous, no distinct colour pattern. Impunctate, except at margins; with fine, irregular punctures, which are lacking medially at base. Slightly matt, with fine, simple reticulation; meshes small and of equal size.

Elytra: Ferrugineous to dark ferrugineous; sometimes with vague, dark ferrugineous to blackish areas (one at scutellar region, one extensive on medial part, and one apically). Dark areas, when present a very diffuse and their delimitation vague (Fig. 384). Submat, finely and equally microsculptured; meshes small and of equal size. Discally with a fine, irregular row of punctures. Dorsolateral row and lateral row of punctures sparse, irregular and especially lateral one is fragmentary. Apically with sparse and irregularly distributed punctures of variable size.

Ventral aspect: Ferrugineous, abdomen in part darker; ferrugineous to dark ferrugineous. Almost impunctate. Scattered punctures on apical ventrite which lacks small knob on one side (Fig. 20). Rather shiny, with very fine, in part, indistinct microsculpture. Apex of prosternal process quite slender, posteriorly moderately extended, apex narrows quite abruptly (approx. as Fig. 1). Metacoxal plates in frontal half provided with, fine, almost transversely located, shallow furrows. Abdomen with fine, curved striae. Metacoxal process ends abruptly; no posterior extensions (Fig. 6).

Legs: Pale ferrugineous to ferrugineous. Pro- and mesotarsus slightly enlarged, provided with suckers. Apex of metatibial spines bifid, although very finely so.

Male genitalia: Penis, lateral aspect, from approximately middle to apex evenly curved; tip somewhat enlarged, ends abruptly (Figs 222–223).

Female: Pro- and mesotarsus slender. Apical sternite (Fig. 21).

Etymology

The species name “ferrugo” is a Latin noun meaning rust (of iron) and relates to the body colour of the new species.

Distribution

Mozambique (Fig. 522).

Collecting circumstances

Unknown.

Species group 2 (L. ruficollis group)

Diagnosis. Quite small sized Laccophilus with length of body 3.2–3.4 mm and width 1.8 mm.

Shape of body oval; body dorsoventrally flattened (Fig. 380). Body dorsally, with distinct colour pattern, formed by two distinct, transverse, pale ferrugineous markings, which are broadly broken by dark suture; apex of elytra also pale coloured (Fig. 385). Body microsculpture on dorsal aspect double, of two kinds (in part meshes reduced).

Prosternal process moderately broad, posteriorly not distinctly extended, apically pointed. Apical sternite not distinctly modified; lacks asymmetrical, small knob (Fig. 22). No stridulatory apparatus on metacoxal plates. Metacoxal process not extended posteriorly (Fig. 6).

Paramere simple, elongate, apically not distinctly enlarged or modified (Fig. 224). Apical half of penis slender, distinctly curved but not distinctly modified (Fig. 224).

Species composition and distribution. One African species is recognized. Only known from Madagascar.

Laccophilus ruficollis Zimmermann, 1919

Figs 22–23, 224, 385, 527

Laccophilus ruficollis Zimmermann 1919: 123 (original description, faunistics); Zimmermann 1920a: 25 (catalogue); Nilsson 2001: 250 (catalogue, faunistics); Nilsson 2015: 217 (catalogue, faunistics).

Type locality

Madagascar.

Type material, studied

(3 exs.). Lectotype (by present designation): male: “Madagascar / Type / Samml. A. Zimmermann / Paratypus” (ZSM; habitus in Fig. 385). – Paralectotypes, male and female: Same data as in lectotype (2 exs. ZSM). [Comment: all three specimens studied are provided with a type and a paratype label and no holotype has been chosen. Two of the specimens are females and one, male. We have chosen the male to be the lectotype.]

Diagnosis

Laccophilus ruficollis is distinguished from all other African species by unmodified apical ventrite and exhibiting distinct, transverse, pale markings on elytra. Furthermore, penis apex is slender and curved and body-microsculpture is a mix of small and large meshes. In combination with small sized body these characters are useful when L. ruficollis is distinguished.

Description

Body length 3.2–3.4 mm, width 1.8 mm. Dorsal colour pattern rather uniform and distinct (Fig. 385).

Head: Pale ferrugineous. Rather shiny although finely microsculptured. Reticulation almost simple, predominantly of one kind. In part reticulation indistinctly double but small and large meshes difficult to distinguish and place in either category. Between eyes, with fine, sparse punctures. At eyes punctures slightly denser.

Pronotum: Ferrugineous, laterally pale ferrugineous (change of colour gradual). Basally with vague, transverse, dark ferrugineous marking. Rather shiny although finely microsculptured; reticulation mostly uniform: small and large meshes difficult to distinguish. Laterally and at anterior margin, finely punctate.

Elytra: Dark ferrugineous, with distinct pale ferrugineous markings (Fig. 385). Slightly mat, finely and densely microsculptured. Reticulation-meshes not clearly forming two distinct groups. Almost impunctate; laterally and at suture with fine punctures.

Ventral aspect: Pale ferrugineous to ferrugineous; lacks distinct colour pattern, but abdomen in part slightly darker. Rather shiny, very finely microsculptured (in part microsculpture hardly discernible). Scattered, curved striae discernible but sometimes rather indistinct. Almost impunctate. Apex of prosternal process comparatively short although pointed. Apical ventrite simple, not distinctly modified (Fig. 22).

Legs: Protarsus slightly extended and enlarged; mesotarsus long and slender. Provided with suckers.

Male genitalia: Rather delicate in size and exhibits hardly any modifications; penis in lateral aspect slender and evenly curved (Fig. 224).

Female: Externally almost as male. Protarsus slender. Apical ventrite as in Fig. 23.

Distribution

Madagascar (Fig. 527).

Collecting circumstances

Unknown.

Species group 3 (L. hyalinus group)

Diagnosis. Large species with body length 4.0–5.3 mm, and width 2.2–3.0 mm.

Shape of body oblong to oval, dorsoventrally flattened (Figs 386–387). Dorsal side unicoloured pale ferrugineous to ferrugineous, generally lacking colour pattern. Some species exhibit vague pattern; elytra ferrugineous with pale ferrugineous, often vague and variable patches (Fig. 389). Body microsculpture double; small and large meshes mixed, in part meshes often reduced and missing.

Prosternal process moderately slender, posteriorly not strongly extended, apex pointed. Apical ventrite not distinctly modified; lacks asymmetric knob on one side (Fig. 24). Two species have in both sexes stridulatory files on metacoxal plates (Fig. 6). Metacoxal process not extended posteriorly (Fig. 6).

Paramere simple, elongate, apically not distinctly enlarged or modified (Fig. 229). Apical half of penis slender, distinctly curved but not distinctly modified (Fig. 228). In L. hyalinus extreme apex of penis finely hooked (Fig. 225).

Species composition and distribution. Five species are recognized in this species group. In Africa they are distributed North of Sahara and most of them exhibit a wider distribution in the Palearctic region.

Key to species (males & females)

1 Metacoxal plates with stridulatory file (Fig. 6) 2
Metacoxal plates lack stridulatory file 3
2 Body shape oval-oblong, narrower (Fig. 386); penis as in Fig. 225 L. hyalinus (p. 28)
Body shape oval, broader (Fig. 387); (female only known) L. demoflysi (p. 31)
3. Smaller species, body length 4.0–4.6 mm; male genitalia slender (Fig. 226) L. minutus (p. 31)
Large species, body length 4.6–5.3 mm; male genitalia robust (Fig. 227) 4
4. Elytra with distinct colour pattern (Fig. 389); male genitalia (Fig. 227) L. mateui (p. 33)
Elytra lack colour pattern or pattern is indistinct (Fig. 390); male genitalia (Fig. 228–229) L. sordidus (p. 35)

Laccophilus hyalinus (De Geer, 1774)

Figs 6, 9–10, 24–25, 225, 386, 528

Dytiscus hyalinus De Geer 1774: 406 (original description, faunistics).

Laccophilus hyalinus (De Geer), Reiche 1872: 23 (faunistics, list.); Sharp 1882: 301 (description, faunistics); Zimmermann 1920a:19 (catalogue); Zimmermann 1930: 14 (description, faunistics); Normand 1938: 343 (catalogue); Guignot 1946b: 186 (discussion, faunistics); Omer-Cooper 1970: 286 (faunistics, discussion); El Alaoui 1983: 133, 135 (faunistics); Brancucci 1983b: 268 (faunistics); Nilsson 2001: 244 (catalogue, faunistics); Nilsson 2015: 212 (catalogue, faunistics).

Laccophilus testaceus Aubé, Reiche 1872: 23 (faunistics, list); Nilsson 2015: 212 (catalogue, faunistics, list synonymy).

Laccophilus interruptus var. testaceus Aubé, Severin 1892: 472 (type material deposition); Régimbart 1895: 132 (faunistics, L. interruptus = L. hyalinus, synonymy).

Laccophilus hyalinus var. testaceus Aubé, Bertrand 1928b: 46 (larva description); Guignot 1946b: 186 (faunistics, discussion).

Laccophilus hyalinus testaceus Aubé, Zimmermann 1920a:19 (catalogue); Lindberg 1939: 11, 13, 29 (biology, faunistics); Angelini 1982: 82 (faunistics); Nilsson 2003: 76 (faunistics, list); Bennas and Sàinz-Cantero 2006: 58, 62 (faunistics, list).

Laccophilus hyalinus ab. testaceus Aubé, Legros 1972: 467 (faunistics).

Laccophilus hyalinus var. inflatus Wollaston, Machado 1987: 50 (description, faunistics); Nilsson 2015: 212 (catalogue, faunistics, list synonymy).

Laccophilus hyalinus inflatus Wollaston, Balke et al. 1990: 361, 369, 370 (discussion, faunistics, biology).

Comment on synonymy

The present list of references is incomplete; selected references with association to Africa and Canary Islands are included. Synonymy of different taxa is based on earlier studies (see World Catalogue, Nilsson 2001, 2015).

Type locality

Sweden.

Type material, studied

(1 ex.). Laccophilus hyalinus: Syntype (unique?): (NHRS). [Comments: no original text labels attached with the specimen; specimen provided with orange label; severely damaged by dermestids.]

Type material, not studied

Laccophilus testaceus: “France, Italy, Spain” (in Brussels).

Additional African material studied

(310 exs.): Morocco: “Tanger 25–29.4. 1926 Lindberg” (7 exs. MZH); “Marrakesh 21–23.5. 1926 Lindberg” (5 exs. MZH); “Marrakech 28.5. 1934, 450 m Ball 20M79 / ab. testaceus” (1 ex. MNHN); “Atlas mai, Reraia 29.5–15.6. 1926 Lindberg” (6 exs. MZH); “Atlas mai., Amismiz 24–25.5. 1926 Lindberg” (1 ex. MZH); “M. Atlas Azrou 1200 m 15.3. 1961 Lindberg” (4 exs. MZH); “Foret de la Mamora 23.3.1961 Lindberg” (1 ex. MZH); “Tiflet-Oulmes 18.2.1961 Meinander” (1 ex. MZH); “Oued Tensift pr., Marrakesh 13.3. 1961 Lindberg” (1 ex. MZH); “nr. Figuig, Defilia 5–20.4. 1966/Hutson” (1 ex. BMNH); “Mar. bor. Tetouan 600 m 25.5. 1994 leg. Majzlan” (14 exs. NMW, 3 exs. MZH); “Ouazaza-Te 12.5. 1975 Eckerlein” (7 exs. NMW, 1 ex. MZH); “Oulad Teima pr. Oued Sous 14.2. 1961 Lindberg” (4 exs. MZH); “Mogador (Essaouira) 12.2. 1961 Meinander” (12 esx. MZH); “Tiferhlal N de Tiznit 18.2. 1961 Meinander” (6 exs. MZH); “Oued Massa N de Tiznit 18.2. 1961 Meinander” (3 exs. MZH); “Maroc sud Torkoz 23–24.2. 1961 Meinander” (7 exs. MZH); “Maroc sud Assa 22.2. 1961 Lindberg” (5 exs. MZH; habitus in Fig. 386). – Algeria: “55 km N of Tamanrasset 16–17.3. 1971 Gruwell leg.” (90 exs. USNM, 10 exs. MZH); “Bouqie 17.7. 1955 Lauck leg.” (1 ex. USNM). – Tunisia: “Totzeur 5.4. 1924 Lindberg” (1 ex. MZH); “Ichkeulsee 1.8. 1991 Schödl” (2 exs. NMW); “Djeb. Ressas / J. Sahlb.” (1 ex. MZH). – Libya: “Libya bor. occ. 653 asl, prov. Yafran 9.5.2002 Ain Az-Zarqa (W of Jadu) / 31°57'21.2"N, 12°00'25.9"E, Reiter A. lgt.” (27 exs. NMPC); “Libya Darnah prov. Wadi Darnah, 117 m, 32°42'06.4"N, 22°36'39.9"E, A. Reiter leg. 15.5. 2002” (17 exx. NMPC); “Libya bor. occ. 605 m asl, prov. Yafran-Ghadamis Nana tala, 10 km W Ar Rhaibat 27.5.2002 / 31°47'09"N, 11°47'07.9” Reiter A. lgt.” (3 exs. NMPC); “Libya bor. occ. 336 m asl, prov. Tarhunah 26.5.2002 Ain Sharshara, 3 km N Tarnuah / 32°27'57.7"N, 13°37'04.7” E. Reiter A lgt” (25 exs. NMPC). – Egypt: “Wadi Kujib 12.6. 1994 Ullrich / L. hyalinus Deg. Hendrich det. 1994” (24 exs. USNM); “Madiba Wala 20.6. 1994 Ullrich / L. hyalinus Deg. Hendrich det. 1994” (4 exs. USNM). – Canary Islands (Spain): “Tenerife, Valle de Masca 12–13.5. 1947 Lindberg” (1 ex. MZH); “Tenerife, Bco Bufadero 10.8. 1949 Fernandez” (4 exs. MZH); “Gran Canaria, Aldea S. Nicolas 1.3. 1949 Lindberg” (9 exs. MZH); “Gran Canaria, Maspalomas 9-10.3. 1950 Lindberg” (1 ex. MZH).

Diagnosis

Laccophilus hyalinus resembles most among African species of L. demoflysi, which also has a similar stridulation apparatus as L. hyalinus. For separation of the two species, see diagnosis of L. demoflysi (p. 31). Stridulation apparatus located on metacoxal plates distinguishes L. hyalinus from L. minutus, L. mateui and L. sordidus, all of which lack similar device. Clear differences between the species are also exhibited in shape of the penis.

Description

Body length 4.7–5.0 mm, width 2.7–2.8 mm. Habitus generally with somewhat paler, longitudinal markings, which often are rather vague, in part indistinct (Fig. 386).

Head: Pale ferrugineous. At eyes, in shallow depression with fine punctures. In short horizontal depressions close to eyes and in connection with shallow depression, with fine punctures. Reticulation double, large meshes contain 2–6 fine meshes. Fine meshes in part weakly developed and indistinct.

Pronotum: Pale ferrugineous. At foremargin and medially with slightly darker areas (dark areas not on surface but “inside” cuticula). Almost impunctate, with a few irregular punctures at frontal margin. Rather finely microsculptured. Reticulation double. Fine meshes in part largely absent, in part very fine to fine. Large meshes, when discernible, contain 4–6 fine meshes.

Elytra: Pale ferrugineous to pale brown, with slightly paler, in part indistinct markings (Fig. 386). Elytral punctation almost absent; discally, dorsolaterally and laterally with a few, very fine, hardly visible punctures placed in vague rows. Slightly mat due to microsculpture. Reticulation double, large meshes contain 2–6 small meshes.

Ventral aspect: Pale ferrugineous. Impunctate. Slightly mat, due to dense, fine microsculpture. Metacoxal plates with about 10 transverse, shallow furrows. Basal segments of abdomen with curved striae. Pronotal process medially slightly enlarged, apex pointed (not strongly extended and sharp). Stridulation apparatus consist of a curved series of about 20 shallow ridges located posteriorly on metacoxa (Fig. 6). Apical ventrite with a shallow depression on one side (Fig. 22).

Legs: Pro- and mesotarsus slightly enlarged, provided with distinct suckers (Fig. 10).

Male genitalia: Extreme apex of penis in lateral aspect strongly curved upwards (Fig. 225).

Female: Provided with similar stridulation apparatus as male. Apical ventrite with uneven surface (Fig. 25). Pro- and mesotarsus rather slender.

Distribution

Morocco, Algeria, Tunisia, Libya, Egypt, Canary Islands (Fig. 528). Additional, African country record is Senegal (Legros 1972).

Collecting circumstances

Information from Africa is rare. In Europe often found in large bodies of running water, in sections with a slow current and some vegetation. Less frequently collected in ponds and lakes (Nilsson and Holmen 1995).

Laccophilus demoflysi Normand, 1938

Figs 26, 387, 529

Laccophilus demoflysi Normand 1938: 343 (original description, faunistics); Nilsson 2001: 242 (catalogue, faunistics); Nilsson 2003a: 76 (faunistics, list); Nilsson 2015: 211 (catalogue, faunistics).

Type locality

Tunisia: El Hamma de Tozeur.

Type material

(not studied): Holotype: female: “El Hamma de Tozeur, 4. 1937 Demoflys” (Coll. Normand, kept in Tunisia, Institut National Agronomique de Tunisie, Tunis, specimen not located).

Material studied

(1 ex.): Tunisia: “Tunisia, centralis oasis Douz env. 31.5.-1.6. 1994 lgt.S. Becvar / Laccophilus demoflysi Norm.det. Rocchi 1998” (1 ex. female CSR; habitus in Fig. 387).

Diagnosis

External characters agree in large with L. hyalinus. Only difference observed was the shape of the body, in L. demoflysi being stouter than in L. hyalinus. Both involved species have a stridulatory apparatus on metacoxal plates, which separates them from L. minutus, L. mateui and L. sordidus. Taxonomic status of L. demoflysi remains open. More specimens (male in particular) are needed to settle this question.

Description

(only differences from description of L. hyalinus are recognized). Body length 4.7 mm, width 2.8 mm. Dorsal colour pattern vague, almost absent (Fig. 387).

Ventral aspect: Apical ventrite (Fig. 26).

Male: Unknown.

Distribution

Tunisia (Fig. 529).

Collecting circumstances

Unknown.

Laccophilus minutus (Linnaeus, 1758)

Figs 27–28, 226, 388, 529

Dytiscus minutus Linnaeus 1758: 412 (original description, faunistics).

Laccophilus minutus (Linnaeus), Leach 1815: 673 (description, biology); Aubé 1838: 417 (description, faunistics); Lucas 1846: 94 (faunistics); Reiche 1872: 23 (faunistics, list); Régimbart 1895: 132 (faunistics; given as L. obscurus Panzer, junior synonym of L. minutus); Zimmermann 1920a: 21 (catalogue); Bertrand 1928b: 274 (juvenile description); Zimmermann 1930: 14 (description, faunistics); Lindberg 1939: 11, 13, 29 (biology, faunistics); Brinck 1943: 154 (faunistics); Balfour-Browne 1951: 193 (discussion); Sanfilippo 1955: 1 (faunistics, biology); Guignot 1956b: 220 (discussion); Guignot 1959a: 579, 583 (redescription, faunistics); Angelini 1982: 82 (faunistics); El Alaoui 1983: 133, 135 (faunistics); Nilsson 2001: 247 (catalogue, faunistics); Nilsson 2003: 76 (faunistics, list); Bennas and Sàinz-Cantero 2006: 59, 62 (faunistics, list); Nilsson 2015: 214 (catalogue, faunistics). [Comments: Only references referring to Africa are listed. The list of synonyms and references is accordingly, not complete. We refer to World catalogue of Nilsson (2015)]

Type locality

Europe.

Type material

(not examined): “Europe”. Type specimen deposited in The Linnean Collections, London, UK (item data: LINN 6347 Dytiscus minutus (Ins. Linn.), www.linnean.org).

African material studied

(49 exs.). Morocco: “Tanger 25-29.4. 1926 Lindberg” (6 exs. MZH); “Tiflet-Oulmes 18.3.1961 Lindberg-Meinander” (4 exs. MZH; habitus in Fig. 388); “Sp. Mor., at Tangier border 8.7.1955” (1 ex. USNM); “Rabat 3–4.5. 1926 Lindberg” (2 exs. MZH); “Gharb 7.7. 1926 Lindberg” (11 exs. MZH); “Marrakesch 21–23.5. 1926 Lindberg” (6 exs. MZH); “Dayet Jerans (lake) 11 km E.N.E. Ifrane 5400 ft. 28.5. 1961 / P.N. Lawrence” (1 ex. BMNH); “Middle Atlas, nr Ifrane 28.5. 1961 Dayet Jerane P.N. Lawrence” (1 ex. BMNH, 1 ex. MZH); “Oued Zad 65 km S Ifrane 21.5. 1961 / P. N. Lawrence / Tiny pond, flooded grass” (1 ex. BMNH); “Fr. Mor., Petit Jean 9.7. 1955 D.L. Lauck” (6 exs. USNM, 2 exs. MZH); “Sp. Mor., El Kaar, El Kebir 8.7.1955 D.R. Lauck” (2 exs. USNM). – Algeria: “Bouqie 17.7. 1955 D.R. Lauck” (1 ex. USNM). – Tunisia: “Tunis J. Sahlb.” (1 ex. MZH); “Tunisien 2.9. 1991 5 km W Utique Schödl” (1 ex. NMW); “Tunisien 3.8. 1991, 22 km N Jendouba Schödl” (1 ex. NMW). – Libya: “Libya bor. or. 495 m a.s.l. prov. Al Jabal Al Akhdar, 5 km SW Al Bayda 18.5.2002 / 32°43'41.9"N, 21°41'14"E, Reiter A. lgt.” (1 ex. NMPC). [Comment: Only material collected from Africa is listed.]

Diagnosis

Laccophilus minutus resembles most of L. mateui and L. sordidus, both species also lacking stridulation apparatus on metacoxal plates. L. minutus is separated from the two close species by having smaller body and more delicate and slender penis (Figs 226, 227, 228).

Description

Body length 4.0–4.6 mm, width 2.2–2.5 mm. Dorsal, aspect of body without distinct colour pattern. On elytra vague, slightly darker areas may sometimes be discerned (Fig. 388).

Head: Pale ferrugineous. Slightly mat to rather shiny, finely microsculptured. Reticulation double. Large meshes slightly more strongly developed than fine meshes. Large meshes may contain 3–6 small meshes. Impunctate, except at inner eye-margin, with an irregular row of punctures. Additionally close to eyes with a vertically located row of a few punctures.

Pronotum: Pale ferrugineous; lacks distinct colour pattern. Impunctate, except at frontal and lateral margins, where fine and sparse, irregular punctures are discernible. Rather shiny to submat, finely microsculptured. Reticulation double. Large meshes a little more strongly developed than small meshes; may contain 3–6 small meshes. Pronotum base posteriorly in middle produced backwards.

Elytra: Pale ferrugineous, without distinct colour pattern. Sometimes elytra with some vague, irregular, slightly darker areas (Fig. 388). Almost impunctate. Discally with sparse, fine and scattered punctures forming a vague row. Additionally, some scattered, fine punctures may be discerned at location of dorsolateral and lateral rows. Pre-apical, lateral row of punctures form a fine furrow, provided with fine hairs. Rather shiny, although finely microsculptured. Reticulation double. Large meshes a little more strongly developed; may contain 3-6 smaller meshes.

Ventral aspect: Pale ferrugineous to ferrugineous, without distinct colour pattern. Almost impunctate, Abdomen apically with some fine punctures. Rather shiny, very finely and in part indistinctly microsculptured. Ventrites with fine, slightly curved striae. Metacoxal plates with about 10 fine, shallow, transversely located furrows. No stridulatory apparatus. Apical ventrite almost symmetric, lacks lateral knob; finely striated, with distinct lateral impressions (Fig. 27). Prosternal process rather slender; posteriorly, moderately extended, apex pointed.

Legs: Pale ferrugineous to ferrugineous. Pro- and mesotarsus somewhat enlarged, provided with suckers.

Male genitalia: Aedeagus almost as in L. mateui but more delicate in lateral aspect (Fig. 226).

Female: Pro- and mesotarsus slender. Apical ventrite lacks distinct lateral impressions; impressions reduced to fine lines (Fig. 28).

Distribution

Morocco, Algeria, Tunisia, Libya (Fig. 529).

Collecting circumstances

According to Nilsson and Holmen (1995), probably focusing from Nordic perspective, main habitat is permanent water body as lakes and ponds with stagnant water. Vegetation of water body is sparse or absent. Lindberg (1939) briefly described some sampling sites of L. minutus in Morocco. The species was collected both in standing and running waters. Collected also from an almost dried up river-bed with rich vegetation. Adults are capable of flight.

Laccophilus mateui Omer-Cooper, 1970

Figs 29–30, 227, 389, 529

Laccophilus mateui Omer-Cooper 1970: 285, 287 (original description, faunistics); Nilsson 2001: 246 (catalogue, faunistics); Nilsson 2003: 76 (catalogue, faunistics); Nilsson 2015: 214 (catalogue, faunistics).

Type locality

Algeria: Sahara, Hoggar, Aguelm, Ymeleulauen.

Type material, studied

(1 ex.). Holotype: male: “Type male / H. B. Leech Collection / det. J. Omer-Cooper Laccophilus mateui sp.n. / Aguelm, Ymeleulauen, Hoggar, Sahara J. Mateu coll. / 18-V- 1951” (AMGS; according to original description, holotype to be deposited in CAS).

Additional material, studied

(5 exs.). Algeria: “55 km N Tamanrasset 16-17 March 1971 J.A. Gruwell” (4 exs. USNM, 1 ex. MZH; habitus in Fig. 389).

Diagnosis

Laccophilus mateui is a close relative to L. minutus and L. sordidus. All three species have similar general appearance and same ground plan regarding penis-shape. Absence of stridulatory apparatus separates it from L. hyalinus and L. demoflysi. Shape of penis distinguishes it from L. minutus (penis apex is broader in L. mateui) and L. sordidus (penis is stouter in L. mateui and longer in L. sordidus). L. mateui (>5 mm) is also larger sized than L. minutus, a species which don’t exceed 5 mm in length.

Description

Body length 5.1–5.3 mm, width 2.8–3.0 mm. Body dorsally pale ferrugineous to ferrugineous, elytral colour pattern vague to fairly distinct (Fig. 389).

Head: Pale ferrugineous, at pronotum darker, ferrugineous to dark ferrugineous (delimitation of colours often vague). Almost impunctate, at eyes with a few, scattered, somewhat indistinct punctures. Submat, finely microsculptured; reticulation double. In central part of head fine reticulation indistinct, in part obliterated; in lateral parts of head fine reticulation clearly discernible; large meshes contain 3–6 fine meshes.

Pronotum: Ferrugineous, laterally pale ferrugineous (gradual change; no distinct delimitation of colours). Frontally, sometimes with a quite distinct dark ferrugineous area. Impunctate, at margin with a few, indistinct, coarser punctures discernible. Submat, rather densely microsculptured; reticulation double, fine meshes in part almost absent or indistinct.

Elytra: Ferrugineous, with vague pale ferrugineous markings (Fig. 389). Elytral colour pattern sometimes quite distinct. Submat, rather finely microsculptured; reticulation double, large meshes contain between 2–6 fine meshes. Each elytron with a discal, dorsolateral and lateral row of punctures, which are sparse and somewhat irregular, in part indistinct. Lateral, pre-apical furrow long, shallow, finely punctate and pubescent.

Ventral aspect: Ferrugineous, metacoxal plate in part dark ferrugineous but no distinct colour pattern formed (delimitation vague). Almost impunctate. Submat, finely microsculptured. No stridulatory apparatus. Metacoxal plates in anterior half with some vague, transversely located, slightly irregular furrows. Abdomen in basal half with rather distinct curved striae. Apex of prosternal process broken in holotype; it is keeled, short and apex pointed. Apical ventrite as in Fig. 29.

Legs: Pro- and mesotarsus somewhat enlarged, provided with suckers.

Male genitalia: Penis comparatively robust; in lateral aspect apical half evenly curved (Fig. 227).

Female: Pro- and mesotarsus slender. Apical ventrite as in Fig. 30.

Distribution

Algeria (Hoggar, Sahara) (Fig. 529).

Collecting circumstances

Unknown.

Laccophilus sordidus Sharp, 1882

Figs 31–32, 228–229, 390, 529

Laccophilus sordidus Sharp 1882: 302 (original description, faunistics); Zimmermann 1920a: 26 (catalogue); Balfour-Browne 1951: 193 (faunistics, discussion, description); Brancucci 1980: 107 (description, faunistics, lectotype designation); Brancucci 1983b: 264, 266 (redescription, faunistics, discussion); Zalat et al. 2000: 39, 40 (description, faunistics, biology); Nilsson 2001: 251 (catalogue, faunistics); Nilsson 2003: 77 (catalogue faunistics, ); Shaverdo et al. 2013: 21, 22 (faunistics); Nilsson 2015: 218 (catalogue, faunistics).

Type locality

Saudi Arabia: El Hedjaz.

Type material, studied

(2 exs.). Lectotype: male, designated by Brancucci (1980a): “Syntype / Type / Hedjaz Millingen / Sharp Coll. 1905-313 / L. sordidus / Laccophilus sordidus Sharp Type (male symbol)” (BMNH; habitus in Fig. 390). – Paralectotype, female: “Hedjaz Millingen / Sharp Coll. 1905-313 / Laccophilus sordidus Sharp Paratype (female symbol) / Syntype” (1 ex. BMNH).

Additional material studied

(1 ex.): Yemen: “Aden Prot., Mukeiras, 85 mls NE of Aden, 7000 ft. 29.12. 1939-6.1. 1940 Hebbert / L. sordidus Shp det J. Balfour-Browne” (1 ex. BMNH).

Diagnosis

Laccophilus sordidus is closely related to L. minutus and L. mateui. From L. minutus, L. sordidus is separated by its larger body and by having a clearly longer penis. From L. mateui, L. sordidus is separated by its lack of elytral markings – L. mateui is generally provided with a clearly discernible elytral colour pattern. Additionally, the penis of L. mateui is distinctly shorter than in L. sordidus.

Description

Body length 4.6–5.2 mm, width 2.6–2.8 mm. Dorsal, aspect of body dark ferrugineous to brownish, no distinct colour pattern exhibited (Fig. 390).

Head: Dark ferrugineous to brownish; no colour pattern discernible. Submat to mat, distinctly microsculptured; reticulation double. Large meshes may contain 3–6 small meshes. At eyes with an irregular row of punctures, which extends a short distance towards middle.

Pronotum: Dark ferrugineous to brownish, no colour pattern formed. Submat, distinctly microsculptured. Reticulation double. Large meshes contain 3–6 small meshes. Pronotal disc impunctate; at margins with punctures. Laterally at side margin, punctures form a slightly irregular row. Anteriorly punctures very fine and scattered. Latero-basally with a few fine, irregular punctures; mediobasally pronotum impunctate.

Elytra: Dark ferrugineous to brownish. No distinct colour pattern exhibited (Fig. 390). Submat, distinctly microsculptured. Reticulation double; large meshes contain generally 3–6 small meshes. Three somewhat irregular rows of punctures formed. Discal row is rather distinct while dorsolateral and lateral rows are quite vague and punctures appear more scattered. Posteriorly lateral row become more condensed and it is located in a shallow pre-apical furrow.

Ventral aspect: Dark ferrugineous to ferrugineous, no distinct colour pattern. Submat, finely microsculptured. Microsculpture of abdomen weaker and in part reduced. Abdomen with fine, curved striae. Metacoxal plates lack stridulatory apparatus. Very shallow, rudimentary transverse furrows discernible on metacoxa. Impunctate, except abdomen with scattered, sparse punctures especially apically. Prosternal process medially slightly enlarged, moderately extended and apex pointed. Apical ventrite almost symmetric (Fig. 31).

Legs: Ferrugineous to dark ferrugineous. Pro- and mesotarsus slightly enlarged and provided with suckers.

Male genitalia: In lateral aspect penis comparatively long, evenly curved; extreme apex slightly bent and it ends abruptly (not rounded) (Figs 228–229).

Female: Pro- and mesotarsus slender. Apical ventrite as in Fig. 32.

Distribution

The species has been described from Saudi Arabia. African records include Libya and Egypt (Zalat et al. 2000). Thus far we have not seen any specimens from Africa, but a few from Arabian Peninsula (Fig. 529).

Collecting circumstances

The habitat of L. sordidus is briefly described in Zalat et al. (2000) as follows “The species occurs in shallow water pools with gravel bottom and sparse vegetation, the water either being fresh or brackish. Considered rare and occurs in Eastern desert of Egypt in August”.

Species group 4 (L. alluaudi group)

Diagnosis. Medium sized to large species; length of body 3.3–5.8 mm, width 1.8–2.9 mm.

Shape of body, oval-oblong to oblong; body dorso-ventrally flattened (Fig. 393). Body dorsally, with distinct colour pattern. Elytra exhibit somewhat broad, dark, longitudinal markings, which in most species fade away before reaching humeral region. Pale stripes between dark markings in most species shaped as small pearls on a narrow string (Fig. 391). One species exhibits hollow, dark elytral markings, i.e. dark, longitudinal marking encloses a narrow, pale area (Fig. 396). Body microsculpture double; two different size-classes can be recognized. Large meshes often in part reduced. One species with meshes in part longitudinally extended.

Prosternal process rather slender, moderately backwards extended, apically pointed. One species with comparatively short prosternal process. Apical ventrite not distinctly modified, lack asymmetric knob (Fig. 33). No stridulation apparatus on metacoxal plates. Metacoxal process not extended posteriorly (Fig. 6). One paramere simple, apically not distinctly modified or enlarged (Fig. 232). One species with slightly modified paramere. Penis slender, lateral aspect, evenly curved or basally angled and quite straight. Appearance of penis quite simple and delicate, lack considerable modifications (e.g. Figs 230–231).

Species composition and distribution. Six species are recognized in this species group, all of which occur on Madagascar or on nearby islands.

Key to species (males only)

1 Large species, body length 5.0–5.8 mm; dark elytral, longitudinal markings hollow (narrow, pale marking enclosed in dark marking) (Fig. 396) L. seyrigi (p. 46)
Small to medium sized species, body length 3.3–4.2 mm; dark elytral marking entirely dark (no enclosed medial pale marking in dark marking) (Fig. 391) 2
2 Elytral dark markings complete or almost complete, reach humeral region (Fig. 391) L. comes (p. 37)
Elytral dark markings fade away before humeral region (Fig. 393) 3
3. Penis, lateral aspect, evenly curved from base to apex (Fig. 273); penis, dorsal aspect, near base on right-hand side with a distinct enlargement (Fig. 238) L. tigrinus (p. 43)
- Penis, lateral aspect, close to base angled (Fig. 230); penis, dorsal aspect, lacks latero-basal enlargement (Fig. 231) 4
4. Large species, body length 4.1–4.2 mm; penis as in Fig. 242–243 L. pseustes (p. 44)
- Small species, body length 3.4-3.9 mm; penis different 5
5. Penis, dorsal aspect, from angle to apex almost evenly broad and almost straight (Fig. 236) L. furthi (p. 42)
- Penis, dorsal aspect, from angle to apex broad to narrow, strongly twisted (Fig. 234) L. alluaudi (p. 39)

Laccophilus comes Guignot, 1955

Figs 33–34, 230–232, 391, 530

Laccophilus comes Guignot 1955f: 141 (original description, faunistics); Rocchi 1991: 86 (faunistics, list); Nilsson 2001: 242 (catalogue, faunistics); Nilsson 2015: 210 (catalogue, faunistics).

Type locality

Madagascar, Tampolo.

Type material, studied

(2 exs.). Holotype: male: “Madagascar Tampolo VIII. 1949 / male symbol / Type” (MNHN). – Allotype (= Paratype), female: “Madagascar Tampolo VIII. 1949 / female symbol / Allotype” (1 ex. MNHN; habitus in Fig. 391).

Additional material studied

(116 exs.): Madagascar: “Andasibe 11.12. 2004 Lat -18.943 Lon 48.4063, Balke & Monaghan / DNA Voucher BMNH <670655> MSL008:B01 / L. comes Bergsten det.” (1 ex. NHRS); same data but “BMNH<670654>” (1 ex. NHRS); “Toam, Ambatondrazaka, Zahamane, Zahamena NP river, P60BI04: N: E: m, 29.12. 2006 leg. Isambert et al / DNA Voucher BMNH <830741> MSL399:F2 / L. comes Bergsten det.” (1 ex. NHRS); same data but “BMNH <830743> MSL399:F4” (1 ex. NHRS); “Toam, Ambatondrazaka, Zahamane, Zahamena NP stream, P60BI15: N: -17.52 E: 48.721: 1075m / m, 29.12. 2006 leg. Isambert et al / DNA Voucher BMNH <830737> MSL399:E10 / L. comes Bergsten det.” (1 ex. NHRS); same data but “Analamaintsoa 3rd stream before Camp 1, stream of pools almost dry P60BI08, 30.12. 2006 N-17,50500, E48,72450, 1054 m Isambert et al (1 ex. NHRS); same data but “BMNH <830748> MSL399:F)/P60BI12: N: -17.517 E: 48.72: 1075 m, 31.12. 2006” (1 ex. NHRS); same data but “Zahamena NP 1st stream btw. Camp 1 and 2, stream of pools, 31.12. 2006, N-17,51733, 48,72067, 1075 m” (85 exx. NHRS, 2 exs. MZH); same data but “Analamaintsoa Forest 4th stream btw Camp 1 and 2, P6OBI15, 31.12. 2006, N-17,52050, E48,721337, 1075 m” (14 exx. NHRS); “IF Anadiana: Sahamalaotra Ranomafana NP: small stream P27MD31, N -21.2359 E: 47.3963, alt. 1123 m, 6.12.2004 leg. Balke et al / BMNH(E) <794196> DNA Voucher” (1 ex. NHRS); same but” BMNH(E) <794198>” (1 ex. NHRS); same but” BMNH(E) <794197>” (1 ex. NHRS); Mahajanga melaky, Tsingy de Bemaraha NP, S19.03419, E044.77499, 41 m.a.o., 15.12. 2009, Water net, field, Bergsten et al. / 000000464 NHRS-JLKB” (1 ex. NHRS); same but “S19.03572, E044.77507, 66 m.a.o., 15.12. 2009 / 000000467 NHRS/JLKB” (1 ex. NHRS); same but “S18.75643, E044.71398, 119 m.a.o., 17.12. 2009 / 000000463 NHRS/JLKB” (1 ex. NHRS); “Ampasimpotsy Moramanga Antsabe 11.12. 2004 N-18,94300, E48,40630, 979 m, Balke et al” (2 exx. NHRS).

Diagnosis

Laccophilus comes forms together with L. alluaudi, L. tigrinus, L. pseustes and L. furthi a distinct group characterized by similar colour pattern of body and male genitalia. Laccophilus comes, L. furthi and L. alluadi are smaller than the other species in the group. The three species are separated by small differences in shape of penis; see diagnosis of L. alluaudi on p. 41 and L. furthi on p. 42.

Description

Body length 3.3–3.9 mm, width 1.8–2.2 mm. Dorsal, colour pattern distinct and uniform (Fig. 391).

Head: Pale ferrugineous. At eyes with fine, irregularly distributed punctures. Rather shiny, although microsculptured. Reticulation double; coarse meshes only slightly stronger developed than fine meshes. Coarse meshes, when discernible, contain 2-3 fine meshes. In part, mesh categories cannot be distinguished.

Pronotum: Pale ferrugineous. Impunctate, except anteriorly, with scattered, fine punctures. Rather shiny, although finely microsculptured. Reticulation double. Coarse meshes only slightly more strongly developed than fine meshes; contain when discernible 3-4 fine meshes.

Elytra: Pale ferrugineous, with distinct, blackish ferrugineous to dark ferrugineous, longitudinal markings (Fig. 391). In a few specimens dark, longitudinal markings close to humeral region strongly reduced as in L. alluaudi. Posteriorly with rather fine, scattered punctation. Submat, distinctly and densely microsculptured. In part, reticulation double. Two kinds of reticulation clearly visible at scutellar region; posteriorly and laterally meshes of reticulation approximately one kind.

Ventral aspect: Metathorax and -coxal plates dark ferrugineous to ferrugineous; otherwise pale ferrugineous. Very fine, sparse punctures discernible on metacoxal plates and abdomen. Slightly mat due to very fine and hardly discernible microsculpture. Prosternal process rather slender, apex pointed but not strongly extended backwards. Abdomen with fine, sparse, curved striae. Apical ventrite not distinctly modified (Fig. 33).

Legs: Pro- and mesotarsus slender, somewhat extended; provided with suckers.

Male genitalia: Penis in lateral aspect medially, straight for a long distance, apex slightly bent; in dorsal aspect, penis from middle, strongly bent right (Figs 230–232).

Female: Pro- and mesotarsus slender. Apex of apical ventrite more angular than in male (Fig. 34).

Distribution

Madagascar (Fig. 530).

Collecting circumstances

Label data indicate that L. comes has been sampled in various sized, running waters as streams and rivers.

Laccophilus alluaudi Régimbart, 1900

Figs 35–36, 233–234, 392, 531

Laccophilus alluaudi Régimbart 1900: 373 (original description, faunistics); Zimmermann 1920a:16 (catalogue); Guignot 1955d: 67 (discussion); Guignot 1955f: 141 (faunistics, discussion); Guignot 1959a: 544, 548 (description, faunistics); Guignot 1959c: 76, 77, 78, 79 (discussion, faunistics); Guignot 1959e: 72 (faunistics); Guignot 1961a: 931 (faunistics); Bertrand and Legros 1971: 244 (faunistics, biology); Wewalka 1980: 724, 726 (faunistics, discussion); Rocchi 1991: 79, 86 (faunistics); Nilsson 2001: 240 (catalogue, faunistics); Pederzani and Rocchi 2009: 95 (faunistics, list); Nilsson 2015: 208 (catalogue, faunistics).

Type locality

Madagascar: Diego Suarez.

Type material, studied

(5 exs). Lectotype (by present designation): male: “Madagascar Diego-Suarez Ch. Alluaud 1893 / male symbol / Cotype / Museum Paris col. Guignot” (MNHN). – Paralectotypes: Same data as in lectotype, but two of the specimens with female symbol (4 exx. MNHN; habitus in Fig. 392).

Additional material studied

(189 exs.): Madagascar: “Forét d’Ambre Lat -12.4754 Lon 49.2173 coll. Balke & Monaghan, BMNH(E)670568_MSL007: B10, 19.11.04 / L. alluaudi Régb. det. Bergsten” (1 ex. NHRS); same data but “BMNH(E)670572_MSL007: CO2” (1 ex. NHRS); “Antsiranana II Mt d’Ambre, Grande cascade stream parallel to GC in deep very steep gorge, mostly isolated pools 17.11. 2004, N: -12,49920 E: 49,17600, 800 m Balke et al” (29 exs. NHRS, 4 exx. MZH); “Antsiranana II Foret d’Ambre, small water hole in dry streambed, gardenland at edge of dry forest 19.11. 2004 N-12,47540, E49,21730, 545m leg. Balke” (2 exs. NHRS); “Antsiranana 1 Mtd French Streampool 12.11. 2004 N-12,33360, E49,35350, 171 m, leg Balke et al” (12 exs. NHRS); “Antsiranana 1 Mt.d’Ambre 16.11. 2004, N-12,52830, E49,17253, 1020 m Balke et al.” (1 ex. NHRS); “Toli, NW Ft Dauphin, forest, watersource, P54E: N: E:: m 19.5. 2006 Bergsten et al. / L. alluaudi det. Bergsten/BMNH(E): <?94192> DNA voucher / L. alluaudi det. Bergsten” (1 ex. NHRS); “Toli, NW Ft. Dauphin, creek with gravel, stones and sand in rainforest along the creek, in small water holes and (Madaglymbus) in waterpool on a large rock with wood and leaves 19.5. 2006 N-24°45.583, E46°51.821, 300 m Bergsten et al.” (2 exs. NHRS); “Ambilobe, Anjiabe Ambony Antsabe: Galoko Mts, hygropetric cascade; alt. 50 m, P2506M N -13.6093 E 48.7212, 23.11. 2004 leg. Monaghan, Andriamparany, Balke / BMNH(E) <794160> DNA voucher / L. alluaudi det. Bergsten” (1 ex. NHRS); “Anjiabe Ambony, Ambilobe Antsabe stairways-like cascade with vertical steps, exposed, extremely hot day 23.11. 2004 N-13,60930, E48,72120, 303 m Balke et al.” (17 exx. NHRS); “Ambilobe, Anjiabe Ambony Antsabe: waterhole in streambed on clearing: 50 m, P25MD11 N - 13.648 E 48.721, 21.11. 2004, leg. Balke et al/BMNH(E) <794189> DNA voucher / L. alluaudi det. Bergsten” (1 ex. NHRS); “Toli, Taolanaro; Isaky Ivondro, Foret Manangotry, running water, P67B: N: -24.859: E: 46.862: 310 m, 9.4. 2007 leg. Ranarilalatiana et al / DNA voucher BMNH <830767> MSL399:H4 / L. alluaudi det. Bergsten” (1 ex. NHRS); “Ants, Sambava, Marojejy, Marojejy NP: Forest stream P57BI01: N: -14.437: E: 49.773: 464 m, 6.12. 2006 leg. Isambert et al. / DNA voucher BMNH <830690> MSL399:A11 / L. alluaudi det. Bergsten” (1 ex. NHRS); “Toam, Ambatondrazaka, Zahamena; Zahamena NP: Stream P60BI08 N: -17.505: E: 48.724: 1054 m, 30.12.2006 leg. Isambert et al. / DNA voucher BMNH <830696> MSL399:B5 / L. alluaudi det. Bergsten” (1 ex. NHRS); same data but “P60BI06 N: -17.508: E: 48.724: 1068 m” and “BMNH <830701> MSL399:B10” (1 ex. NHRS); same data but “Zahamena NP, Analamaintsoa 1st stream before Camp 1 within a few m from the GPS point P60BI06, 30.12. 2006 N-17,50617, E48,72400, 1068 m” (4 exs. NHRS); “TOAM Ambatondrazaka Zahamena NP on the way to camp 2 to Fenerive Est “Route des contrebandiers” P60BI29 02.1. 2007, N: -17,54167 E: 48,72183, 1322m leg. Isambert et al” (37 exs. NHRS); same data but “Analamaintsoa, 2nd stream btw Camp 1 and 2, stream of pools 31.12. 2006 N-17,51850, E48,72217, 1075 m” (4 exs. NHRS); same data but “Analamaintsoa, 3rd stream before Camp 1, stream of pools, almost dry P60BI08, 30.12. 2006, N-17,50500, E48,72450, 1054 m Isambert et al” (10 exx. NHRS); “Montagne des Francais, Lat. -12.3336 Lon. 49.3535 leg. Balke & Otke, BMNH(E)_671210 MSL014; 1/1/1904 / DNA voucher BMNH <671210> MSL014: H02/L. alluaudi det. Bergsten” (1 ex. NHRS); “Saratanana, leg. Lees & Ranaivosolo MNH(E)_672835_MSL028; 1/1/1904 /DNA voucher BMNH <672835> MSL028: B05 / L. alluaudi det. Bergsten” (1 ex. NHRS); “Antsabe Lat -13.6093 Lon 49.7212, Balke leg. BMNH(E) _670700_MSL008 23.9. 2004/1904/DNA voucher BMNH <670700> MSL008: E10/L. alluaudi det. Bergsten” (1 ex. NHRS); “Ants, Nosy Be: Lokobe R.N.1, 50 m, 2004/DNA voucher BMNH(E) <794171> / L. alluaudi det. Bergsten” (1 ex. NHRS); “R.N.I. Lokobe Nosy Be, 50 m, 15.12. 2004 Ravo” (1 ex. NHRS); “Mahajanga: Boeny Mahavavy Kinkony RS. S16.05648, E045.76371, 55 m.a.o., 5.12. 2009 water net, field, Bergsten et al. (19 exs. NHRS); same data, add: “000000465 NHRS-JLKB” (1 ex. NHRS); “Mahajanga: Melaky: Tsingy de Bemaraha NP, S19.03572, E044.77507, 66m m.a.o., 15.12. 2009, water net, field, Bergsten et al.” (12 exs. NHRS); same data but “S18.75643, E044.71398, 119 m.a.o., 17.12. 2009” (2 exs. NHRS); same data but “S19.03419, E044.77499, 41 m.a.o., 15.12. 2009” (2 exs. NHRS); same data but “S18.75724, E044.71239, 72 m.a.o., 17.12. 2009” (3 exx. NHRS); same data, add: “000000468 NHRS-JLKB” (1 ex. NHRS); same data but “S19.14114, E044.81245, 45 m.a.o., 14.12. 2009/000000466 NHRS-JLKB” (1 ex.NHRS); “Prov. Antsirarana, P.N. Montagne d’Ambre, elev. 960 m 26–29.1. 2001/N -12*30'52” E 49*10'53” leg. Irwin & al, malaise trap” (1 ex. CAS); “Prov. Antsirarana, P.N. Montagne d’Ambre, elev. 1125 m 30.5.-6.6. 2001/N -12°31'13” E 49°10'45” leg. Irwin & Hala, malaise trap” (1 ex. CAS); “Prov. Antsirarana, P.N. Montagne d’Ambre, elev. 1125 m 21-26.4. 2001/N -12°31'13” E 49°10'45” leg. Irwin & Hala, malaise trap” (1 ex. CAS); “Prov. Antsirarana, Sakalava beach, dwarf littoral forest 10 m, 13–20.8. 2001/N -12°15'46” E 49°23'51” Irwin & Hala leg., malaise trap – across sandy trail” (1 ex. CAS); “Andjamangirana (Majunga) 19.10. 2001/stream in dry forest, upstream. Rice field area (road to Tsaratanana) 200 m a.s.l., 30,8 °C, 0.008 mS/cm/Gerecke & Goldschmidt leg.” (1 ex. BMNH, 1 ex. MZH); “Anjozorobe (Antananarivo) Ravoandrina Riv. Ampanakamonty 21.7.2001 / 1280 m asl, 12,8 °C, 0,078 mS/cm/Gerecke & Goldschmidt leg.” (1 ex. BMNH); “SE-Mad. Rés. Nat. Integr. de Andohahela (NW Ft. Dauphin) Parcelle 1 (versante E) – 300 m foresta pluviale 26.5. 1991/Bartolozzi, Taiti, Raharimina leg. / L. alluaudi Rég. det. Rocchi 1991”(2 exs. CSR); “E-Mad. Ampamoho nr Andilamena 1200-1300 m asl, 18-20.1. 1995 Dunay & Janak” (1 ex. MZH); “Fiananrantsoa, Mania River S Ilaka, 900 m 27 rd km NNW Ambositra 23.10. 2001 Schuh leg.” (1 ex. NMW); “Foret de Fito, ex. coll. Dr. Breuning” (1 ex. MRAC); “Prov. de Tamatave, Foret de Perinet 17.7. 1970 Pederzani / L. lateralis Sharp det. Pederzani” (1 ex. CSR; determination uncertain).

Diagnosis

Laccophilus alluaudi forms together with L. comes, L. tigrinus, L. pseustes and L. furthi a distinct group of species characterized by quite similar colour pattern of body and male genitalia exhibiting same ground plan. Penis of L. alluaudi, L. comes and L. furthi is not evenly curved in lateral view as in L. tigrinus, but angled. Longitudinal markings of L. comes reach humeral region while in L. alluaudi and L. furthi corresponding markings fade away before reaching humeral region. Finally, penis (dorsal aspect) in L. alluaudi is strongly twisted, while almost straight in L. furthi. Furthermore L. comes penis is in lateral view medially slightly depressed and immediately after angle towards apex there is a minute, sharp knob which is lacking in L. alluaudi.

Description

(See description of L. comes; only diagnostically important differences noted): Body length 3.4–3.9 mm, width 1.9–2.1 mm. Dorsal, colour pattern (Fig. 392); slightly less pronounced in comparison with L. comes (Fig. 390).

Elytra: Longitudinal markings brownish to ferrugineous; less pronounced in comparison with material of L. comes; especially humeral region with reduced dark markings (Fig. 392).

Ventral aspect: Apical ventrite as in Fig. 35.

Male genitalia: Penis in lateral aspect from base region, almost straight to extreme apex, which is slightly bent upwards; in dorsal aspect, penis bent at right but less so than in L. comes (Figs 233–234).

Female apical ventrite as in Fig. 34.

Distribution

Madagascar (Fig. 531). Also reported from the Comoros (e.g. Guignot 1958b).

Collecting circumstances

Label data indicate that L. alluaudi occurs in both standing and running waters. Collected in a creek with gravel, stones and sand in rainforest along the creek, in small water holes and together with Madaglymbus in water pool on a large rock with wood and leaves.

Laccophilus furthi sp. n.

Figs 37, 235–236, 393, 532

Type locality

Madagascar: Prov. Fianarantsoa, 7 km West of Ranomafana.

Type material

(2 exs.): Holotype: male: “Madagascar: Prov. Fianarantsoa, 7 km W Ranomafana, 1100 m 8-21. October 1988 W.E. Steiner / From stream with mossy rocks and sandy bottom, montane rainforest” (USNM; habitus in Fig. 393). – Paratype: “Madagascar 19–22.1. 2000 Toamasina distr. (Périnet) Analamazaotra S Andasibe 18°56'09"S, 48°24'48"E, O. Hovorka leg., black light” (1 ex. NMPC).

Diagnosis

Laccophilus furthi resembles most of L. alluaudi, L. comes and L. tigrinus. Distinguishable by study of the penis, the shape of which is almost straight and comparatively broad in L. furthi, while sinuate and less evenly broad in the three resembling species mentioned above. Vide diagnosis of L. alluaudi (p. 41).

Description

Body length 3.7–3.8 mm, width 2.0 mm. Dorsal, colour pattern of body as in Fig. 393. Dark, longitudinal lines on elytra gradually fade away towards base of elytra.

Head: Pale ferrugineous. Submat to rather shiny, finely microsculptured. Reticulation double, but size classes of meshes difficult to distinguish. In part meshes mixed and sculpture appears irregular, consisting of variable shaped meshes. Impunctate, except at eyes; with fine, irregular punctures, which extend a short distance towards middle of head-disc.

Pronotum: Pale ferrugineous; no distinct colour pattern. Submat to rather shiny, finely microsculptured. Reticulation double but size classes of meshes difficult to distinguish. In part meshes mixed, and sculpture appears irregular, consisting of variable shaped meshes. Impunctate, except frontally and laterally; with very fine, scattered punctures.

Elytra: Pale ferrugineous, with dark ferrugineous to brownish, longitudinal areas, which anteriorly, gradually fade away in the holotype while quite distinct in paratype (Fig. 393). Rather shiny, although finely and densely microsculptured. Reticulation double, but large meshes almost absent because strongly reduced (only rudiments discernible). Very fine, irregular punctures form a somewhat vague, discal row. Dorsolateral and lateral rows indistinct; indicated by some scattered fine punctures. Pre-apical row consists of fine, slightly impressed punctures provided with fine hairs. In apical quarter of elytra fine punctures mixed and no separate rows discernible.

Ventral aspect: Pale ferrugineous, except metathorax and -coxal plates; blackish to dark ferrugineous. Shiny to rather shiny, microsculpture almost absent. Only very fine rudimentary microsculpture can sporadically be discerned. Abdomen with very fine, curved striae. Impunctate, except apical ventrite; with some fine, scattered punctures; shape of ventrite almost symmetric (Fig. 35). Metacoxal plates with 3-4, very fine, in part reduced, transverse furrows. Lateral impression on metacoxal plate moderate but clearly discernible. Prosternal process rather slender, posteriorly moderately extended, apically pointed.

Legs: Pale ferrugineous. Pro- and mesotarsus slightly enlarged and extended, with suckers.

Male genitalia: Penis exhibits few modifications, being almost straight both in lateral and dorsal aspects (Figs 235–236).

Female: Unknown.

Etymology

The name is a noun in its genitive form based on the name of Dr. David Furth, Washington D.C., USA, who kindly assured the loan of large African Laccophilus collections for this study, deposited in USNM.

Distribution

Madagascar (Fig. 532).

Collecting circumstances

Collecting label informs that L. furthi has been “collected from stream with mossy rocks and sandy bottom in montane rainforest”. The single paratype was collected by black light.

Laccophilus tigrinus Guignot, 1959

Figs 38–39, 237–239, 394, 532

Laccophilus tigrinus Guignot 1959c: 76, 78, 79 (original description, faunistics); Guignot 1961a: 931 (faunistics); Wewalka 1980: 724, 726 (faunistics); Nilsson 2001: 251 (catalogue, faunistics); Nilsson 2015: 218 (catalogue, faunistics).

Type locality

Comoro Islands: Anjouan, Foret de M’Remani.

Type material, studied

(5 exs.). Holotype: male: “Type / F. Guignot det. 1955 Laccophilus tigrinus sp. n. Type, male symbol” (MNHN). – Paratypes: Same data as holotype but labelled “Paratype” (1 ex. MNHN); “Anjouan Fet de M’Remani X-1953 (Millot) / male symbol / Paratype” (1 ex. MNHN; habitus in Fig. 394); “Moheli Foret de Fomboni 600 m 2eme torrent 6.54 (JM) / male symbol / Paratype” (1 ex. MNHN); Same data but provided with label “Coll. Guignot” and a peculiar label with the following text: “R. Mouchamps det. 63 L. mohelicus sp. n. paratype” (1 ex. MNHN).

Additional material studied

(5 exs.): Comoro Islands: “Grande Comore Nioumbadjou 9.8. 1981 R. Joqué”(2 exs. MRAC, 1 ex. MZH); “Moheli Foret de Fomboni 600 m 2eme torrent 6.54 (J.M.)” (2 exs. IRSNB).

Diagnosis

Laccophilus tigrinus resembles most of all species of L. comes, L. alluaudi and L. furthi, but it is often slightly larger and penis in lateral view almost evenly curved and not angled. Additionally, penis of L. tigrinus on one side, provided with a latero-basal expansion which is absent in L. comes, L. alluaudi and L. furthi. Resembles also of L. pseustes but size of body smaller.

Description

(only diagnostically important differences to description L. alluaudi are recognized):

Body length 3.6–4.0 mm, width 2.1–2.3 mm. Dorsal, colour pattern of body distinct (Fig. 394).

Head: Posteriorly head often becomes gradually a little darker.

Pronotum: Frontally in middle with a vague, somewhat darker area.

Elytra: Very fine, sparse punctures form a discal, a dorsolateral and a lateral row of punctures discernible on each elytron. Pre-apical, lateral furrow rather shallow; punctate with fine hairs.

Ventral aspect: Metacoxal plates in part blackish. Apical ventrite as in Fig. 38.

Legs: Pro- and mesotarsus slightly enlarged and extended, provided with distinct suckers.

Male genitalia: Penis, in lateral aspect evenly curved towards apex, in dorsal aspect, slightly sinuate; basally provided with a distinct enlargement (Figs 237–239).

Female: Pro- and mesotarsus slender. Apical ventrite as in Fig. 39.

Distribution

Comoro Islands (Fig. 532).

Collecting circumstances

Unknown.

Laccophilus pseustes Guignot, 1955

Figs 40, 240–241, 395, 532

Laccophilus pseustes Guignot 1955d: 67 (original description, faunistics); Rocchi 1991: 86 (faunistics, list); Nilsson 2001: 249 (catalogue, faunistics); Pederzani and Rocchi 2009: 95 (faunistics, list); Nilsson 2015: 216 (catalogue, faunistics).

Type locality

Madagascar: Isalo sur Pandamus.

Type material, studied

(1 ex.). Holotype: male: “Isalo sur Pandamus, Inst. Sci. Madagascar VIII.48 RP / Type / Guignot det. 1955 Laccophilus pseustes Type, male symbol” (MNHN; habitus in Fig. 395).

Additional material studied

(23 exs.): Madagascar: “Fian: Isalo, source of piscine naturelle, waterhole, P41K: N -22.553: E: 45.368: 859 m 12.5. 2006 leg. Bergsten et al / BMNH(E) <794199> DNA voucher / L. pseustes det. Bergsten” (1 ex. NHRS); same data but “BMNH(E)<745062> DNA voucher ” (1 ex. NHRS); “Fian: Isalo, source of piscine naturelle, small water holes at beginning of stream P41K, 12.5. 2006 N-22°33.206, E45°22.089, 859 m, Bergsten et al.” (5 exs. NHRS); “Fian: Isalo, Canyon de Makis: River: P41E: N: -22.548: E: 45.408: 780 m, 11.5. 2006 leg. Bergsten et al / BMNH(E) <745068> DNA voucher / L. pseustes det. Bergsten” (1 ex. NHRS); “Fian: Isalo, Canyon de Makis, sandy bottom of river, with side pools and hygropetric sections at sides, wood in water, P41E 11.5. 2006 N-22°32.922, E45°24.064, 780 m Bergsten et al.” (6 exs. NHRS, 2 exs. MZH); “Fian, Isalo, Namaza R.: stagnant waterpool P41I: N: -22.539: E: 45.377: 794 m, 12.5. 2006 leg. Bergsten et al / BMNH(E) <745060> DNA voucher / L. pseustes Bergsten det.” (1 ex. NHRS); “Fian, Isalo: P41O: Trib. to Namaza R.: Waterhole, N: -22.543: E: 45.377, 842. 1624 m, 13.5.2006 leg. Isambert et al / DNA voucher BMNH(H) <831017> MSL 402:E2 / L. pseustes det. Bergsten” (1 ex. NHRS); same as but “DNA voucher BMNH(E) <831016> MSL402:E1” (1 ex. NHRS); same as but “DNA voucher BMNH(E) <831019> MSL402:E4” (1 ex. NHRS); same as but “DNA voucher BMNH(E) <831020> MSL402:E5” (1 ex. NHRS); “Fian: Isalo Namaza R. stagnant water pool with lots of woody debris and leaves 12.5. 2006, N-22°32.348, E45°22.626, 794 m, Bergsten et al. (2 exs. NHRS).

Diagnosis

Laccophilus pseustes resembles most L. alluaudi and L. comes and also of some other species in this group but its body is generally clearly larger. Additional diagnostic features are found in the shape of the penis: In lateral aspect, penis long and narrow and extreme apex slightly curved upwards; in dorsal aspect, penis quite broad and somewhat sinuate with narrow slightly curved tip.

Description

Body length 4.1–4.2 mm, width 2.3–2.4 mm. Elytral colour pattern slightly vague (Fig. 395).

Head: Pale ferrugineous. Rather shiny, finely microsculptured. Reticulation indistinctly double. Large meshes only slightly more strongly developed in comparison with small meshes. In part, small meshes reduced and hardly visible. Impunctate, except at eyes; with fine, scattered punctures; closely towards centre of head, there is an additional small group of fine punctures located in a small depression.

Pronotum: Pale ferrugineous. Submat, finely microsculptured. Reticulation double; large meshes contain 3–5 small meshes. Laterally and frontally, with fine, scattered punctures.

Elytra: Pale ferrugineous, with somewhat vague, dark ferrugineous to ferrugineous colour pattern (Fig. 395). Slightly mat, finely microsculptured; reticulation double, but small meshes distinct while large meshes strongly reduced and only in part discernible. Fine, scattered punctures form a somewhat irregular, discal row. Dorsolateral and lateral rows indistinct; simply indicated by few scattered punctures. Postero-laterally, with a pre-apical, pubescent furrow.

Ventral aspect: Pale ferrugineous to ferrugineous, no distinct colour pattern. Submat, very finely and in part indistinctly microsculptured. Abdomen with fine, curved striae. Metacoxal plates with some 8–9 transversely located, shallow furrows. Almost impunctate, except on apical ventrite; with a few scattered punctures. Apical ventrite symmetric and lacks lateral knob (Fig. 40). Prosternal process quite narrow, apex short, only slightly extended, apically pointed.

Legs: Pro- and mesotarsus somewhat enlarged, extended and provided with suckers. Hindlegs quite robust.

Male genitalia: Penis in dorsal aspect comparatively broad with narrow, slightly curved apex; in lateral aspect, penis quite slender and long with tip curved slightly upwards (Figs 240–241).

Female: Pro- and mesotarsus rather slender.

Distribution

Madagascar (Fig. 532).

Collecting circumstances

Mainly unknown. Label data simply indicate that the species has been collected in a river with sandy bottom, with side pools and hygropetric sections at sides, wood in water. Additionally, recorded in stagnant water pool with lots of woody debris and leaves.

Laccophilus seyrigi Guignot, 1937

Figs 41–42, 242–244, 396, 532

Laccophilus seyrigi Guignot 1937: 140 (original description, faunistics); Guignot 1959a: 544, 545, 550 (description, faunistics); Rocchi 1991: 86 (faunistics, list); Nilsson 2001: 250: (catalogue, faunistics); Nilsson 2015: 217: (catalogue, faunistics).

Type locality

Madagascar: Békily.

Type material, studied

(6 exs.). Holotype: male: “Madagascar Békily III 1936 – S / male symbol / Type” (MNHN). – Paratypes: Same data as holotype but “Paratype” (1 ex. MNHN; habitus in Fig. 396); Same data as holotype but “female symbol / Paratype” (3 exs. MNHN, 1 ex. IRSNB).

Diagnosis

Laccophilus seyrigi forms together with L. comes and some other morphologically similar species an own group of species. L. seyrigi is, however, a deviating species in the group, and it is separated from the other species by clearly larger body size, by peculiar elytral colour pattern, by longitudinally extended meshes of microsculpture and by species-characteristic shape of penis; in dorsal aspect being long, slender and straight; in lateral aspect basally, with a distinct enlargement.

Description

Body length 5.0–5.8 mm, width 2.8–2.9 mm. Dorsal, colour pattern of body rather distinct and stable; only minor variation exhibited (Fig. 396).

Head: Pale ferrugineous. At eyes with dense and fine punctures. Additionally with fine punctures in a short transverse impression located close to each eye. Submat, finely microsculptured. Reticulation double. Coarse meshes distinct; fine reticulation reduced, only in part discernible. Fine meshes extensively obliterated.

Pronotum: Pale ferrugineous, frontally in middle with distinct dark ferrugineous area; posteriorly in middle with a vague, bilobed ferrugineous to dark ferrugineous spot. Almost impunctate, except frontally and laterally with fine scattered punctures. Rather shiny, distinctly microsculptured. Reticulation double. Large meshes distinct; especially in middle meshes longitudinally extended. Fine meshes clearly discernible laterally; medially fine reticulation absent or almost totally obliterated.

Elytra: Pale ferrugineous, with distinct, dark ferrugineous markings (Fig. 396). Almost impunctate; discally and laterally with a few, fine punctures. Rather shiny, although distinctly microsculptured. Reticulation double. Coarse meshes distinct, in frontal half meshes longitudinally extended. Fine meshes frontally almost totally obliterated; in posterior half fine meshes clearly discernible. When discernible, coarse meshes contain 3–6 fine meshes.

Ventral aspect: Pale ferrugineous, laterally gradually darker, or with quite distinct, dark, lateral spots; dark ferrugineous to blackish. Abdomen pale ferrugineous. Ventrites latero-posteriorly with darker areas (dark ferrugineous to blackish). Apical ventrite pale except for latero-basally, with dark ferrugineous areas. Apical ventrite not modified (Fig. 41). Almost impunctate. Apical ventrite with fine punctures. Rather shiny, although very finely microsculptured. Microsculpture in part reduced, obliterated. Abdomen with fine striae. Metacoxal plates with 5–6 very fine, shallow furrows, which are almost transversely located. Prosternal process rather slender and comparatively short, apically pointed.

Legs: Pro- and mesotarsus rather slender and extended, with protruding suckers.

Male genitalia: Penis in dorsal aspect long, slender and straight; in lateral aspect basally, with a distinct enlargement (Figs 242–244).

Female: Pro- and mesotarsus slender, somewhat extended. Apical ventrite uniform (Fig. 42).

Distribution

Madagascar (Fig. 532).

Collecting circumstances

Unknown.

Species group 5 (L. isamberti group)

Diagnosis. Medium sized species with body length 3.7–4.0 mm, width 2.2–2.4 mm.

Shape of body oval-oblong, body dorsoventrally flattened (Fig. 397). Dorsal colour pattern distinct and peculiar. Elytra with blackish to dark ferrugineous base from which comparatively broad blackish to dark ferrugineous, longitudinal lines start, leaving pale ferrugineous, somewhat vague lines between them. Apically dark lines become somewhat irregular (Fig. 397). Body microsculpture double, although division in two size-classes difficult. Large meshes obscure, extensively strongly reduced (only fragments of large meshes discernible), while small meshes rather distinct.

Prosternal process quite narrow, apex only moderately extended, apex pointed. Apical ventrite simple, not distinctly modified; no asymmetrical knob located on one side of ventrite (Fig. 44). No stridulatory apparatus discernible on metacoxal plates. Metacoxal process posteriorly expanded (not truncate as in other African Laccophilus) (Fig. 7).

Paramere quite simple but clearly enlarged in apical half (Fig. 246). Penis, lateral aspect, in apical half evenly curved, exhibits no distinct modifications (Fig. 245).

Species composition and distribution. One species recognized in this species group. Only recorded from Madagascar.

Laccophilus isamberti sp. n.

Figs 7, 43–44, 245–246, 397, 533

Type locality

Madagascar: Zahamena N.P., Ambatondrazaka. (N: -17,50800 E: 48,72283).

Type material studied

(23 exs.): Holotype: male: “MAD TOAM: Ambatondrazaka Zahamena: Zahamena N.P. close to Camp site 1 Manambota River, on the Rocks. PB60BI01: N: -17,50800 E: 48,72283: 943 m 28.XII. 2006 Leg. Isambert et. al. / Laccophilus sp.n. lateralis gr. Det. J. Bergsten. 2008” (NHRS, habitus in Fig. 397). – Paratypes: Same data as holotype (4 exs. NHRS, 2 exs. MZH); same data as holotype and “DNA VOUCHER BMNH(E) <834433> MSL:430:G02” (1 ex. BMNH); same data as holotype, but ”<834432> MSL:430:G01” (1 ex. BMNH); “MAD TOAM: Ambatondrazaka Zahamena: Zahamena N.P. Analamaintsoa Forest between Camp Site 1 and Camp Site 2 Manambota River. PB60BI02: N: -17,50750 E: 48,72250: 1071 m 29.XII. 2006 Leg. Isambert et al. / Laccophilus sp.n. lateralis gr. Det. J. Bergsten 2008/DNA VOUCHER BMNH(E) <834434-7> MSL:430:G03-6” (3 exs. BMNH, 1 ex. MZH); same data but not vouchers (1 ex. NHRS, 1 ex. NMW); same data but”P60BI04” (2 exs. NHRS, 1 ex. NMW); “MAD TOAM: Ambatondrazaka Zahamena: Zahamena N.P. Analamaintsoa Forest Manambota Rv, 500 m between Camp 1 & Camp 2 PB60BI11 30.XII. 2006 N: -17,50717 E: 48,72400 leg. Isambert et al. / Laccophilus sp.n. lateralis gr. Det. J. Bergsten, 2008/DNA VOUCHER BMNH(E) <834438> MSL:430:G07” (1 ex. BMNH); same data but not vouchers (2 exs. NHRS); “MAD TOAM: Ambatondrazaka Zahamena: Zahamena N.P. Analamaintsoa Forest 5th stream between Camp 1 and Camp 2 PB60BI16 31.XII. 2006 N: -17,52183 E: 48,72067 1092 m leg. Isambert et al. / Laccophilus sp.n. lateralis gr. Det. J. Bergsten, 2008/DNA VOUCHER BMNH(E) <834439> MSL:430:G08” (1 ex. BMNH).

Diagnosis

A deviating species, which on the basis of external appearance and shape of penis may be closely related to species group 4 (L. alluaudi). Laccophilus isamberti, however, exhibits peculiar modification on metacoxal process, being posteriorly expanded (Fig. 7). This feature is lacking in all other African Laccophilus species and accordingly the location within the genus Laccophilus can also be discussed. Further study is definitely needed to establish the status of the species L. isamberti.

Description

Body length 3.7-4.0 mm, width 2.2-2.4 mm. Dorsal colour pattern of body (Fig. 397); minor variation observed in width of elytral, longitudinal markings.

Head: Posteriorly dark brown; anteriorly head becomes gradually paler. Rather shiny, although finely microsculptured. Reticulation indistinctly double; large meshes generally quite distinct while small meshes in part indistinct. When discernible, large meshes contain 2-4 small meshes. At eyes with fine, irregular punctures. Area with punctures extended from eyes towards middle of head but they don’t meet medially.

Pronotum: Blackish ferrugineous to dark ferrugineous, laterally broadly paler; ferrugineous to pale ferrugineous. Slightly mat, finely microsculptured. Reticulation indistinctly double. Small meshes distinct; large meshes strongly reduced and almost absent, only slightly stronger developed than small meshes. Large meshes, when discernible, contain 2-4 small meshes. Impunctate, except at margins with fine, irregular punctures (mediobasally, punctures also absent).

Elytra: Blackish to dark ferrugineous, with pale ferrugineous, longitudinal markings. Posteriorly markings undulate (Fig. 397). Width of longitudinal markings slightly variable. Slightly mat, finely microsculptured. Reticulation double, but large meshes strongly reduced and almost absent. Fine, scattered punctures form a discal row. Dorsolateral and lateral rows indicated by a few scattered punctures. Lateral, pre-apical furrow shallow, rather finely pubescent.

Ventral aspect: Blackish to dark ferrugineous; without distinct colour pattern. Rather shiny, very finely microsculptured. Reticulation in part absent. Abdomen basally with fine, curved striae. Almost impunctate; ventrites with fine punctures. Apical ventrite as in Fig. 43. Prosternal process quite narrow, apex only moderately extended, apex pointed, Metacoxal plates in anterior half with four outwards curved, distinct furrows; in posterior half furrows strongly reduced. Metacoxal process posteriorly expanded (Fig. 7).

Legs: Pro- and mesotarsus slightly enlarged, with suckers.

Male genitalia: Shape of penis (Figs 245–246) resembles some of the species located in species group 4 and especially the species L. pictipennis, placed in an own species group 6.

Female: Pro- and mesotarsus slender. Apical ventrite as in Fig. 44.

Etymology

The name is a noun in its genitive form based on the name of Dr. Benjamin Isambert, Toulouse, France, who collected the type material during his PhD studies.

Distribution

Madagascar, so far only known from Zahamena National Park (Fig. 533).

Collecting circumstances

This is a lotic species occurring in rivers and streams. The known localities are at an altitude of 1000-1100 m.

Species group 6 (L. pictipennis group)

Diagnosis. Large species with body length 4.4–4.7 mm and width 2.5–2.8 mm.

Shape of body oval-oblong; body dorsoventrally flattened (Fig. 398). Dorsal side with distinct colour pattern; pale ferrugineous to ferrugineous with dark ferrugineous, often somewhat vague patches (Fig. 399). Body microsculpture double; both small and large meshes exhibited. Small meshes extensively reduced and weakly developed; sometimes totally missing.

Prosternal process moderately slender, posteriorly not strongly extended, apex pointed. Apical ventrite with posterior end excavated on both sides and medially ventrite moderately produced backwards; lacks asymmetric knob on one side (Fig. 45). No stridulatory files on metacoxal plates. Metacoxal process not extended posteriorly (Fig. 6).

Paramere simple, elongate, apically not enlarged or modified (Fig. 248). Penis rather slender, clearly curved and apex not distinctly modified (Fig. 247).

Species composition and distribution: One species recognized in this species group. In Africa it occurs in north-eastern part; also recorded from Arabian Peninsula.

Laccophilus pictipennis Sharp, 1882

Figs 45–46, 247–248, 398–399, 534

Laccophilus pictipennis Sharp 1882: 305 (original description, faunistics); v. d. Branden 1885: 23 (catalogue, faunistics); Régimbart 1895: 131, 132 (description, faunistics, discussion); Zimmermann 1920a: 24 (catalogue); Zimmermann 1930: 21, 23 (description, faunistics); Balfour-Browne 1951: 193 (discussion, faunistics); Guignot 1952d: 4, 5 (discussion); Guignot 1959a: 533, 536 description, faunistics); Brancucci 1979: 159 (faunistics, description, discussion); Brancucci 1983b:274, 394, 416 (description, faunistics; lectotype designation); Rocchi 1984: 447 (faunistics.); Nilsson and Persson 1993: 79, 94 (faunistics, discussion); Nilsson 2001: 248 (catalogue, faunistics); Nilsson 2003: 77 (faunistics, list); Angus 2003: 16 (synonymy, discussion); Hajek and Reiter 2014: (faunistics, biology); Nilsson 2015: 168 (catalogue, faunistics).

Laccophilus wehnckei Sharp 1882: 306 (original description, faunistics); v. d. Branden 1885: 24 (catalogue, faunistics); Régimbart 1895: 131 (description, faunistics, discussion); Zimmermann1920a: 28 (catalogue); Guignot 1943: 99 (description, faunistics); Guignot 1946c: 270, 273, 277, 312, 315 (L. wehnchei Sharp, misspelling: description, faunistics, discussion); Guignot 1952d: 4 (discussion: misidentification by Guignot 1946c); Legros 1954: 268 (faunistics); Legros 1958: 211 (faunistics); Guignot 1959a: 536, 567 (listed as synonym of L. pictipennis Sharp, discussion); Nilsson and Persson 1993: 79 (list, synonymy); Nilsson 2001: 249 (catalogue, faunistics, list, synonymy); Nilsson 2015: 168 (catalogue, faunistics, list, synonymy). Confirmed synonym.

Type localities

Laccophilus pictipennis: Saudi Arabia: Hedjaz.

Laccophilus wehnckei: Tanzania: Zanzibar.

Type material studied

(6 exs.): Laccophilus pictipennis: Lectotype (designated by Brancucci (1983b)): Male: “Lectotype / Lectotypus / male-label / 566 / Hedjaz Millingen / Sharp Coll. 1905-313 / Laccophilus pictipennis Shp M.E. Bacchus det. 1977 Syntype / Lectotype Laccophilus pictipennis Sharp des. Brancucci 81” (BMNH; habitus in Fig. 398). – Paralectotypes: same data as lectotype but labeled as “Paralectotype” (1 ex. BMNH); “Paralectotype / Paralectotypus / Abyssinia / Sharp Coll. 1905-313 / Type 566 L. pictipennis / Laccophilus pictipennis Shp M.E. Bacchus det. 1977 Syntype” (1 ex. BMNH); same data but add: “Raffray” (1 ex. BMNH).

Laccophilus wehnckei: Lectotype (by present designation): female: “Type / E. Africa / Sharp Coll. 1905-313 / Type 620 Laccophilus wehnckei sp. n. Zanzibar” (BMNH; habitus in Fig. 399). – Paralectotype: female: principally same data as lectotype but labelled as “cotype” (1 ex. BMNH).

Additional material studied

(15 exs.). Ethiopia: “Saati Levander” (1 ex. MZH); “Abyssinia” (1 ex. ZMHB). – Somalia: “Daragodleh 25.6. 1963 Linnavuori” (6 exs. MZH); “Lasgori / L. pictipennis Sharp det. Brancucci 1982” (5 exx. ZMHB, 1 ex. NHMB). Non-African record: – Yemen: “W. Aden Prot. nr Lahej 9-15.7. 1963 Linnavuori” (1 ex. MZH).

Comments on synonymy

Earlier established synonymy of L. pictipennis and L. wehnckei is confirmed by study of external characters; no diagnostically important differences detected. As no males are available of L. wehnckei we could not in this case undertake comparison of male genitalia. L. discretus Sharp, 1882, described from Saudi Arabia, has earlier been synonymized with L. pictipennis. It has never been recorded from Africa by its own name and accordingly, it is outside the scope of this study.

Diagnosis

Laccophilus pictipennis is characterized by peculiar elytral colour pattern in combination with penis, which is slightly and evenly curved, tapering gradually towards its apex. Note also that male apical ventrite lacks asymmetrically located knob, although excavated on each side of midline and slight medial extension (Fig. 45).

Description

Body length 4.4–4.7 mm, width 2.5–2.8 mm. Dorsal, aspect of body with fairly distinct colour pattern. African specimens seem to have vaguer dorsal colour pattern (Figs 398–399).

Head: Pale ferrugineous. Punctation indistinct, almost absent; close to eyes with two minute deptressions with irregular, fine punctures. Shiny, although irregularly and rather finely reticulated. In part double reticulation weakly discernible (delimitation in two distinct size classes of meshes vague).

Pronotum: Pale ferrugineous. At frontal margin with a vague, ferrugineous, almost bilobed marking. Punctation fine, sparse to rather sparse and irregularly distributed. Punctures frontally densest. Rather shiny although microsculptured. Reticulation indistinctly divided into two kinds; smaller meshes sometimes discernible within large meshes. In part, only large meshes well-developed.

Elytra: Pale ferrugineous, with extensive, distinct, dark ferrugineous to brownish markings (Fig. 398). Sometimes elytral colour pattern rather vague (Fig. 399). Reticulation double; large meshes contain generally 3–6 small meshes. Small meshes fine, sometimes weakly developed and indistinct. Irregular, discal, dorsolateral and lateral rows of punctures are discernible. All rows of punctures rather sparse, not forming straight rows.

Ventral aspect: Pale ferrugineous to ferrugineous. Rather shiny, although extensively, finely microsculptured. Besides microsculpture metacoxal plates with shallow, transverse furrows and abdomen especially basally with distinct striae. Apical ventrite lacks knob (Fig. 45). Apex of prosternal process rather narrow and pointed.

Legs: Pro- and mesotarsus slightly enlarged, with fine suckers.

Male genitalia: Penis in lateral aspect quite broad, from base slightly and evenly curved to apex (Figs 247–248).

Female: Pro- and mesotarsus slender. Apical ventrite (Fig. 46).

Distribution

Somalia, Ethiopia, Tanzania (Zanzibar) (Fig. 534). Material examined also from Arabian Peninsula (Saudi Arabia and Yemen). Only personally examined specimens accepted.

Collecting circumstances

Almost unknown in Africa. Hajek and Reiter (2014) report the species from Oman being mostly associated with running water, especially in relatively permanent side pools of streams and river at lower and middle altitudes.

Species group 7 (L. taeniolatus group)

Diagnosis. Medium to large sized species; body length 3.5–5.3 mm, width 1.9–2.9 mm.

Shape of body oval-oblong, dorsoventrally flattened (Fig. 403). Body dorsally with distinct colour pattern. Elytron provided with dense irrorations, which often cover whole disc. In a few species dark irrorations “hollow”; i.e. single irroration encloses a pale and narrow irroration (Fig. 401). One species has a sub-basal, pale area with reduced and sparse irrorations (Fig. 415). Some species have a moderate, mediobasal area with no or reduced irrorations (Fig. 412). Finally, sometimes, irrorations in part merged into larger, dark areas. Dorsal microsculpture almost simple/indistinctly double, of one kind; large meshes strongly reduced and hardly discernible; sometimes discernible but only slightly more strongly developed than small meshes. In two species, small meshes are reduced or weakly developed, while large meshes discernible.

Prosternal process rather slender, extended, apically pointed. Apical ventrite posteriorly on each side excavated, medially posteriorly extended, but asymmetrical knob always absent (posterior outline of ventrite “undulate” with medial extension) (Fig. 47). No stridulation apparatus on metacoxal plates. Metacoxal process posteriorly truncate, not posteriorly extended (Fig. 6).

Paramere simple, somewhat enlarged but not distinctly modified (Fig. 250). Penis more or less evenly curved, apically often enlarged and provided with minor processes. One species with penis apex lacking modifications (Fig. 256).

Species composition and distribution. Nine species are recognized; two of them occur in Madagascar and seven in mainland Africa, South of Sahara. To observe, that from point 3 in the key below, external characters are variable and male genitalia must be studied.

Key to species (males)

1 Large species, length of body 4.8-5.3 mm; elytra with vague but clear, dark, longitudinal lines of which medial lines enclose an undulate pale marking (Fig. 416) L. rivulosus (p. 85)
Smaller species, length of body 3.5-4.6 mm; elytral colour pattern different 2
2 Dark irrorations sparse at elytra-base forming a sub-basal, transverse pale area (Fig. 415); apical half of penis, strongly curved, simple, exhibits no distinct modifications (Fig. 256) L. irroratus (p. 83)
Dark irrorations dense at base; if sparse no transverse, pale area formed (Figs 403, 412); apex of penis exhibits modifications 3
3 Dark irrorations at least partly “hollow” with pale irroration-area enclosed (Fig. 401) 4
Dark irroration almost completely dark (Fig. 412) 6
4 Penis apex broad; ends abruptly and exhibits no extension (Fig. 251) L. inobservatus (p. 63)
Penis apex less broad; apex externally somewhat extended (Fig. 249) 5
5 Penis apex delicate, less pronounced (Fig. 249) (African mainland) L. continentalis (p. 53)
Penis apex robust, pronounced (Fig. 250) (Madagascar) L. posticus (p. 58)
6 Penis apex enlarged on both sides close to truncate apex (Fig. 252) L. simplicistriatus (p. 66)
Penis apex enlarged on one (marginal) side or not enlarged close to truncate apex (Fig. 253) 7
7 Apical process of penis apex curved upwards (Fig. 255) (Madagascar) L. complicatus (p. 80)
Apical process of penis apex not curved upwards (Fig. 253) (African mainland) 8
8 Penis long with apical process distinct (Fig. 253) L. taeniolatus (p. 72)
Penis shorter with vague apical process (Fig. 254) L. propinquus (p. 79)

Laccophilus continentalis Gschwendtner, 1935

Figs 47–48, 249, 400–401, 535

Laccophilus posticus continentalis Gschwendtner 1935a: 16, 18 (original description, faunistics).

Laccophilus continentalis Gschwendter, Guignot 1946c: 279, 281, 283, 312, 316 (discussion, description, faunistics); Capra 1952: 6: (faunistics); Omer-Cooper 1956: 21 (faunistics, biology); Omer-Cooper 1957: 20, 21 (discussion, description); Omer-Cooper 1958a: 57, 59 (faunistics, biology); Omer-Cooper 1958b: 37, 43, 44, 45 (description, discussion, faunistics, biology); Guignot 1959a: 570, 573, 575 (redescription, faunistics); Omer-Cooper 1965: 77, 81 (description, faunuistics); Omer-Cooper 1970: 288, 289, 290 (discussion, description, faunistics); Rocchi 1975: 48 (faunistics); Bilardo 1976: 190 (faunistics, biology); Bilardo and Rocchi 1987: 104 (faunistics, biology); Rocchi 1990:442 (faunistics); Nilsson 2001: 242 (catalougue, faunistics); Nilsson 2015: 210 (catalougue, faunistics).

Laccophilus perplexus Omer-Cooper 1970: 287, 288, 289, 290 (original description, faunistics, discussion.); Nilsson 2001: 248 (catalogue, faunistics); Nilsson 2015: 215 (catalougue, faunistics). New synonym.

Type localities

Laccophilus continentalis: Botswana: Kalahari, Tsotsorogo Pan.

Laccophilus perplexus: Mozambique: Umbeluzi River near Goba.

Type material, studied

(21 exs.). Laccophilus continentalis: Lectotype (by present designation): male: “V.-L. Kal. Exp. Tsotsorogo Pan 17/6-9/7/30 / Type male (symbol) Gschw. / Laccoph. posticus continentalis det. Gschwendtner” (TMSA; habitus in Fig. 400). – Paralectotypes: Same data as lectotype, but “Type female (symbol)” (1 ex. TMSA); “V.-L. Kal. Exp. N’Kate Makarikari 6-23/8/1930 / Paratypus Laccophilus posticus continentalis ssp. L. Gschwendtner” (10 exs. TMSA, 1 ex. OLML); same as lectotype but labelled as “Paratype Gschw.” (5 exs. OLML, 2 exs. AMGS).

Laccophilus perplexus: Holotype: male: “Type / L. perplexus sp. n. / Mozambique Umbeluzi River near Goba 4.12. 1948 J.O.C.” (AMGS; according to original description holotype preserved in BMNH).

Additional material, studied

(518 exs.). Senegal: “Sumpf von Peykone, Senegal 9. 08 Riggenbach S.V.“ (1 ex. ZMHB). – Gambia-S. Senegal: “Stream of Selety 13°10'N-16°36'W 19.2. 1976 Holmen leg.” (2 exs. ZMUC). – Sudan: “Wad Medani a. Bl. Nil 29.10. 1979 Hieke“ (1 ex. ZMHB); same but “12.10. 1979 lux“ (2 exs. ZMHB); same but “18.10. 1979“(1 ex. ZMHB); same but “8.10. 1979“ (2 exs. ZMHB); same but “9.10. 1979“ (1 ex. ZMHB); same but “30-31.10. 1979“ (1 ex. ZMHB); same but “20.10.1979“ (1 ex. ZMHB); same but “15.10. 1979“ (1 ex. ZMHB); same but “22.10.1979“ (2 exs. ZMHB); same but “leg. Königsmann“ (1 ex. ZMHB); “Senaar a.Bl. Nil, lux 21.10.1979 Hieke“ (5 exs. ZMHB); “Umm Banein, light trap 14.11. 1962 Linnavuori” (1 ex. MZH). – Ghana: “N Region Nyankpala 183 m N9°25’-W1°00’ Dr. S. Endrödy-Younga / shore washing 10.2. 1970“ (1 ex. CGW). – Nigeria: “Samaru 17.5. 1959, Sands / light trap” (1 ex. BMNH); “Nt Kano 1.5. 1928 Lindwer Madsen” (1 ex. ZMUC). – Somalia: “Somali Rep. 1961 Roffey”(3 exs. BMNH, 1 ex. MZH). – Kenya: “Kibwezi Scheffler leg.“ (1 ex. ZMHB); same as but “1906“ (1 ex. ZMHB); “Eastern Mwingi, Nguni env. 28.11. 1999 Snizek” (1 ex. NMW); “SE Kenya ShimbaHills 20 km S Mombasa 5.6. 1985 Lödl” (1 ex. NMW, 1 ex. MZH); Mafisini, pond, Kwale District 19.9. 1976 Holmen leg.” (1 ex. ZMUC); “Maji ya Chumvi River, Kwale District 16.9. 1976 Holmen leg.” (1 ex. ZMUC); “Maji-Chumwi (Wa Nyika) Alluaud 7. 1903” (2 exs. NHMB); “Athi River, Machakos District 14.9. 1976 Holmen det“ (1 ex. ZMUC); “Kombeni River, Mazeras, Kilifi distr 15.9. 1976 Holmen” (2 exs. ZMUC, 2 exs. MZH); “Mariakani dam, Kilifi District 16.9. 1976 Holmen” (4 exs. MZH); “Mandera R. Dana 23.10. 1970 Brown” (1 ex. BMNH); “Wajir 27.10. 1970 Brown” (1 ex. BMNH); “Malindi, alle luci, 15.11.-5.12. 1989 / L. continentalis Gschwendtner det. Rocchi 1990” (3 exs. CSR); “Voi 11. 1997 Snizek M. / L. continentalis Gschwendtner det. Rocchi 1990” (3 exs. CSR); “Voi Mtito Andei, light trap roof Tsavo Inn 24-25.11. 1990 Päts & Viklund” (1 ex. MZH); “Fort Hall / Coll. E. Häuser / L. continentalis Gschwendtner det. Brancucci” (1 ex. ZMHB); “Afr. or Jkurha” (1 ex. ZMHB). – Tanzania: “Daressalaam, Pangani und Hinterland Regner” (1 ex. ZMHB); “Daressalaam leg. Methner” (1 ex. NHMB, 1 ex. ZMHB); “Reg. Morogoro Mikumi 17-20.12. 1993 Bednatik” (1 ex. NMW); “Mts Uluguru, Morogoro Campus Fac. Agric., UV, 600 m 5-6. 1971 / L. continentalis Gschw. det. Bilardo” (1 ex. NHMB); “Ponds S of Korogwe, Korogwe District 24.9. 1976 Holmen” (2 exs. ZMUC, 1 ex. MZH); “Rice field S of Tanga, Tanga District 26.9. 1976 Holmen” (1 ex. ZMUC); “Tanga / Sjöstedt” (1 ex. NHRS); “Tanga Reimer S.” (1 ex. ZMHB); “Tanganyika Pond in stream 103 (?) miles from Dodoma 15.2. 1954 JOC” (1 ex. AMGS); “Tanga Prov. 4-5. 1950 Sweeney / At light” (2 exs. BMNH); “Usa River 3900 feet Dr. J. Szunyoghy / Light trap 15.11.-31.12. 1965” (1 ex. CGW); ”2 mi to L. Manyara, seashore 3150 feet Dr. J. Szunyoghy / singled material 1–26. 1965” (1 ex. CGW); “Usagara” (1 ex. ZMHB); “Umgb Urumba 15.9. D.O.A. leg. Methner / L. continentalis Gschw. det. M. Brancucci” (3 exs. ZMHB); “J. Kurha” (1 ex. ZMHB); “Zanzibar Pemba 23. Sept. 1955 Fowler” (5 exs. AMGS); “Zanzibar 13th Sept. 1955 JOC.” (2 exs. AMGS); “Zanzibar 5. 1954 Brown” (2 exs. BMNH, 1 ex. MZH); “Zanzibar / Reimer S. & Schultz. – Zambia: “S Luangwa NP, Mfuwe Crocodile Farm, 13.06.03S, 31.47.32E, 450 m, lux 23.3. 1993 Uhlig” (45 exs. ZMHB, 2 exs. MZH); same but “21.3. 1993” (4 exs. ZMHB); same but “24.3. 1993” (6 exs. ZMHB); same but “13.06S, 31.47S, 21-24.3. 1993 Göllner” (2 exs. ZMHB); same but “Deckert” (1 ex. ZMHB). – Namibia: “E Caprivi: Katima Mulilo, lux, 17.29S, 24.17E, 3-8.3. 1992 Uhlig” (65 exs. ZMHB, 5 exs. MZH; habitus in Fig. 401); same but “Deckert leg.” (2 exs. ZMHb); same but “Göllner leg.” (2 exs. ZMHB); “E Caprivi: 3 km E Katima Mulilo, 17.29S, 24.18E, Hippo Camp, in Swimming Pool 6.3.1992 Uhlig” (1 ex. ZMHB); “E Caprivi: 30 km SE Katima Mulilo 17.31S, 24.25E, Zambesi – Altwasserarm, lux 6.3. 1992 Uhlig” (7 exs. ZMHB, 1 ex. NMNW); “E Caprivi: Mudumu NP, Nakatwa, 18.10S, 23.26E, 8-13.3. 1992 Uhlig” (3 exs. ZMHB); same but “Buffalo Trails Camp lux, ca. 18.10S, 23.26E, 12.3. 1992” (3 exs. ZMHB); same but “Kwando-Ufer, Phragmites, schlammig” (1 ex. ZMHB); “Kavango Popa Falls 18.07S, 21.35E, lux 26.2.-3.3. 1992 lux Deckert” (1 ex. ZMHB); same but “Uhlig” (1 ex. ZMHB); “Kavango Mahango Game Reserve 18.17S, 21.43E, lux 2.3. 1992 Göllner” (1 ex. ZMHB); Okavango Distr., Mutompo, 60 km S Rundu, 18.18.38,7S, 19.15.29,4E, 1180 m NN, 13.3. 2003, hand light trap, Frisch & Voland” (1 ex. ZMHB). – Botswana: “Serowe, sevage ponds, Farmer’s Brigade 1.6. 1987, SE22 26 BD, Forchhammer leg.” (3 exs. MZH); same but “7.6. 1987” (3 exs. MZH); same data but ”4. 1988 / L. continentalis Gschwendtner det. Rocchi 1993” (2 exs. CSR); “Kasane, Chobe Safari lodge, 17.48.32S, 25.08.39E, 26.11.1993 lux, Uhlig” (5 exs. ZMHB, 3 exs. MZH); “5 km NW San-ta-wani Safari Lodge, 19.27.01S, 23.38.46E, lux 8-9.3.1993 Uhlig” (27 exs. ZMHB); “5 km NW San-ta-wani Safari Lodge, 19.27S, 23.38E, lux 8-9.3. 1993 Göllner” (1 ex. ZMHB); “6 km E Kalkfontein, 22.04S, 20.56E, 7.3. 1993 lux Göllner” (1 ex. ZMHB); “Okavango Delta, Moremi Wildlife Res. Third Bridge Campsite, lux 10.3. 1993, 19.14.22S, 23.21.24E, 10.3. 1993 Uhlig” (3 exs. ZMHB); “Chobe NP Savuti Camp 18.33.55S, 24.03.53E, lux 11.3. 1993 Uhlig” (5 exs. ZMHB); “R. Thamalakane 7 mls NE Maun 20.4. 1972/at light” (1 ex. BMNH). – Zimbabwe: “Victoria Falls Zambezi NP –Camp 11-12.12. 1993, 17.53S, 25.49E, lux, Uhlig” (1 ex. ZMHB); “S. Rhodesia Pool Lundi 22. N. 1948 JOC.” (6 exs. AMGS); “S. Rhodesia Wankie Reserve water holes 3.9. 1948 / L. continentalis Gschw. Det. JOC.” (1 ex. AMGS); “Wankie Game Res. 5 Sept. 1948 JOC. Pools at Robins rest camp / L. continentalis Gschw. Det. JOC.” (1 ex. AMGS); “Wankie Game Res. 4 Sept. 1948 JOC.” (4 exs. AMGS, 1 ex. BMNH); “Wankie Reserve water hole Sept. 1948” (13 exs. AMGS); “Wankie Game Reserve Shapi pond 5.IX. 1948 JOC. / L. continentalis Gschw. Det. JOC.” (9 exs. AMGS); “Wankie Game Reserve Shapi pan 5.IX. 1948” (3 exs. AMGS); “Wankie Reserve Musumu dam 14.IX. 1948 JOC. / L. continentalis Gschw. Det. JOC.” (1 ex. AMGS); “Wankie Game Reserve Sept. waterhole 1948 JOC. / L. continentalis Gschw. Det. JOC.” (7 exs. AMGS);”5 mi SE Wankie 7.4.1968 Spangler” (34 exs. USNM, 6 exs. MZH); “Marandellas 2. N. 1948 JOC. / L. continentalis Gschw. Det. JOC.” (1 ex. AMGS); “Gwai River 3.4. 1968 Spangler” (8 exs. USNM); “Shamgani 60 km SW of Gweru 2.12. 1998 F. Kantner leg.” (1 ex. NMPC);”Tongwe 30 km N Beitbridge 7.12. 1998 Kantner leg.” (1 ex. NMPC). – Mozambique: “Mozambique Beira 7. Sept. 1955 JOC.” (12 exs. AMGS); “Moz., Dambo Pan 30.6. 1960” (2 exs. AMGS); “Port. E Afr. Lorenco Marques 3.12. 1948 JOC.” (2 exs. AMGS); “Lourenco Marques Dec. 1948 JOC.” (1 ex. AMGS); “Umbuluzi R. nr. Goba 4.12. 1948 J.O-C.” (2 exs. AMGS). – Swaziland: “Muddy pond nr Stegi 4.12. 1948 / L. continentalis Gschw. Det. JOC.” (1 ex. AMGS). – South Africa: “Kruger Nat. Pk. Mathlakusapan 22.37S-31.22E / shorewashing 9.2. 1994 Endrödy-Younga leg.” (9 exs. TMSA, 2 exs. MZH); “Kruger Nat. Pk. Skukuza 12 km S, 25.04S-31.37E / UV light, 6.3. 1996 Endrödy-Younga leg.” (10 exs. TMSA, 2 exs. MZH); “Kruger Nat. Pk, Skukuza Res. camp 24.59S-31.36E / UV light & trap 25.2. 1995 Endrödy-Younga” (1 ex. TMSA); “Kruger Nat. Pk, Skukuza Res. camp 24.59S-31.35E / UV light 3.3. 1996 Endrödy-Younga” (1 ex. TMSA); “Kruger Nat. Pk, Skukuza Res. camp 25.00S-31.35E / UV light & trap 19.2. 1995 Endrödy-Younga” (1 ex. TMSA); “Kruger Nat. Pk, Skukuza, 40 km S dam 20.04S-31.36E/Shorewashing 23.2. 1995 Endrödy-Younga” (1 ex. TMSA); “Kruger Nat. Pk, Skukuza Res. camp 24.59S-31.35E / UV light 7.3. 1996 Endrödy-Younga” (1 ex. TMSA); “Kruger Nat. Pk, Pafuri research camp 4 km W, 22.25S-31.09E / 1.2.1994 UV light & trap Endrödy-Younga leg.” (4 exs. TMSA); “Kruger Nat. Pk, Pafuri res. camp, 22.25S-31.12E/30.1. 1994 UV light & trap Endrödy-Younga leg.” (3 exs. TMSA, 1 ex. MZH); “Kruger Nat. Pk, Pafuri research camp, 22.25S-31.10E / 2.2. 1994 UV light & trap Endrödy-Younga leg.” (1 ex. TMSA); “Kruger Nat. Pk Levuvu River 22.27S-31.10E / 12.2. 1994 shorewashing Endrödy-Younga leg.” (4 exs. TMSA); “Kruger Nat. Pk, Malonga springs 22.36S-31.20E / 8.2. 1994 shorewashing Endrödy-Younga leg.” (1 ex. TMSA); “Kruger Nat. Pk Punda Maria Ngots Dam 21.26S-31.14E / 7.2. 1994 shorewashing Endrödy-Younga leg.” (1 ex. TMSA); “Kruger Nat. Pk Letaba Riv. bel. dam 23.46S-31.30E/1.3. 1995 shorewashing Endrödy-Younga leg.” (1 ex. TMSA); “Transvaal Kruger National Park, Leeu Pan NE Skukuza 1.5. 1951 / L. continentalis Gschw. det. J. Omer-Cooper” (1 ex. MZLU); “Trsvl, Waterhole nr Police Picket, KNP. 30.6. 1960” (25 exs. AMGS); “Trsvl, Kumana Pan 24 km S Satara Camp, KNP, N-24.610, E31.800, 18.6. 1960” (1 ex. AMGS); “Gauteng Tswaing 25.24S-28.06E / 16.2. 2003 light trap TMSA staff leg.” (2 exs. TMSA); “Trsvl Mmabolela Estate 22.40S-28.15E / 21.11. 1991 shorew. Limpopo Riv. Klimazewski leg.” (1 ex. TMSA); “Trsvl 5 mi W Warmbad 24-25.2. 1968 Spangler” (1 ex. USNM); “Trsvl, Hartebeespoortdam, N-25.730, E27.820, 30.5. 1971 Reavell” (1 ex. AMGS); “N. Prov. Messina Nat. Res. 22.21S-30.03E / 13.12. 2000 light trap Müller, Burger leg.” (1 ex. TMSA); “Kw. Natal, Sivai Lagoon 10.3. 1981 Reavell” (1 ex. AMGS); “Zululd. Ndumu Banzi fresh. wat. pan 26.53S-32.16E / 16.2. 1989 shorewashing Endrödy & Klimazew” (1 ex. TMSA); “Natal Zululand Mtuba-tuba 23.9. 1947 JOC”. (1 ex. AMGS); “Natal Durban Umgeni Trägårdh” (1 ex. MZLU); “Zululand Hhu-Hluve 18.IX. 1947” (1 ex. AMGS); “Natal, roadside puddles, ca. 2 km S Mbazwana to Hluhluwe nr Sodwana 9.1. 1997 Turner” (9 exs. NHMB); same data but ”5.3. 1997” (6 exs. NHMB); “Natal, Mkuze NP 17.36S, 32.13E, 2-3.2. 1994 lux Uhlig (1 ex. ZMHB);”Natal, Waterton Timber Co. N-28.20.5, E32.14, at light Atkinson” (5 exs. NHMB).

Specimens with uncertain location

“Lowrie 17.5. 1955” (1 ex. AMGS); “Kotsch Lu.(?) Kohu” (1 ex. NMW).

Comments on the synonymy

The lectotype of L. continentalis and the holotype of L. perplexus have been examined and compared. Minor difference is present in appearance of the elytral colour pattern and shape of the penis. Differences observed are, however, superficial and clearly falls within the variation exhibited by one species. L. continentalis being the older name is the valid name of the species.

Diagnosis

Externally L. continentalis resembles much of L. posticus. Useful diagnostic character is the shape of the penis. In L. continentalis apical part of penis is less prominent in comparison with rest of the penis when L. posticus is characterized by distinctly more prominent apex of penis. The shape of penis separates the two species from all other African Laccophilus species.

Description

Body length 3.6–4.1 mm, width 1.9–2.2 mm. Pale ferrugineous, with dark ferrugineous, extensive but variable elytral irrorations (Figs 400–401). Sometimes single irroration is formed only by distinct outlines leaving the middle pale coloured. Other extreme is that single irroration is solid and totally dark.

Head: Frontally pale ferrugineous; posteriorly head becomes gradually slightly darker; at pronotum ferrugineous to pale brown. Head often uniformly pale coloured. Submat, finely microsculptured; reticulation almost uniform. Indistinct fragments of double reticulation discernible (large meshes incomplete). Almost impunctate; at eyes with fine, hardly discernible punctures.

Pronotum: Pale ferrugineous, frontally with a vague dark ferruginous to dark brownish marking. Darker marking on disc sometimes reduced and visible in frontal part of pronotum. Submat, finely microsculptured. Reticulation almost simple; indistinct fragments of coarse reticulation discernible: large meshes incomplete. Almost impunctate; frontally and laterally with scattered, very fine punctures discernible.

Elytra: Pale ferrugineous, with variable, dark ferrugineous irrorations. Irrorations sometimes in part reduced; outlines only distinct (Figs 400–401). Submat, finely microsculpture; reticulation of one kind. Discally with sparse, irregular row of very fine punctures. Laterally with sparsely scattered, very fine punctures. Fine lateral furrow formed of fine punctures located somewhat posterior to middle towards elytral apex.

Ventral aspect: Pale ferrugineous. Abdomen in part ferrugineous to dark ferrugineous, with lateral and apical areas paler. Almost impunctate. Extensively very finely microsculptured. Prosternal process slender; apex extended and pointed. Metacoxal plates in anterior half with some transversely located, shallow furrows. Abdomen with sparse but distinct striae. Apical ventrite (Fig. 47).

Legs: Pro- and mesotarsus rather long and slender. Pro- and mesotarsus provided with distinct suckers.

Male genitalia: Extreme apex of penis forms a short, sharp and small extension (Fig. 249).

Female: Apical ventrite (Fig. 48).

Distribution

Gambia, Senegal, Sudan, Ghana, Nigeria, Somalia, Kenya, Tanzania, Namibia, Botswana, Zimbabwe, Mozambique, South Africa (Fig. 535). Additionally, L. continentalis is recorded from Swaziland (Omer-Cooper 1958b). Record outside Africa but in close neigborhood is Yemen (Socotra) by Hájek and Reiter (2014).

Collecting circumstances

The biology and habitats of L. continentalis are not well documented. Scattered observations can be gathered from literature and from collection data written on the labels. Label data give that the species is capable of flying and attracted by light. L. continentalis has also been sampled from various water-bodies as ponds and streams. Omer-Cooper (1956) reports the species in Mozambique from pools, a ditch and a slow flowing stream with vegetation as Marsilia sp., Lagarosiphon sp., Limnophyton sp., water lilies and duckweed. Also reported from Zimbabwe in water holes, springs and dams used by the game. Additionally, taken from streams e.g. with a pool in the river bed, in ponds with rock and gravel bottom and some mud deposition; blue water lilies and weed growing in the pools. Guignot (1946c) assumed that the species is a character-species of steppes and savannes.

Laccophilus posticus Aubé, 1838

Figs 49–50, 250, 402–405, 536

Laccophilus posticus Aubé 1838: 428 (original description, faunistics); Erichson 1843: 205 (faunistics); Boheman 1848: 245 (faunistics, description); Sharp 1882: 309 (redescription, faunistics); van den Branden 1885: 23 (catalogue, faunistics); Severin 1892: 472 (type deposition); Régimbart 1895: 136, 137, 138, 141 (description, faunistics, discussion); Alluaud 1897: 212 (faunistics); Peschet 1917: 23, 24, 26, 55 (description, faunistics, discussion); Zimmermann 1920a: 24 (catalogue); Gschwendtner 1935a: 17, 18 (faunistics, description); Guignot 1946c: 279, 281, 283 (discussion, description, faunistics); 1952e: 170 (discussion); Guignot 1955d: 67 (discussion); Vinson 1956: 28 (faunistics, list, biology); Omer-Cooper 1958b: 43, 45 (discussion, description); Guignot 1959a: 570, 572, 573, 575 (description, discussion, faunistics); Guignot 1961a: 931 (faunistics); Vinson 1967: 314 (faunistics, list); Omer-Cooper 1970: 290 (description); Bertrand and Legros 1971: 245 (faunistics, biology); Wewalka 1980: 730 (faunistics list); Bameul 1984: 94 (faunistics,); Rocchi 1991: 80, 86 (faunistics); Nilsson 2001: 249 (catalogue, faunistics); Pederzani and Rocchi 2009: 95 (faunistics, list); Nilsson 2015: 216 (catalogue, faunistics).

Type locality

Mauritius (Ile de France).

Type material studied

(1 ex.). Holotype: female: “Data in NHRS JLKB 000030279/Ex – Museo Dejean / D. Sharp Monogr. / irroratus après Dej. / Laccophilus posticus var. / Dr. Régimbart vidit 1893 / Coll. Oberthur” (MNHN). [Comment: in the original description Aubé (1838) mentions existence of an additional specimen from Philippines. We have not located it and accordingly the exact taxonomic status of this specimen remains unknown.]

Additional material studied

(486 exs.). Mauritius: “Ile de Maurice Avril 1908 d’Emmerez / Museum Paris 1945 Coll. R. Peschet / Laccophilus posticus Aubé” (4 exs. MNHN); “Ile de Maurice Bambous Carié Déc. 1912 / Museum Paris 1945 Coll. R. Peschet / Laccophilus posticus Aubé R. Peschet det. 1917” (1 ex. MNHN); “Balaclava 4.5. 2007 Madl” (2 exs. NMW, 1 ex. MZH); “Nr. Triolet, nr. Fond du Sac, temp. new pool at roadside, terr. plants flooded by water, ephemeral aquatic habitat” (1 ex. CCT); “Ins. Mauritius Westw.” (1 ex. ZMHB); “Mauritius” (2 exs. ZMHB). – Madagascar: “Mahajanga: Boeny: Ankarafantsika NP, S16.30341, E046.81073, 74 m.a.o. 29.11. 2009, 22W black light, field Bergsten et al. leg” (13 exs. NHRS); same data, add “NHRS-JLKB 000000511” (1 ex. NHRS); same data, add “NHRS-JLKB 000000514” (1 ex. NHRS); same data but “S16.30270, E046.80996, 75 m.a.o. 30.11. 2009” (12 exs. NHRS); same data, add “NHRS-JLKB 000000515” (1 ex. NHRS); same data but “S16.30653, E046.81227, 108 m.a.o. 28.11. 2009” (4 exs. NHRS); same data, add “NHRS-JLKB 000000494” (1 ex. NHRS); same data but “S16.31215, E046.81523, 76 m.a.o. 29.11. 2009” (17 exs. NHRS); same data, add “NHRS-JLKB 000000513” (1 ex. NHRS); “Mahajanga: Boeny: Mahavavy Kinkony RS. S16.14653, E045.94926, 9 m.a.o. 4.12. 2009 water net, field leg. Bergsten et al” (15 exs. NHRS); Same data; add “NHRS-JLKB 000000501” (1 ex. NHRS); same data but “S16.06651, E045.77627, 24 m.a.o. 5.12. 2009” (10 exs. NHRS); same data, Andasibe Adasibe add “NHRS-JLKB 000000510” (1 ex. NHRS); same data but “S16.15502, E045.91878, 10 m.a.o. 3.12. 2009” (10 exs. NHRS); same data but “S16.13337, E045.95778, 19 m.a.o., 4.12. 2009 / NHRS-JLBK 000000509” (1 ex. NHRS); same data but “NHRS-JLKB 000000496” (1 ex. NHRS); same data but “S16.15890, E045.93967, 3.12. 2009 / NHRS-JLBK 000000508” (1 ex. NHRS); same data but “S16.14147, E045.93661, 12 m.a.o., 3.12. 2009 / NHRS-JLKB 000000502” (1 ex. NHRS); “Toliara: Menabe: Menabe RS, S19.92773, E045.52253, 102 m.a.o. 10.12. 2009 water net, field” (13 exs. BMNH); same data, add “NHRS-JLKB 000000516” (1 ex. NHRS); same data but “S20.09034, E044.56400, 45 m.a.o. 11.12. 2009 / NHRS-JLKB (1 ex. NHRS); “Toliara: Menabe: Kirindy RS. S20.07641, E044.67478, 65 m.a.o., 11.12. 2009 water net, field” (1 ex. NHRS); same data, add “NHRS-JLKB 000000497” (1 ex. NHRS); same data but “000000504” (1 ex. NHRS); same data but “S20.07655, E044.67532, 57 m.a.o., 12.12. 2009/NHRS-JLKB 000000505” (1 ex. NHRS); “Mahajanga: Melaky btw. Morafenobe-Ambohijanahary, S18.19091, E45.19986, 290 m.a.o. 19.12. 2009 water net, field Bergsten et al.” (9 exs. NHRS); same data, add “NHRS-JLKB 000000512” (1 ex. NHRS); “Mahajanga: Melaky: Tsingy de Bemaraha NP. S18.75724, E044.71239, 72 m.a.o., 17.12. 2009 water net, field Bergsten et al.” (1 ex. NHRS); same data, add “NHRS-JLKB 000000498” (1 ex. NHRS); same data but “S18.775797, E044.71289, 81 m.a.o. 17.12. 2009, 22W black light, Field” (2 exs. NHRS); same data, add: “NHRS-JLKB 000000493” (1 ex. NHRS); same as, except “S19.14210, E044.81309, 59 m.a.o., 14.12. 2009, 22 w black light, field / NHRS-JLBK 000000499” (1 ex. NHRS); “Mahajanga: Melaky: betw. Bekopaka-Antsalova, 18.91556, E044.55546, 47 m.a.o., 16.12. 2009 water net, field Bergsten et al.” (66 exs. NHRS); Same data, add “NHRS-JLKB 000000506” (1 ex. NHRS); “Mahajanga: Melaky: btw. Antsalova-Maintirano S18.30233, E044.18071, 37 m.a.o., 18.12. 2009 Bergsten et al. / NHRS-JLKB 000000503” (1 ex. NHRS); “Ankarana Lat -12.947 Lon 49.0119 27.11. 2004 / DNA voucher BMNH 675044, MSL045:E07 / L. posticus Aubé det. Bergsten” (1 ex, NHRS); “Antsabe, Lat -13.648, Lon 48.721, 21.11. 2004, Balke, Lees & Monaghan / DNA voucher BMNH 672769, MSLO27:D 11 / L. posticus det. Bergsten” (1 ex. NHRS); same but “DNA voucher BMNH 672774, MSL027: E04” (1 ex. NHRS); “Anjiabe Ambony, Ambilobe, Antsabe near camp, ¾ moon, dry, very many water beetles: P25MD12: 21.11. 2004, N: -13.6518, E: 48.7267, 49 m Balke et al” (1 ex. NHRS); “Ambilobe 4. 1951 R.P. / Paratype” (ab. pseudotaenilatus Guignot – not available name) (1 ex. MNHN); “Androka 5. 51 / Paratype” (ab. pseudotaenilatus Guignot – not available name) (1 ex. MNHN); “Isaky Ivondro Ampasy, rice paddies P66, 9.4. 2007 N-24,93056, E46,86317,64 m Ranarilalatiana et al” (41 exs. NHRS); “Isaky Ivondro, Foret Manangotry (rte towards Ranomafana) running water P67C, 9.4. 2007, N-24.7994, E46,86244, 406 m Ranarilalatiana et al.” (4 exs. NHRS); “TOLI, Taolanaro: Isaky Ivondro, Foret Managotry, running water P67C: N -24.799 E 46.862, 406 m 9.4. 2007, leg. Ranarilalalatiana et al./L. posticus Aubé det. Bergsten” (1 ex. NHRS); “TOLI, muddy waterhole, N -23.242, E 44.229, 415 m, 17.5. 2006 Bergsten et al. / BMNH(E) 794210 DNA voucher / L.posticus Aubé Bergsten det. (1 ex. MZH); same as but “BMNH(E) 794228 DNA voucher” (1 ex. NHRS); same but “BMNH(E) 794252 DNA voucher” (1 ex. NHRS); “TOLI NW Ft. Dauphin, rice paddies, P54F, N -24.824, E 46.866, 34.44 m, 19.5. 2006, leg. Bergsten et al / L. posticus det. Bergsten/BMNH(E): <74511> DNA voucher” (1 ex. NHRS); Same but “<794236> DNA voucher” (1 ex. NHRS); “Toli NW Ft Dauphin, rice paddies with water somewhat running under road, 19.5. 2006 N-24°49.472, E46°51.974 34 m Bergsten et al.” (12 exs. NHRS); “TOLI Zombitse Ankilemiletsy, muddy zebu waterhole, some emergent vegetation P42B, 14.5. 2006, N-22°52.112, E44°34.616, 545 m Bergsten et al.” (1 ex. NHRS); “Toli Zombitse Ambiamena, edge PN Zombitse, stagnant zebu-visited marshland, muddy and lots of vegetation, 14.5. 2006 N-22°51.605, E44°37.035, 533 m Bergsten et al.” (4 exs. NHRS); “Toli Zombitse Andranomena R. (Anomena R. ?) near Ranomena, PN Zombitse section Isoky. Pools of muddy & vegetation, stagnant waters in the river basin, among ricefields and Phragmites ? 15.5. 2006 -22°38.407, E44°51.866, 578 m Bergsten et al.” (16 exs. NHRS); “Toli MK Manakaralahy, Manakaralahy R. Dried out river with waterhole on sandy bottom with algal mats, 18.5. 2006, N-24°28.162, E44°35.683, 210 m Bergsten et al.” (3 exs. NHRS); “Toli Sakondry Sakondry R, near RN 10 bridge at Satria river with sandy bottom, wide (50m +) and shallow, algal mats along the edges 17.5. 2006, N-23°20.807, E44°20.353, 214 m Bergsten et al.” (5 exs. NHRS); “Toli Menarandra Menarandra R, 49 km from Ampanihy pools beside a river close to village, algae in pools and sandy bottom with some wood 18.5. 2006, N-24°43.104, E45°2.859, 227 m Bergsten et al.” (9 exs. NHRS); “Fian Isalo, Menamaty R.: degraded river, P41AMO1 N: -22.55, E: 45.401, 757 m, 11.5. 2006 leg. Bergsten et al. / L. posticus det. Bergsten/BMNH(E) <745103> DNA voucher” (1 ex. NHRS); “Fian Isalo, Menamaty R., sandy/stony bottom with some vegetation at edges, zebu crossing, degraded P41C, 11.5. 2006 N-22°29.359, E45°23.505, 715 m Bergsten et al.” (3 exs. NHRS); “Fian Isalo Menamaty Riv. degraded with lots of vegetation, used by women to wash clothes in P41AM01, 11.5. 2006 N-22°33.001, E45°24.074, 757 m Bergsten et al.” (40 exs. NHRS); “Tanandava, lum. Schmitz” (8 exs. MRAC, 1 ex. MZH; habitus as in Figs 404–405); “Marovoay, lampe UV 8. 1962 Dubois” (1 ex. MRAC, 1 ex. MZH); “TAM Morarano-Chrome foret 25 km W. bac j. 4. 1992 Pauly” (1 ex. MRAC); “Diego Suarez Alluaud 10. 1893 / Museum Paris coll. Maurice Régimbart 1908 / posticus Aubé” (1 ex. MNHN); “Maromandia (Antalaha, Antsiranana) 30.10. 2001 / R. Ankavia nr village, 40 m asl / Gerecke & Goldsmith leg.” (1 ex. BMNH, 1 ex. MZH); “Maroambihy (Sambava, Antsiranana) left affl. R. Lokoho upstr. from village 12.11.2001/90 m a.s.l. / Gerecke & Goldsmith leg.” (1 ex. BMNH; habitus in Fig. 402); “E Mad. Fenerive, foret Tampolo 28.12. 1998 Moravec” (1 ex. NMW); “Mad. east Tampolo 17.17S – 49.25E / 12.11. 1998, E-Y: 3372 light trap, Müller leg.” (1 ex. TMSA); same but “E-Y: 3364, 10.11. 1998” (1 ex. TMSA); “S Mad. Umg. Beloha Franz 1969 / L. posticus Aubé det. Wewalka 1969” (1 ex. NMW); “S Mad. Ambilalilalika, Rd Betioky-Beneloka 50 m asl, 27.1. 1995 Dunay & Janak” (1 ex. NMW); “Betsiboka Bas. 53 km Maevatanana, 47°04'33"E, 16°42'13"S, Alt. 49 m, 2.4. 1993 leg. ORSTOM” (4 exs. NMW); “Betsiboka Bas. Manjakavaradrano, Mamokomita Riv., 46°54'20"E, 17°38‘00"S, Alt. 625 m, 16.4. 1991 leg. ORSTOM” (1 ex. NMW); “Betsiboka Bas., Ambalanbongo, Affl. de Betsiboka Riv., 47°00'30"E, 16°48'00"S, 30.3. 1993 leg. ORSTOM” (2 exs. NMW); “Betsiboka bas., Ambohimanalrika, Kamoro Riv., 47°10'06"E, 16°28'55"S, alt. 40 m, 1.4. 1993 leg. ORSTOM” (10 exs. NMW); “Betsiboka Bas., Ambohimanatrika, Kamoro Riv., 47°10'06"E, 16°28'55"S, 4.11. 1995 Elouard & Oliarinony leg.” (1 ex. NMW; habitus in Fig. 403); “Betsiboka Bas., Andriantoany Riv., 46°56'23"E, 17°19'40"S, 5.11.1995 Elouard & Oliarinony leg.” (1 ex. NMW); “Mandrare bas., Betanimena, Manananara Riv., 46°39'20"E, 24 48 17"S, alt. 118 m, 23.5. 1994 leg. ORSTOM” (3 exs. NMW); “Sahankazo Bas., 5 km au Nord de Antsandrangotika Riv. 49°23'46"E – 12°28'40"S, alt. 50 m, 4.4. 1994, leg. Edouard et Sartori” (1 ex. NMW); “Anove à Ivondro Bas., Tampolo, Affl. non nommé Riv. 49°25'40"E, 17°17'07"S, Alt. 8 m, 12.4. 1997, Gibon F. -M, Randriamasimanana D.” (3 exs. NMW); “Onilahy Bas., Ambatofotsy (Horombe), Aff. de Ihazofotsy Riv., 45°40'43"E, 22°30'49"S, 1.6. 1995, Elouard J-M.” (1 ex. NMW); “Onilahy Bas., Onilahy Riv., 44°33'53"E, 23°32'18"S, leg. ORSTOM” (1 ex. NMW); “Tsiribihina Bas., Antazoa, Manampanda Riv., 45°35'04"E, 20°21'40"S, Alt. 145 m, 29.5. 1996 Elouard & Sambatra leg.” (1 ex. NMW); “Antseranana distr., Sambirana Riv., Marovata vill. 5-12.12. 2001 Horak leg.” (2 exs. NMW); “Fianarantsoa Pr. Foret d’Analalava 29.6 km 280˚ W Ranohira, elev. 700 m, 1-5.2. 2003/22°35'30"S, 045°07'42"E, at light in tropical dry forest, Fischer, Griswold et al leg.” (1 ex. CAS); “Tollara Prov., Rés. privé Berenty, Foret de Bealoka, Mandraré Riv. 14.6 km 329˚ NNW Amboasary, elev. 35 m3-8.2. 2002/24°57'25"S, 46°16'17"E, at light in gallery forest, Fischer, Griswold et al leg.” (2 exs. CAS); “Maroansetra, Restaurant La Baquette d’or, because of light trap 20.12. 2006, N-15,42467, E49,73800, 12 m Isambert et al” (3 exs. NHRS); “Madagask., Kaudern / 19.1. / L. posticus Aubé det. Zimmermann” (1 ex. NHRS); same but “febr.” (1 ex. NHRS); same but “Tamatave / febr.” (1 ex. NHRS); “Tamatava distr., Andasibe J. Rolschik leg. 17-30.12. 2001” (1 ex. NMPC); “Katsepy (Majunga) 24-31.12. 1997 Pacholátko” (35 exs. NHMB, 8 exs. MZH); “Majunga, Cirque Rouge 22-23.12. 1997 Pacholátko” (1 ex. NHMB); “Moramanga env. 10-18.12. 1997 Pacholátko” (1 ex. NHMB); “Mangily N of Tulear 12.1. 2005 Bergsten” (5 exs. NHRS); “Kap Diego 1916 Friederichs S.G.” (1 ex. ZMHB); “Andranohinaly Voeltzkow S. / L. posticus Aubé det. Brancucci” (1 ex. ZMHB); “Res. Spec. Beza Mahafaly 44°32'E, 23°40'S, ca. 150 m, 18.5. 199/leg. Bartolozzi et al / L. posticus Aubé det. Rocchi 1991” (2 exs. CSR); “Madagascar, Fairmaire / L. posticus Aubé” (1 ex. NMW); “Madagascar, Fruhstorfer” (1 ex. NMW). – Aldabra: “Aldabra-Ins. N v. Madagascar 4.5. 1895 A. Voeltskow S.” (9 exs. ZMHB); “Ald. Atoll, 9°24'S, 46°20'E, Takamaka Camp 14.2. 1968 Shaffer J.C. / Black light / L. posticus Aubé det. Bameul 1986” (1 ex. USNM).

Diagnosis

Laccophilus posticus and L. continentalis form a pair of species with considerable resemblance. The two species are separated by differences in shape of the penis, being distinctly more prominent in L. posticus than in L. continentalis; especially, anterior part is more extended in L. posticus. The penis of L. posticus and L. continentalis resembles also the penis of L. rivulosus but this species exhibits clear differences in the elytral colour pattern (irrorations are formed as longitudunal lines).

Description

Body length 3.5–4.4 mm, width 2.0–2.5 mm. Dorsal, colouration of body as in Figs 402–405. Quite stable, although colour pattern exhibits some variation.

Head: Pale ferrugineous, posteriorly sometimes slightly darker. Submat, finely, evenly and distinctly microsculptured; reticulation indistinctly double. Large meshes rudimentary, weakly indicated and only in part discernible. Impunctate, except at eyes; with fine, scattered punctures.

Pronotum: Pale ferrrugineous. Frontally and basally in middle with rather vague dark ferrugineous to ferrugineous areas. Submat, finely, evenly and distinctly microsculptured; reticulation indistinctly double. Large meshes rudimentary, weakly indicated and only in part discernible. Impunctate, except anteriorly and laterally; here with fine, scattered punctures.

Elytra: Pale ferrugineous, with dark ferrugineous irrorations. Elytral colour pattern quite stable, exhibits some variation. Single dark marking sometimes only defined by its dark outlines while centre is of pale colour. Rarely the dark, longitudinal markings are in part mixed with each other forming a larger dark area (Figs 402–405). Submat, finely, evenly and distinctly microsculptured; reticulation indistinctly double. Large meshes rudimentary, weakly indicated and only in part discernible. Rows of punctures indistinct, indicated by fine, irregular punctures. Elytron with pre-apical lateral furrow, which is rather discrete and moderately pubescent.

Ventral aspect: Blackish ferrugineous to ferrugineous, prothorax pale ferrugineous to ferrugineous; no distinct colour pattern. Rather shiny, although very finely microsculptured. Abdomen with fine, curved striae. Almost impunctate. Metathorax with about 10 very fine, shallow and in part transversely located furrows. Prothorax moderatly broad, apex distinctly extended, apically pointed. Metacoxal process not distinctly modified. Apical ventrite (Fig. 49); symmetric and lacks lateral knob.

Legs: Pro- and mesotarsus slightly enlarged, provided with distinct suckers.

Male genitalia: Penis in lateral aspect prominent; apex extended to a sharp process (Fig. 250).

Female: Apical ventrite as in Fig. 50. Pro- and mesotarsus slender.

Distribution

Mauritius, Madagascar, Aldabra (Fig. 536).

Collecting circumstances

Insufficiently documented. Vinson (1956) reports the species to occur in stagnant water. On the other hand label data indicates that the species also occur in running water. Sometimes the species has been collected at light in forests (gallery forest and tropical dry forest). Obviously a lowland species the highest elevation for collection being 700 m a.s.l. Laccophilus posticus has also been recorded from rice paddies.

Laccophilus inobservatus sp. n.

Figs 51–52, 251, 406–408, 537

Type locality

Chad: Near Bongor.

Type material

(234 exs.). Holotype: male: “Chad nr Bongor 27.5. 1973 R. Linnavuori” (MZH; habitus in Fig. 406). – Paratypes: Gambia: “Abuko Nature Reserve, at light at the bamboo pool 18.30-20.30, 18.11. 1977 UTM 28 PCK2181 Loc. 24 / Cederholm-Danielsson-Hammarstedt-Hedquist-Samuelsson” (1 ex. MZLU, 1 ex. NHMB); “Outside Abuko Nature Reserve at waterworks. At light 19.00-22.00, 26.2. 1977 Loc. No. 6 UTM 28 PCK 214812 / Cederholm-Danielsson-Hammarstedt-Hedquist-Samuelsson” (1 ex. MZLU); “Tendeba Camp at light in semiarid veg. near River Gambia 18.30-20.30, 14.11. 1977, UTM 28 POK1285 Loc. 12a / Cederholm-Danielsson-Hammarstedt-Hedquist-Samuelsson” (1 ex. MZLU); “3.5 km S Georgetown, Hilltop at Sankuli Kunda, alt. about 30 m, at light 18.30-20.15, 15.11. 1977 UTM 28PEK2593 Loc. 37 /-Danielsson-Hammarstedt-Hedquist-Samuelsson” (1 ex. MZLU); “2 km S Kitty, 7 km SSW Brikama Road junction. In and at Fresh Water Stream 13.11. 1977 UTM 28PCK 1761 Loc 7/-Danielsson-Hammarstedt-Hedquist-Samuelsson” (1 ex. MZLU, 1 ex. MZH, 1 ex. NHMB); “Gam Bathurst Jan. 68 Palm / L. taeniolatus Reg. det. Sven Persson” (4 exs. MZLU); “Bathurst Jan. 1968 T-E Leiler” (5 exs. NHRS, 1 ex. MZH); “Kuntaur NW Georgetown 21.11. 2003 B. Vondel / L. taeniolatus Régb. det. Rocchi 2004” (1 ex. CSR); Gambia: Bakau 6-26.11. 1984 leg. Palm / L. taeniolatus Rég. det. Sven Persson” (1 ex. MZLU); “Gambia Oil Palm and mangrove veg. close to the beach, about 5 km SSW Gunjur, at light 18.45-20.30 13.11. 1977 UTM 28PCK0554 Loc. 8 / Cederholm & al.” (1 ex. NHMB); “Gambia-southern Senegal 13°10'N 16°36'W stream N of Selety 19,2, 1976 M. Holmen” (1 ex. ZMUC). – Senegal: “70 km W Tambacounda 13°57.4'N, 14°15.9'E, 29.6. 2004 Halada leg.” (2 exs. NMPC); “Senegal, Parc National de Niokolo Koba 16.2. 1989” (1 ex. NHMB); “Senegal Cayare II. 46 A. Villiers” (1 ex. NHMB). – Mali: “NW Afr., K. Macina 10.11. 1973 D.R. Reynolds (C. O. R.R.) BM 1974-222” (1 ex. BMNH, 1 ex. MZH); “Kogoni X. 1966 G. Schmitz” (8 exs. MRAC, 2 exs. MZH); “Korioume, Niger Riv. 18.2. 2000 18°40'N, 3°00'W, leg. Komarek & Mayer / L. taeniolatus Régb. det. Wewalka 2001” (1 ex. NMW); “Markala Niger River, 13°40'N, 6°05'W, leg. Komarek & Mayer 9-4 / L. taeniolatus Régb. det. Wewalka 2001” (1 ex. NMW); “S Tombouctou 16°40'N, 3°00'W, 18.2. 2000 leg. Komarek & Mayer 18-1 / L. taeniolatus Régb. det. Wewalka 2001” (1 ex. NMW, 1 ex. MZH); “Mopti Niger Riv. 14°30'N, 4°12'W, 21.2. 2000 leg. Komarek & Mayer 21-2 / L. taeniolatus Régb. det. Wewalka 2001” (1 ex. NMW, 1 ex. MZH); “Goundaka, Bandiagara Riv. 14°29'N, 3°56'W, 12.2. 2000 leg. Komarek & Mayer 12-1 / L. taeniolatus Régb. det. Wewalka 2001” (2 exs. NMW) “SE Douna, Bani Riv. 13°13'N, 5°54'W, 10.2. 2000 leg. Komarek & Mayer 10-1 / L. taeniolatus Régb. det. Wewalka 2001” (1 ex. NMW). – Niger: “Nr. Boureimi 9.11. 1973 R. Linnavuori” (1 ex. MZH); “Rég. de Zinder, Sultanat de Damagherim Dungass, Mission Tilho Dr. R. Gaillard 1910” (6 exs. NHMB). – Sudan: “Blue Nile Ingessana Mts. 17-22.11. 1962 Linnavuori” (6 exs. MZH); “Nile Blue Nile Singa-Roseiras 15-17.11. 1962 Linnavuori” (1 ex. MZH); “Upper Nile Malakal 5-20.1. 1963 Linnavuori” (6 exs. MZH); “Kordofan Lake Keilak 8-11.2.1963 Linnavuori” (4 exs. MZH); “Bahr el Gazal, Wau 19.2. 1963 Linnavuori” (1 ex. MZH); “Bahr el Abiad Trägårdh / L. taeniolatus Régt var.” (1 ex. NHRS); “Torit 2.7. 1980 Armstrong” (1 ex. USNM); “Nairege River 27.2. 1980 Armstrong” (1 ex. USNM); “Gilo water tank (pumped up from stream) 20.3. 1980 Armstrong (8 exs. USNM, 3 exs. MZH); “Kinyetti River at Imeila 19.3. 1980 Armstrong” (5 exs. USNM, 1 ex. MZH); “Sudan Wad Medani am Bl. Nil, 18.10, 1979 lux, leg. Hieke” (1 ex. NHMB); same but “21.10. 1979” (1 ex. NHMB); same but “29.10. 1979” (1 ex. MZH); same but “12.10. 1979” (1 ex. NHMB); same but “30+31.10. 1979” (1 ex. NHMB); “Tombe 17.1. 1954 Omer-Cooper” (1 ex. AMGS). – Chad: Same data as holotype (5 exs. MZH); “Distr. Kanem N’Gouri X-XI. 1958 P. Renaud ex. coll. Breuning” (21 exs. MRAC, 2 exs. MZH). – Ethiopia: “Shoa, Awash NP, Filwoha Hot Springs 25.12. 1988, 1500 m leg. S. Persson / L. taeniolatus Régb. det. A. Nilsson” (2 exs. MZLU); “Shoa, Metahara 2.10. 1988 950 m, water hole in lava field, leg. S. Persson / L. taeniolatus Régb. det. A. Nilsson” (6 exs. MZLU, 2 exs. MZH; habitus in Fig. 407); “Hora Harsadi, Addas 7000 ft 2.12. 1926 J. Omer-Cooper” (1 ex. AMGS). – Burkina Faso:”Pundu Mte Volta1927-1928 Dez.-Juni Olsufiew” (2 exs. NHRS); “Pundu Olsufiew” (38 exs. NHRS, 3 exs. MZH); “Yatenga Gourcy Barrage 300 m 14.1. 1995 Trockengefallen lehmiges Ufer leg. B. Maier / L. taeniolatus Rég. det. Rocchi 2002” (1 ex. CSR). – Ivory Coast: “Comoé NP, N8,5°, W3,5° leg. N. Reintjes, det. F. Pederzani / 1.3. 1999 rock pool in Comoé river bed” (1 ex. NMW). “Comoé NP, N8,5°, W3,5° leg. N. Reintjes, det. F. Pederzani / 3.4. 1999 temporary Pond” (1 ex. NMW). – Ghana: “Upper E Prov. Navrongo env. 11-13.6. 2006 Pokorny leg.” (5 exs. NMPC, 1 ex. MZH). – Nigeria: “Kano St. Wudil-Kari 17.5. 1973 R. Linnavuori” (1 ex. MZH); “NW St. Gummi-Anka 24.7. 1973 R. Linnavuori” (1 ex. MZH); “NE St. Gombe-Bauchi 27.8. 1973 R. Linnavuori” (4 exs. MZH; habitus in Fig. 408); “Detritus pond, Jos-Bauchi rd 9.4. 1963 JOC”(4 exs. AMGS); “Samaru 17.5. 1959 W. Sands B.M. 1961-525 / Light trap” (1 ex. BMNH); “Marsk, road Katsina-Daura 6.4. 1963 JOC” (14 exs. AMGS). – Cameroon: “Maroua (lumière) X-XI. 1965 G. Schmitz” (1 ex. MRAC); “Maroua, Miss. Cath. 26.8. 73” (1 ex. NHMB); “Maroua 5 Aout 71” (1 ex. NHMB); “Tokombere, dint. Maroua, 12.7. 1979 Onore / L. congener O-C. det. Rocchi 1980” (1 ex. CSR). – Zaire: “P.N.G. Ndelele. K.117/14S, 19.3. 1952 H, De Saeger, 3199” (2 exs. MRAC). P.N.G. II/fd/13, 5.5. 1952 H, De Saeger, 3421” (1 ex. MRAC). – Non-African records: Yemen: “Wadi am Rija W Lahj Al Hulah by road, 13.01.57N, 44.33.30E (GPS) 25-26.10. 2007, 297 m a.s.l., Reitter leg.” (7 exs. NMPC, 2 exs. MZH).

Specimens with unclear labelling

Two specimens in NHRS labelled “Egypten” also belong to this species. The material is fairly old and the exact location of these records is somewhat unclear. The specimens are therefore not included in type material.

Diagnosis

Laccophilus inobservatus is closely related especially to L. continentalis, L. simplicistriatus and L. taeniolatus. The species can be distinguished by study of penis apex-shape, which is peculiar and stable in all four species. Penis apex of L. inobservatus is cut off straight and lacks any signs of anterior processes – resembling species have all at least minor kinds of modifications/processes on penis apex.

Description

Body length 3.6–4.0 mm, width 1.9–2.2 mm. Colour pattern dorsally, reasonably uniform; rarely reduced so that elytral irrorations are in part fragmentary (Figs 406–408).

Head: Pale ferrugineous, lacks darker areas. Slightly mat, finely microsculptured. Reticulation double. Large meshes only slightly more strongly developed in comparison with small meshes. Large meshes may contain 2–6 small meshes. Impunctate, except at eyes, with fine, irregular punctures.

Pronotum: Pale ferrugineous. At foremargin between eyes with a dark ferrugineous to blackish, slightly vague marking. At base in middle with two narrow, blackish spots. Sometimes dark areas on pronotum may be reduced. Slightly mat, finely microsculptured. Reticulation double. Large meshes only slightly more strongly developed than small meshes. Large meshes may contain 2–6 small meshes. Impunctate, but at margins except basally in middle with very fine scattered punctures.

Elytra: Pale ferrugineous, with dense, blackish to dark ferrugineous irrorations. Irrorations generally quite evenly distributed; sometimes irrorations reduced and at least in part separate irrorations rudimentary. Posterior to middle irrorations can be strongly reduced forming an irregular pale spot on each elytron (Figs 406–408). Slightly mat, finely microsculptured; reticulation double. Large meshes only slightly more strongly developed than small meshes. Large meshes contain 2–6 small meshes. Punctures very fine, sparse and irregularly distributed; on disc, irregular punctures form a vague row of punctures. Pre-apical, lateral row of punctures form a shallow furrow provided with hairs.

Ventral aspect: Pale ferrugineous, abdomen distinctly darker; dark to blackish ferrugineous. Almost impunctate, except apical ventrite; with some, fine, irregular punctures. Apical ventrite lacks lateral knob (Fig. 51). Rather shiny although finely microsculptured; microsculpture in part indistinct and reduced. Ventrites with fine, slightly curved striae. Metacoxal plates with some 10 almost transversely located, shallow and in part reduced, furrows. Prosternal process slender, posteriorly distinctly extended, apex pointed.

Legs: Pro- and mesotarsus somewhat extended and enlarged, provided with distinct suckers.

Male genitalia: Extreme penis apex, blunt, abruptly broken and lacks any kinds of processes (Fig. 251).

Female: Pro- and mesotarsi slender. Apical ventrite as in Fig. 52.

Etymology

The species name inobservatus is a Latin adjective meaning “unobserved”. It here refers to the peculiar situation that the species remained overlooked for a long time due to misinterpretation, although it is widespread and common.

Distribution

Africa: Gambia, Senegal, Mali, Niger, Sudan, Chad, Ethiopia, Burkina Faso, Ivory Coast, Ghana, Nigeria, Cameroon, Zaire and Asia: Yemen (Fig. 537). Only personally verified records are included in the map. Egypt is also among examined material but exact location of this record is unknown and therefore record is not mapped.

Collecting circumstances

Label data provide some information on the living habits of L. inobservatus. Accordingly, in Gambia collected at light at a bamboo pool and in semiarid vegetation near a river. Moreover the species has been collected in and at a fresh water stream and in a rock pool in river bed.

Laccophilus simplicistriatus Gschwendtner, 1932

Figs 53–54, 252, 409, 538

Laccophilus simplicistriatus Gschwendtner 1932b: 260 (original description, faunistics); Gschwendtner 1935a: 16, 17, 18 (description, faunistics); Gschwendtner 1938a: 5 (faunistics); Guignot 1943: 99 (faunistics); Guignot 1946c: 271, 273, 279, 280, 281, 312 (faunistics, biology); Guignot 1948: 15 (faunistics); Balfour-Browne 1950: 360 (faunistics); Guignot 1951: 215 (faunistics); Guignot 1952a: 533 (faunistics); Guignot 1952c: 522 (faunistics, biology); Guignot 1953b: 234 (faunistics); Legros 1954: 268 (faunistics); Guignot 1954: 26, 27 (faunistics, discussion); Guignot 1956b:219 (faunistics); Omer-Cooper 1956: 21 (faunistics, biology); Omer-Cooper 1957: 20, 21 (description, discussion); Omer-Cooper 1958b: 37, 43, 45 (description, discussion, faunistics, biology); Legros 1958: 211 (faunistics); Guignot 1959a: 570, 571, 573 (description, faunistics); Omer-Cooper 1965: 81, 82 (description, discussion, faunistics); Omer-Cooper 1970: 289, 290, 291 (description, discussion); Legros 1972: 466 (faunistics); Bilardo and Rocchi 1987: 104 (faunistics, biology); Curtis 1991: 186 (faunistics); Nilsson and Persson 1993: 81, 94 (discussion, biology); Nilsson 2001: 251 (catalogue, faunistics); Nilsson 2015: 218 (catalogue, faunistics).

Laccophilus monas Guignot 1953a: 238 (original description, faunistics); Guignot 1954: 27 (description, faunistics); Guignot 1959a: 571 (list as synonym of L. simplicistratus); Guignot 1959d: 162 (discussion, faunistics); Omer-Cooper 1965: 82 (list, synonymy); Nilsson 2001:251 (catalogue, list, synonymy, faunistics); Nilsson 2015: 218 (catalogue, list, synonymy, faunistics). Confirmed synonym.

Type localities

Laccophilus simplicistriatus: Zaire: Lusindol.

Laccophilus monas: Zaire: Route Shangugu-Usumbura, riv. Lua.

Type material studied

(12 exs.). Laccophilus simplicistriatus: Lectotype (by present designation): male: “Paratypus / Musée du Congo Lusindol 15-VIII-1911 L. Burgeon / R. Det. 2093 C” (MRAC; habitus in Fig. 409). – Paralectotypes: Same data as lectotype but “7-VIII-1911” (1 ex. MRAC); “Paratypus / Musée du Congo Albertville 20-X-1925 Dr. H. Schouteden / R. Det. 2093 C” (1 ex. MRAC); “Paratypus / Musée du Congo Karemi V-1912 Dr. Bayer / R. Det. 2093 C / Laccophilus simplicistriatus Gschw. det. Gschwendt.” (1 ex. MRAC); same data as preceding but no determination label (1 ex. MRAC); “Musée du Congo Haut Uelé: Moto 1920 L. Burgeon / R. Det. 2093 C” (2 exs. MRAC); “Musée du Congo Riv. Lobozi 5.11. 1912 Dr. Stappers 1548/ R. Det. 2093 C” (1 ex. MRAC); “Musée du Congo Bavrengura Haut Uelé L. Burgeon / R. Det. 2093 C” (1 ex. MRAC); “Musée du Congo Katanga: Katompe 1/15-VI-1930 Dr. P. Gerard / R. Det. 2093 C” (1 ex. MRAC); “Musée du Congo Kil. 345 de Kindu, nuit Dr. Russo / R. Det. 2093 C” (1 ex. MRAC). [Comments: no clarification exists why five of the specimens above have earlier been provided with a paratype label? Regarded as a case of mislabeling.]

Laccophilus monas: Holotype: male: “Holotypus / I.R.S.A.C. –Mus. Congo/Route Shangugu-Usumbura riv. Lua 5-VIII 1949 G. Marlier / Type / Eaux thermals rivier Lua 5-8-49/R Dét. I. 6182 / Guignot det., 1953 Laccophilus monas Guign. Type, male/= simplicistriatus Gschw. det. J. Omer-Cooper May 25th 1954” (MRAC).

Additional material studied

(396 exs.). Sudan: “Meya Saku 43 mi. from Amadi, Juba rd. 29.I. 1954 JOC” (2 exs. AMGS); “L. Yirol 6,33N, 30,3E 24.I. 1954 JJOC” (2 exs. AMGS); “Nimule, Fula rapids 4.XI. 1954 JOC.” (1 ex. AMGS); “Nimule Ferry 4.XI. 1954 JJOC.” (3 exs. AMGS); “L. Nyibor 23.I. 1954 JJOC.” (3 exs. AMGS); “Sandy river 50 mi. NW of Juba JJOC.” (5 exs. AMGS); “Aluakluak 30,5E, 6,30N 15.IV. 1954” (2 exs. AMGS); “Stream from hot springs Nyangwara 30,5E, 4,39N 29.I. 1954 JJOC.” (3 exs. AMGS); “Equatoria Tali Post 8.IV. 1954” (1 ex. AMGS); “L. Shambe 21.I. 1954 JJOC.” (3 exs. AMGS); “Upper Nile, Malakal 5-20.1. 1963 Linnavuori / ad lucem” (4 exs. MZH); “Upper Nile, Malakal 5-20.1. 1963 Linnavuori” (10 exs. MZH); “Upper Nile Malakal Linnavuori” (1 ex. MZH); “Upper Nile Pr. Malakal, nr junction Nile – Sobat 21.9. 1957 Forsberg/L. simplicistriatus Gschw. det. Nilsson 1996” (1 ex. MZLU); “Blue Nile Ingessana Mts. 17-22.11. 1962 Linnavuori” (2 exs. MZH); “Weisser Nil bei Tonga 10-13.4. 1914 Ebner”(1 ex. NMW); “Mongalla 50 Werner” (2 exs. NMW); “Kadugli at light, 11. 1954 Sweeney” (1 ex. BMNH); “Torit 2.7. 1980 Armstrong” (11 exs. USNM, 2 exs. MZH); “Gilo water tank (pumped of from stream) 20.3. 1980 Armstrong” (13 exs. USNM, 3 exs. MZH); “Kinyetti Riv. at Imeila 19.3. 1980 Armstrong” (4 exs. USNM); “SW Sudan nr Yambio Abbott” (1 ex. USNM); “Nairege Riv. 27.2. 1980 Armstrong” (1 ex. USNM). – Ethiopia: “Arsi, Dehra 40 km N Assella 25.9. 1988 1800 m, temp pool, Persson / L. simplicistriatus Gschw. det. Nilsson” (2 exs. MZLU); “Shoa, Soddere 16.10, 1988, 1500 m, temp. pool without vegetation Persson / L. simplicistriatus Gschw. det. Nilsson” (1 ex. MZLU); “Shoa, Soddere 25.9. 1988, 1500 m Persson / L. simplicistriatus Gschw. det. Nilsson” (2 exs. MZLU); “Shoa Dobre Zeit Hora lake 15.3. 1989, 2200 m, polluted water Persson /L. simplicistriatus Gschw. det. Nilsson” (1 ex. MZLU); “7000 ft. Hora Harsadi Addas 2.XII. 1926 J. Omer-Cooper” (1 ex. AMGS); “West marsh L. Zwai 5500 ft. 2-3.XI. 1926 JOC.” (2 exs. AMGS); 5000 ft. small pond Hora Shala 21.XI. 1926 JOC.” (1 ex. AMGS); “7000 ft. Mt. Chilalu 8.XI. 1926 JOC.” (2 exs. AMGS); “Baher Dar 8.10.1968 Horde leg. /Lichtfang” (7 exs. NHMB, 2 exs. USNM); “Bahar Dar, at light 4.4. 1967 P. Stys leg.” (1 ex. NMPC). – Zaire: “Kigoma V. 1930 / Paratype / L. simplicistriatus Gschw. det. Gschwendtner” (1 ex. OLML; not type material); same as preceding but no determination label (1 ex. MRAC; not type material); “Karemi V- 1912 Dr. Bayer / L. simplicistriatus G. J. Balfour-Browne det. 1963” (3 exs. MRAC); “Riv. Lobozi 5.11.1912 / L. taeniolatus Rég. var. R. Peschet det. 1914“ (1 ex. MAC); “Kivu: Luvungi XII-1932” (2 exs. MRAC); “Elisabethville (a la lumière) X/XI-1950 / L. simplicistriatus Gschw. det. Guignot 1953” (1 ex. MRAC); “PNG I/c/4, 15.3. 1950 Demoulin 234 / Paratype / L. monas Guign. det. Guignot” (1 ex. IRSNB; not type material); “PNG Napokomweli 18.X. 1950 G. Demoulin 893 / L. monas Guign. det. Guignot 1957” (1 ex. AMGS); “PNG, I/a/2, 21.4. 1950 Demoulin 452” (1 ex. NHMB); “PNG, Ndelele 19.3. 1952, 3199” (1 ex. MRAC); “PNG II/fd/12, 10.3. 1952, 3180” (2 exs. MRAC, 1 ex. MZH); “PNG PpK/14/g/14s, 4.4. 1952, 3290” (3 exs. MRAC, 1 ex. MZH); “PNG II/fd/14s, 3.4. 1952, 3278” (1 ex. MRAC); “PNG II7fd/Gar 29.2. 1952, 3152” (1 ex. MAC). – Uganda: “Mabira Forest Tinga 19.7. 1970 Brown” (1 ex. BMNH). – Kenya: “Lambwe Valley on light 11.6.1974 van Etten” (1 ex. RMNH); “Aberdares NP 5.12. 1989 Jäch” (1 ex. NMW); “Thika 7.12. 1989 Jäch” (2 exs. NMW, 1 ex. MZH); “Meru Distr., Gatunga 5.4. 1987 Mourglia” (1 ex. NHMB). – Rwanda: “Rumonge, Regenwald Jan. 1986 Heiss” (1 ex. NHMB). – Tanzania: “Ukerewe Tang. Terr. VIII. / L. simplicistriatus Gschw. det. Gschwendtner” (4 exs. OLML); “TPC S of Moshi canals 28.9. 1976 Holmen” (1 ex. ZMUC); “Mwanza nr. Lake Victoria 31.7. 1957 / sweet potato channels” (2 exs. BMNH); “Mwanza nr. L. Victoria 1957 / Marginal pools and ditches” (7 exs. BMNH); “Tanganyika 1959 Eccles” (1 ex. BMNH); “SW Tanganyika Mpanda (dans ruisseau) 6. 1960 Leleup” (1 ex. MAC); “T.T. Rukwa Milepa 25.4. 1951 Water in road- tracks Backlund / L. simplicistriatus Gschw. det. Nilsson 1996” (1 ex. MZLU); “T.T. Rukwa Tumba 29.1. 1951 Backlund / L. simplicistriatus Gschw. det. Nilsson 1996” (1 ex. MZLU); “T.T. Rukwa Tumba 12.1. 1951 T. river Backlund / L. simplicistriatus Gschw. det. Nilsson 1996” (1 ex. MZLU); “T.T. Rukwa Kipangati 28.11. 1950 sulphurous pools, shallow in rich woodland Backlund / L. simplicistriatus Gschw. det. Nilsson 1996” (1 ex. MZLU); “Tang. terr. Nzega, Naro 19.8. 1951 Backlund / L. simplicistriatus Gschw. det. Nilsson 1996” (1 ex. MZLU); “Rukwa, Rungwa Riv. 18.5. 1950 Backlund / L. simplicistriatus Gschw. det. Nilsson 1996” (1 ex. MZLU); “T.T. Shinyanga 21.2. 1935 Burtt” (1 ex. BMNH); “T.T. Rukwa Mkumbwa 12.5. 1950 Backlund / L. simplicistriatus Gschw. det. Nilsson 1996” (1 ex. MZLU); “Iringa Prov., 100 km NE Iringa 07°37'S, 36°17'E, 9.1. 2007, 660 m, J. Halada leg.” (2 exs. NMPC, 1 ex. MZH); “Mbeya prov., 120 km E Mbeya 08°51'S, 34°00'E, 1220 m, 6.1. 2007 m, J. Halada leg.” (1 ex. NMPC). – Angola: “Namakunda 6. 1948, 16.15E, 18.50 S Koch” (13 exs. BMNH, 1 ex. MZH); “Namakunda 6. 1948, 16.15 E. 1850S, Koch” (11 exs. BMNH); “Mongua 4.6. 1954, shallow reedy vlei” (1 ex. BMNH); “Mossamedes Distr., Rio Coroca 23.6. 1954/small clear pool with Chara (1 ex. BMNH); “Rio Coroca 8 m. N of Porto Alexandre 22-23.6. 1954/Pond with Algae & Lemna, fringing Juncus (1 ex. BMNH); “Pediva, ca, 30 mi. E of Porto Alexandre 400 ft. 26-27.6. 1954 / Ponds in warm, saline river; thick weed” (1 ex. BMNH); “Angola Schönlein” (1 ex. ZMHB). – Zambia: “Central Pr. Lusaka 8.1. 1982 Selander / rain pond” (1 ex. MZH); ”29.3.1993. Kafue NP., Chunga Camp, 15°02'35"S/26°00'09"E, lux Uhlig” (1 ex. ZMHB). – Malawi: “Stream 20 mi. from Dedza on Lower Lilongwae rd 30.IX. 1948” (8 exs. AMGS); “R. Diedma Lilongwe rd 30.IX. 1948” (1 ex. AMGS); “Bua R. 2.X. 1948 JOC.” (1 ex. AMGS); “Dallys Hotel nr. Ft. Johnstone 23.VIII. 1948” (3 exs. AMGS); “Zomba plateau res.7.XI. 1948” (3 exs. AMGS); “Stream, Zomba plateau 6000 ft. 7.XI. 1948” (1 ex. AMGS); “Gomba plateau (?) 7.XI. 1948” (3 exs. AMS); “Stream 6 mi. N of R. Mtiti 2.X. 1948” (1 ex. AMGS); “Swampy pool nr. L. Nyasa 9.6. 1946” (1 ex. BMNH). – Zimbabwe: “Stream at Salisbury17.IX. 1948” (2 exs. AMGS); “Marandellas 2. XI. 1948 JOC.” (2 exs. AMGS); “Small stream nr. Halfway Hotel Salisbury-Gatooma 14.IX. 1948” (2 exs. AMGS); “Wankie Reserve water holes 3.IX. 1948” (5 exs. AMGS); “Wankie Game Res. IX. 1948 water holes” (10 exs. AMGS); “Wankie Game Res. IX. 1948 water hole / L. simplicistriatus Gschw. det. Omer-Cooper” (3 exs. AMGS); “Wankie Game Res. Mazume Dam 4.IX. 48 / L. simplicistriatus Gschw. det. Omer-Cooper” (5 exs. AMGS); “Wankie game reserve, Shapi pan 5-6.IX. 1948 (7 exs. AMGS); “Wankie Game Reserve 4.9. 1948 / J. OmerCooper” (1 ex. BMNH); “Wanki Game Reserve 4-5.9. 1948 J. Omer-Cooper / L. simplicistriatus Gschw. det. J. Omer-Cooper”(4 exs. NHMB, 9 exs. USNM); ”5 mi SE Wankie 7.4. 1968 Spangler” (9 exs. USNM, 3 exs. MZH); “Victoria Falls rainforest 6.X. 1948” (2 exs. AMGS); “Gwai River 3.4. 1968 Spangler” (7 exs. USNM, 1 ex. MZH). – Mozambique: “Niassa Prov., S12°17'28.8”, E34°46'31.4” Mandambuzi Marsh, Watson 6.4. 2009” (1 ex. CGF); “Niassa Prov. Cmimulimuli River, S12°11.520’, E34°42.288’, Watson 10.2. 2008” (1 ex. CGF). – Namibia: “Windhoek Town Dam 7.VII. 39” (1 ex. AMGS); “Okahandja Distr. Toggekry 250, Omatako Ranch, 55 km NNW, NNW Okahandja, thornbush savannah / 7.2. 2001, 21°30'43"S/16°43'00” lux 22°-14°, 25.4. 2001, 17,45-20,00 Uhlig & Ebert” (1 ex. ZMHB, 1 ex. NMNW, 1 ex. MZH); “Damaral. Okahandja 21.59S-16.52E / 12.9. 1974 shore washing, Endrödy-Younga” (2 exs. TMSA, 1 ex. MZH); “Damaraland Oshikango 5. 1948 15.55 E 17.25 S Koch” (1 ex. BMNH); “Kavango:Popa Falls 18°07'S-21°35'E, 26.2.-3.11.1992 lux Uhlig leg.” (1 ex. ZMHB); “Ovamboland Namutoni 31.5. 1954 / weedy waterhole and stream” (16 exs. BMNH, 2 exs. MZH); “Oshikango SE of frontier post 2.6. 1954/shallow water, svampy marsh” (27 exs. BMNH, 2 exs. MZH); “Ca. 7 mls N.E. of Grootfontein / waterhole in dolomite” (4 exs. BMNH); “Okarupa, ca. 17 mi. E of Okahandja, 4900 ft, 22.5. 1954 / pools in overflow stream from dam, much weed & algae” (2 exs. BMNH); “Kro, ca. 15 mi. SE of Namutoni 30.5. 1954 / shallow & muddy with algae” (1 ex. BMNH); “Etosha Pan Okaukujo camp 19.11S-15.55E/28.12. 1974 shore washing Endrödy-Younga” (1 ex. TMSA); “Etosha Pan, 60 m. NW Namutoni 5. 1937”(1 ex. TMSA); “Etosha Game Res., Namutoni 27.5. 1937” (3 exs. TMSA, 1 ex. MZH); “Kaokoweld Kowares, 90 mi SE Ohopoho 3.6. 1951” (1 ex. MZLU); “Kaokoweld Sanitatas, abt 85 mi. WSW Ohopoho 14-16.6. 1951” (1 ex. MZLU); “Kaokoweld Anabib (Orupembe) 100 mi. W Ohopoho 12-13.6. 1951” (1 ex. MZLU); “Kaokowelld Omutati, 70 mi. WSW Ohopoho 5.6. 1951” (2 exs. MZLU); “Kaokoweld 17.10. 1963 Gaerdes” (1 ex. MZLU); “Kaokoweld Sesfontain, 17 km WSW, 19.12S-13.32E/1.2. 1975 singled in riv. bed, Endrödy-Younga” (1 ex. TMSA); “Distr. Grootfontein leg. Irish / Farm klein Nosib 19.28S-14.50E Anfang April 1989” (1 ex. ZMHB); “Okomite R, temp. pool, N-17.4305, E14.1666, 12.11. 1997 De Moor” (5 exs. AMGS); “Omapapurawe Guard Post, 200 m from campsite, Kunene R., N-17.218, E13.645, 15.11. 1997 Bethune & al.” (3 exs. AMGS); “Kunene R., stream from cave into pool, N-17.00.07, E12.59.54, 20.6. 1997 De Moor & al.” (1 ex. AMGS). – Botswana: “Tsotsorogo Pan 17.VI-9. VIII. 1930 / Type male / female / paratype / L. simplicistriatus Gschw. det. Gschwendtner” (2 exs. AMGS, 2 exs. TMSA, 1 ex. OLML; not type material); “Chobe Park Savuti Camp11.3. 1993, 18°33'S/24°03'E, lux Göllner” (1 ex. ZMHB); ”5 km NW San-ta-Wani Safari Lodge 19°27'01”/23°38'46”lux, Uhlig” (1 ex.ZMHB). – South Africa: “Trsvl 5 mi W Warmbad 24-25.2. 1968 Spangler” (1 ex. USNM); “Trsvl Randburg, N-26.070, E27.950, 6.6.1971 Reavell” (1 ex. AMGS); “Caffraria / J. Wahlb.” (1 ex. RMS); “Gauteng Cullinan Premier Mine Res. 25,40S–28,29E / 17.1.2002 Endrödy-Younga, light trap” (1 ex. TMSA); “Gauteng Tswaing 25.24S-28.06E / 16.2. 2003 light trap, TMSA staff leg.” (1 ex. TMSA); “Roodeplaat Pretoria distr. 10. 1960 Neubecker” (1 ex. TMSA); “Xolo R, small stream, riverbed, trib. Kunene R, 15.11. 1997 De Moor” (1 ex. AMGS). – Swaziland: “Eranchi 5-10.1. 1955 Capener / L. simplicistriatus J. Omer-Cooper det.” (3 exs. MZH). – Lesotho: “Nazareth M. S., 20 mi.ESE Maseru 24.3. 1951” (1 ex. MZLU).

Specimen with uncertain determination

Cameroon: female “Yaounde, Bor to Kosti by boat 13-14.3. 1978 Perkins” (1 ex. USNM).

Specimen with uncertain labelling

Mauritius: male “Insel Mauritius Westw. Nr. 9984 / L. simplicistriatus Gschw. det. Brancucci 82” (1 ex. ZMHB). Until additional specimens from Mauritius are available, this record is considered a case of mislabelling.

Comments on synonymy

Confusion regarding the original description and the type material of L. simplicistriatus followed when Gschwendtner (1932b) in a faunistic paper listed and mentioned L. simplicistriatus and at the same time he provided a short description of the species. Originally this act was not ment to be the original description, which it is in fact. Accordingly, the report (Gschwendtner 1932b) is the original description and not the later publication of Gschwendtner (1935a). Designation of type in the later article is accordingly invalid. A lectotype has been chosen from the valid type material of L. simplicistriatus and it has been compared with the holotype of L. monas. This examination confirms earlier proposed synonymy of the two taxa.

Diagnosis

Laccophilus simplicistriatus externally resembles most of L. taeniolatus and L. complicatus. From L. taeniolatus it is distinguished by elytral irroration, which is complete (uniform coverage) and not reduced frontally at suture. In L. taeniolatus elytral irroration frontally at suture is always sparser, often formed as a pale somewhat irregular spot. Shape of penis apex exhibits differences separating L. simplicistriatus from L. complicatus (apex of penis sharp and strongly curved) as well as from all other Laccophilus species.

Description

Body length 3.6–4.3 mm, width 1.9–2.4 mm. Habitus, dorsal aspect as in Fig. 409. Dorsal colour pattern quite uniform exhibiting only minor variation.

Head: Pale ferrugineous. Finely microsculptured; reticulation indistinctly double. In part, coarser meshes not discernible. When visible one large mesh contains 2–6 fine meshes. At eyes with fine, irregular punctures; a few scattered punctures medially between eyes.

Pronotum: Pale ferrugineous, anteriorly and posteriorly in middle with narrow ferrugineous to dark ferrugineous area. At margins, except basally in middle, with fine, irregular punctures. Microsculpture fine; reticulation indistinctly double (only in part, discernible). Slightly coarser, large meshes contain 2–6 fine meshes.

Elytra: Pale ferrugineous, with quite coarse and distinct, dark ferrugineous irrorations (Fig. 409). Irrorations laterally sometimes slightly reduced and less distinct than at suture. Slightly mat, finely microsculptured; double reticulation indistinct, only in part discernible; coarser meshes extensively rudimentary and hardly visible. Very fine, sparse punctures laterally and discally (form an irregular, discal row of punctures) discernible. Lateral, pre-apical furrow fine, finely pubescent.

Ventral aspect: Ferrugineous, apical half of abdomen dark ferrugineous. Rather shiny, although extensively, very finely microsculptured. Almost impunctate. Metacoxal plates with some transversely located, shallow furrows. Abdomen with fine striae. Prosternal process narrow, apically pointed. Apical ventrite not distinctly asymmetric (no process or knob on one side) (Fig. 53).

Legs: Pale ferrugineous, hindlegs a little darker. Pro- and mesotarsus rather slender, claws not especially long, slightly curved. Pro- and mesotarsus with suckers.

Male genitalia: Penis in lateral aspect, with apical half, somewhat enlarged on both sides of apex; extreme apex with two small processes (Fig. 252).

Female: Externally as male but apical ventrite less impressed on both side of midline and apex more extended backwards and rounded (Fig. 54).

Distribution

Sudan, Ethiopia, Zaire, Uganda, Kenya, Rwanda, Tanzania, Angola, Zambia, Malawi, Zimbabwe, Mozambique, Namibia, Botswana, South Africa, Swaziland, Lesotho (Fig. 538). Uncertain female record from Cameroon. One male record from Mauritius is considered a probable case of mislabelling.

Collecting circumstances

Very little, detailed information is available on the biology of L. simplicistriatus. In Malawi L. simplicistriatus was e.g. collected in a reservoir surrounded by a marshy area with small areas of open water and red mud and from a swift river with a wide coarse gravel bed where the species was collected in a pool. Additionally from Malawi, the species was sampled in a clear river with white water lilies, reed beds and patches of swamp. In Zimbabwe the species was collected in a number of water holes, springs and dams used by the game, and also in a dam with reeds and water plants, the bottom largely covered with dark mud, but sand in places. Finally, also in Zimbabwe collected in a series of pools with a small stream connecting them. The bottom of the water body was rock, sand and gravel with some deposited mud. Some vegetation growing in the pools. Also recorded from streams (Omer-Cooper 1958b). Data of collection labels is also rather scarce and simply describe method of sampling or kind of water body; light collection or temporary pool, stream from hot spring, pond in warm, saline river etc.

Laccophilus taeniolatus Régimbart, 1889

Figs 56–57, 253, 410–412, 539

Laccophilus taeniolatus Régimbart 1889: 52 (original description, faunistics); Severin 1892: 472 (incorrect type deposition); Régimbart 1894: 237 (description, faunistics); Régimbart 1895: 136, 137 (description, discussion, faunistics); Sharp 1904: 3 (faunistics); Régimbart 1904: 66 (faunistics); Régimbart 1906: 248 (description variation, faunistics); Régimbart 1908: 5 (faunistics); Zimmermann 1919: 122 (faunistics); Zimmermann 1920a: 26 (catalogue, faunistics); Peschet 1920: 250 (discussion, faunistics); Régimbart 1922: 532 (faunistics); Peschet 1922: 374 (faunistics); Zimmermann 1926: 23 (faunistics, description); Gschwendtner 1930: 88 (faunistics); Gschwendtner 1931: 180 (faunistics); Omer-Cooper 1931: 757 (biology, description, faunistics); Gschwendtner 1932b: 260 (discussion); Gschwendtner 1935a: 16, 17, 18 (description, discussion, faunistics); Guignot 1943: 99 (faunistics); Guignot 1946c: 271, 273, 278, 280, 312 (description, discussion, faunistics); Guignot 1951: 215 (faunistics); Guignot 1952a: 533, 535 (discussion, faunistics); Capra 1952: 6–7 (faunistics); Guignot 1954: 29 (faunistics); Guignot 1955d: 67 (discussion); Guignot 1955g: 865 (faunistics); Guignot 1956b: 219 (faunistics); Omer-Cooper 1957a: 20, 21 (description, discussion, incorretc association); Omer-Cooper 1958b: 43, 45 (description, discussion, incorretct assocation); Guignot 1959a: 570, 573, 574, 575 (description, faunistics, incorrect association); Bruneau de Miré and Legros 1963: 874, 883 (faunistics); Omer-Cooper 1965: 81, 82 (list); Omer-Cooper 1970: 288, 289, 290 (description, discussion); Legros 1972: 466 (faunistics, list.); Medler 1980: 155 (faunistics, list.); Forge 1981: 500, 501 (description, faunistics); Nilsson and Persson 1993: 58, 81, 88, 94 (biology, discussion, faunistics); Nilsson et al. 1995: 506 (discussion, faunistics); Nilsson 2001: 251 (catalogue, faunistics); Bilardo and Rocchi 2002: 174 (faunistics, list); Reintjes 2004: 68 (faunistics); Aistleitner and Jäch 2014: 47, 49 (faunistics, biology); Nilsson 2015: 218 (catalogue, faunistics). [Comment: This species was quite early misinterpreted and accordingly, any information in articles listed above on L. taeniolatus should be considered carefully before acceptance.]

Laccophilus congener Omer-Cooper 1957a: 19, 21, 90 (original description, discussion, faunistics); Omer-Cooper 1958a: 59 (faunistics); Omer-Cooper 1958b: 37, 43, 44, 45, 46 (biology, description, discussion, faunistics); Guignot 1959a: 570, 571, 573 (biology, description, discussion, faunistics); Omer-Cooper 1962: 295 (faunistics); Omer-Cooper 1965: 76, 81, 82 (description, discussion, faunistics); Omer-Cooper 1970: 288, 289, 290 (description, discussion); Bilardo and Pederzani 1978: 119 (description, faunistics); Medler 1980: 155 (faunistics, list); Pederzani and Rocchi 1982: 72 (faunistics); Bilardo 1982a: 447 (description, faunistics. Spelled Laccophylus); Pederzani 1988: 107 (biology, faunistics); Bilardo and Rocchi 1990: 162, 177 (biology, faunistics); Franciscolo and Sanfilippo 1990: 145 (biology, description, faunistics); Curtis 1991: 186 (faunistics); Nilsson et al. 1995: 506 (faunistics); Rocchi 2000: 24 (faunistics); Nilsson 2001: 242 (catalogue, faunistics); Bilardo and Rocchi 2002: 156, 161, 162, 174 (faunistics, list); Reintjes 2004: 67 (faunistics, list); van Vondel 2005: 130 (biology, faunistics); Bilardo and Rocchi 2006: 130 (faunistics); Bilardo and Rocchi 2013: 141 (faunistics, biology); Nilsson 2015: 210 (catalogue, faunistics). New synonym.

Type localities

Laccophilus taeniolatus: Angola: Humpata.

Laccophilus congener: South Africa: Transvaal, Belfast.

Type material studied

(12 exs.). Laccophilus taeniolatus: Holotype: male: “P.J. vd Kellen Humpata Afr. trop. / Laccophilus taeniolatus sp. n. type Régb. / taeniolatus sp. n. Régimb.” (RMNH; habitus in Fig. 410). [Comment: type material contains only one specimen.]

Laccophilus congener: Holotype: male: “Type / Transvaal Belfast pond 23. N. 1948 Omer-Cooper / Brit. Mus. 1957-660 / L. congener O-C.” (BMNH). – Paratype: female: Same information as holotype but labelled as “Allotype” (1 ex. BMNH); additional paratypes: same information as holotype but labelled as “Paratype” (4 exs. AMGS); “Paratype / Transvaal Belfast 30 Dec. 1948 J. Omer-Cooper” (1 ex. AMGS); “Paratype / Transvaal Standerton 8.12. 1948 J.O-C. /L. congener O-C. det. J. Omer-Cooper” (1 ex. AMGS); “Paratype / Transvaal R. Nyl at Num Num 23 Aug. 1948 J.O-C. (1 ex. AMGS); Paratype / Transvaal Misselburg 29 N. 1948” (2 exs. AMGS).

Specimens with type status uncertain

(not given in original description) (2 exs.): “Paratype / Transvaal Deel Kraal 20.8. 1948 J. O-C.” (1 ex, AMGS); “Paratype / Transvaal Poerzya R., Waterberg Distr. 19.8. 1948” (1 ex. AMGS).

Additional material studied

(457 exs.): Gambia: “Stream N of Selety 19.2.1976 Holmen leg.” (2 exs. ZMUC, 1 ex. MZH); “Bathurst Jan. 1968 Leiler T.” (1 ex. NHRS); “Outside Abuko Nature Reserve at water works, at light 18.30–21.00, 4.11. 1977 / Cederholm-Danielsson-Hammarstedt-Hedquist-Samuelsson” (1 ex. MZLU); same data but ”19.00–22.00, 26.11. 1977” (1 ex. NHMB); same data but ”in and at Lamin stream 25–26.11. 1977” (1 ex. NHMB); “Abuko Nature Reserve, at light at the bamboo-pool 18.30-20.30, 18.11. 1977 / Cederholm-Danielsson-Hammarstedt-Hedquist-Samuelsson” (1 ex. MZLU, 1 ex. NHMB); “2 km S Kitty, 7 km SSW Brikama rd., junction in and at fresh water stream 13.2. 1977 / Cederholm-Danielsson-Hammerstedt-Hedquist-Samuelsson” (1 ex. MZLU). – Senegal: “Swamps ca. 3 km SW of Ziguinchor 8.3. 1977 / Cederholm-Danielsson-Larsson-Mireström-Norling-Samuelsson” (1 ex. MZLU); “In forest 1,5 km NE Djibélor, ca 6.5 km SW Ziguinchor, 8.3. 1977, at light 19.00-21.30 / Cederholm-Danielsson-Larsson-Norling-Samuelsson” (1 ex. MZLU); “Village Saré Sara 21 km ESE Kolda, in and at the junction of Rivers Koring and Tiángol, Dianguina 6.3. 1977 6.3. 1977 / Cederholm-Danielsson-Larsson-Norling-Samuelsson” (1 ex. MZLU). – Guinea Bissau: “Cacheu, 12 km E Varela, earth pit pond 9.4. 1993 S. Persson leg.” (8 exs. MZLU); “Cacheu, Bula, temporary pools 16.7. 1992 S. Persson leg.” (2 exs. MZLU); “Cacheu, 5 km W Bula, ponds 25.7. 1992 S. Persson leg.” (3 exs. MZLU). – Guinea: “Seredoux, lux 7-8.4. 1975 Zott” (14 exs. ZMHB, 1 ex. MZH); same data but ”4.5. 1975” (5 exs. ZMHB); same data but ”5.4. 1975” (2 exs. ZMHB); same data but ”4.4. 1975” (4 exs. ZMHB, 3 exs. MZH); same data but ”16.4. 1975” (1 ex. ZMHB); same data but ”18.4. 1975” (1 ex. ZMHB). – Burkina Faso: “Karfiguéla 29.10. 1973 Linnavuori leg.” (1 ex. MZH); “Niangoloko 26.10. 1973 Linnavuori leg.” (1 ex. MZH); “Ouagadougou 7.10-11. 1926” (1 ex. NHMB); “Nadiagow MV August 2005 Moretto” (1 ex. NHMB). – Chad: “Bebedja 28-31.5. 1973 Linnavuori leg.” (7 exs. MZH); “Nr Bongor 27.5. 1973 Linnavuori leg.” (1 ex. MZH). – Central African Republic: “Bambari UV 2. 1964 Pierrard” (1 ex. MRAC). – Sudan: “Equatoria, Yambio 18-25.4. 1963 Linnavuori leg.” (1 ex. MZH); “Dahr el Ghazal, Wau 19.2. 1963 Linnavuori leg.” (1 ex. MZH); “Dahr el Ghazal R. Malmul 21.2. 1963 Linnavuori leg.” (1 ex. MZH); “Aluoklua Riv. 30.5E-6.30N, 15.4. 1954 Reid T.” (1 ex. AMGS); “L. Shambe 21.1. 1954 Omer-Cooper” (1 ex. AMGS); “Alel R. Lau 14 mi.NE of Yirol 17.1. 1954 Omer-Cooper” (1 ex. AMGS); “L. Nyibor 23.1. 1954” (1 ex. AMGS); “Rain ponds S of Rumbek nr Wulu 19.1. 1954”(3 exs. ZMHB); “R. Maila 30.57E, 4.39N, 29.1. 1954” (5 exs. AMGS); “Minkammon 31.31E-6.2N, 16–17.1.1954 O-C.” (1 ex. AMGS). – Sierra Leone: “Makeni 28.11. 1993 light trap/Cederhalm-Danielsson-Hall” (4 exs. MZLU); same data but ”27.11. 1993” (7 exs. MZLU; habitus in Fig. 412); “Kalangba 8.11. 1980 Jump leg.” (1 ex. USNM, 1 ex. MZH); same data but ”9.11. 1980” (1 ex. USNM); same data but ”25.10. 1980” (4 exs. USNM). – Liberia: “Suakoko 26.4. 1952 light trap Blickenstaff” (1 ex. USNM); “Suakoko 8.4. 1953 Blickenstaff” (1 ex. USNM). – Ivory Coast: “Nord CI, Ferkessedougou 10-20.5. 1964 Decelle leg.” (1 ex. MRAC). – Ghana: “Upper East Pr. Navrongo Env. 11-13.6. 2006 Pokorny S. leg.” (3 exs. NMPC, 1 ex. MZH). – Togo: “Reg. Plateaux, Pref. Kloto, nr. Kpimé, 1 km NW Seva village 10.2.2006 Komarek & Hounguè leg. / 300 asl, agricultural irrigation ditch” (1 ex. NMW); “Sokodé- Kmpamgalam, FL HQ (niedere vegetat.) N-22.132 S, 4. 1980/Krell leg.” (1 ex. NHMB). – Benin: “Dép. Zou, commune de Zogbodomè 29.1. 2006 Goergen leg. / Lokoli forest 17 m asl, light trap” (1 ex. NMW); “Penessoulou, pond, forest area Oct. 2003 Goergen leg.” (1 ex. NMW, 1 ex. MZH); “SE Benin, 15 km SE Save 8–11.4.2000 Halada leg.” (1 ex. NMW). – Nigeria: “NW St. Badeggi rice fields 8-9.8.1973 Linnavuori leg.” (70 exs. MZH); “Ilorin Prov., Ilorin15–18.2 1949 Malkin / small clear pond” (3 exs. BMNH, 1 ex. MZH); “NW St. Yelwa 23.7. 1973 Linnavuori leg.” (5 exs. MZH); “W St. Igboho-Kishi 19.7.1973 Linnavuori leg.” (1 ex. MZH); “R St. nr Mbiama 4-5.7. 1973 Linnavuori leg.” (1 ex. MZH); “MW St. Sapoba forest 1.9.1973 Linnavuori leg.” (1 ex. MZH); “NE St. Gombe-Bauchi 27.8.1973 Linnavuori leg.” (1 ex. MZH); “Ibadan ca. Jan.-Juni 1954 Stenholt-Clausen leg. / L. congener O-C. J. Balfour-Browne det. 1961” (2 exs. ZMUC); “R. Kaduna at Kadoura ? 1964” (1 ex. AMGS); “Little stream Oyo-Ibadan 28.3. 1963 JOC.” (1 ex. AMGS); “Pools in dry stream bed Kontagora 5.4. 1963” (1 ex. AMGS); “Pools in river bed Kontagora 3.4. 1963 JOC.” (1 ex. AMGS); “Stream at Assob 36 miles from Jos 13.4. 1963” (2 exs. AMGS); “Stream reservoir Jos 10.4. 1963” (1 ex. AMGS); “Pond in stream bed Kontagora-Kaduna rd 5.4. 1963 JOC.” (1 ex. AMGS); “Stream 86 miles from Makurdi at Jos road 24.4. 1963 JOC.” (1 ex. AMGS); “Detritus pond, Jos-Bauchi rd 9.4. 1968” (3 exs. AMGS); “Stream, escarpment, rd Jos-Wambe 13.10. 1963 JOC.” (1 ex. AMGS); “Stream 64 miles from Bida on Jebba rd 15.IV. 1963 JOC.” (9 exs. AMGS); “Stream Zaria rd about 3 miles from Kaduna 4.4. 1963 JOC.” (1 ex. AMGS); “Stream nr Zaria 4.4. 1963 JOC.” (3 exs. AMGS); “Stream 3,5 miles from Oyo 28.3. 1963 JOC.” (10 exs. AMGS); “Trib. of R. Gagere (?) Zaria-Katsina 5.4. 1963 JOC.” (1 ex. AMGS); “Zaria 1969” (1 ex. NHMB); “Rivercrossing rd about 49 miles from Makurdi in Enyogo direction 24.4. 1963 JOC.” (4 exs. AMGS); “New Calabar River nr Port Harcourt 13.1. 1989 Umeozor leg.” (1 ex. USNM). – Cameroon: “SW Kumba-Mamfe 23.6. 1973 Linnavuori leg.” (2 exs. MZH); “Maroua 10/11.1965 (lumière) Schmitz leg.” (1 ex. MZH); “Boura, partially parched small stream with riverine forest in savannah, at light 13.1. 1978/Gärdenfors-Hall-Samuelsson” (1 ex. MZLU); “Obala Juin 1969” (2 exs. NHMB); “Tsang Monatiele 5.1. 1970” (1 ex. NHMB). – Gabon: “Lagune Iguelá, Fortét à Est. Tassi Gen. 97 Bilardo / L. congener O-C. det. Rocchi S. 1998” (1 ex. OLML). – Congo: “Brazzaville P.K. Rouge G. Onore 4. 1979 G. Onore / L. congener O-C. det. Rocchi S. 1998” (1 ex. OLML); “Djili P.K. Rouge 3. 1979 Onore” (2 exs. NHMB); “Loudima, sur la route en savane 20.3. 1980 Onore” (3 exs. NHMB). – Zaire: “Parc National Garamba 1.9.1952 De Saeger leg.” (1 ex. MRAC); PNG II/fc/13, 7.3. 1952 De Saeger 3257” (1 ex. IRSNB); “PNG 30.1. 1950 G. Demoulin 240” (1 ex. NHMB); same data but ”8.5. 1950, 494” (1 ex. NHMB); “PNG II/fd/12, 10.3. 1952 De Saeger 3180” (1 ex. NHMB); “Tshuapa-Mbandaka 1964, A.B. Stam leg.” (3 exs. RMNH); “Dilolo 8-9. 1931 de Witte” (2 exs. NHMB); “Haut Uele Moto 1920 L. Burgeon / L. congener O-C. det. Wewalka 1981”(1 ex. OLML); “Lusindoi 15.7. 1911 L. Burgeon/L. congener O-C. det. Wewalka 1981” (1 ex. OLML); “Elisabethville 1.1956-1.1957 à la lumière Seydel leg.” (1 ex. MRAC, 1 ex. MZH, 1 ex. NHMB); same but ”1957-1958”(1 ex. NHMB); “Katanga Kansenia 6. 1925 G.F. de Witte / L. congener O-C. det. Wewalka 1981” (1 ex. OLML). – Kenya: “Mariakani Dam, Kilifi distr. 16.9.1976 Holmen leg.” (1 ex. ZMUC); “Machakos district, Athi River 14.9.1976 Holmen leg.” (1 ex. ZMUC); “Djili P.K. Rouge 3. 1979 G. Onore” (2 exs., NHMB). – Tanzania: “Petukiza, ponds, Tanga Distr. 23.9. 1976 Holmen leg.” (1 ex. ZMUC); “Deforested place nr Mangula, 297 m, N-07°52'20”, E36°55'06”, 18.7. 2004, 297 m at light, Sprecher” (4 exs. AMGS, 1 ex. MZH). – Angola: “Pediva ca. 30 mi. E of Porto Alexandre 400 ft. 26-27.6. 1954 / Pools in warm, saline river, thick weed” (1 ex. BMNH); Mossamedes Distr. Rio Coroca 23.6. 1954 / Small clear pool in sand with Chara(1 ex. BMNH); “Rio Coroca 8 m N of Porto Alexandre 22-23.6. 1954 / Pond with Algae & Lemna, fringing Juncus (6 exs. BMNH, 1 ex. MZH); “Ca. 10 mls. W of Cainde, ca 3500 ft 15.6. 1954 / Stagnant water hole, nitellid algae and muddy silt” (2 exs. BMNH). – Zambia: “Central Prov., Lusaka 8.1. 1982 Selander leg. / rain pond” (2 exs. MZH); “Lusaka 8.5. 1974 Lange” (3 exs. CGC); same data but ”23.4. 1974” (1 ex. CGC); “Kafue NP Chunga Camp, 15.02.35S-26.00.09E, 29.3. 1993 lux Uhlig leg.” (2 exs. ZMHB, 1 ex. MZH); “Kackhalola 820 m, 14.45.43S-30.35.46E, 19.3. 1993 lux Uhlig leg.” (1 ex. ZMHB); “Copperbelt Pr. Mwekera 23.1. 1982 Selander leg. / rain pond” (2 exs. MZH); “Abercorn 13.4. 1951 H.O. Backlund leg. / L. propinquus O.-C. det. A.N. Nilsson 1996” (1 ex. MZLU); “Kasanka Nat. Res. N-12.30, E30.15, rain-filles dams Reavell” (4 exs. NHMB). – Malawi: “Fort Hill 8.10. 1945 JOC.” (1 ex. AMGS); “Mulanje Mts.,env. 22.-26.12. 2001 Kantner” (1 ex. NHMB); “Dambo below Livingstonia, lakeshuve 21.9. 1945” (1 ex. IRSNB). – Zimbabwe: “Wankie Res. waterhole 3.9. 1948” (5 exs. AMGS); “Wankie Game Res. 9. 1948 waterhole JOC: / L. congener det. J. Omer-Cooper” (10 exs. AMGS); “Wankie Game Res. Shapi Pan 5.9. 1948 / L. congener det. J. Omer-Cooper” (2 exs. AMGS); “Wankie Game Res. Shapi Pond 5.9. 1948” (1 ex. AMS); “Wankie Game Res. 4.9. 1948” (3 exs. AMGS); “Wankie Res., Masumu Dam 4.9. 1948” (3 exs. AMGS); “Wankie Res., Masumu Dam 9. 1948” (2 exs. AMGS); “Wankie Res., Masumu Dam 9. 1948, waterhole” (2 exs. AMGS); ”5 mi. SE Wankie 7.4. 1968 Spangler” (2 exs. USNM, 1 ex. MZH); “Marandellas 2.9. 1948 JOC. / L. congener det. J. Omer-Cooper” (1 ex. AMGS); “Sinkukwe 30.12. 1948 JOC.” (1 ex. AMGS); “Small stream, Halfway Hotel Salisbury-Gatooma 14.9. 1948” (3 exs. AMGS); “Gwaai River 1.9. 1948 JOC.” (1 ex. AMGS); “Gwai R. 3.4. 1968 Spangler” (4 exs. USNM). – Mozambique: “Niassa Prov. S12°17'28.8”, E34°46'31.4”, Mandambuzi Marsh Watson 6.4. 2009” (1 ex. CGF); “Jangamo Block N-24.3212, E35.280, 7.6. 2008 De Moor” (1 ex. AMGS). – Namibia: “Windhoek, New Dam 7.7. 34 JOC:” (3 exs. AMGS); “Windhoek Town Dam 7.7. 1939” (1 ex. AMGS); “SW Prot. Windhuk Parch 1919” (1 ex. SAMC); “E Capriwi, Katima Mulilo lux, 17.29S-24.17E, 3-8.3. 1992 Uhlig leg.” (3 exs. ZMHB, 1 ex. NMNW); “Kavango: Kaudom-Camp, Wasserloch, Schilf + Gras-Gesiebe, 18.31S-20.43E, 22-25.2. 1992 Uhlig leg.” (1 ex. ZMHB); “E Capriwi Mudumu NP: Nakatwa 18.10S-23.26E, 8-13.3. 1992 lux Uhlig leg.” (1 ex. ZMHB); “E Capriwi Mudumu NP: Buffalo Trails Camp, lux, ca. 18.10S-23.26E, 12.3. 1992 Uhlig leg.” (1 ex. ZMHB, 1 ex. MZH); “Kavango Mahango Game Reserve, piknik site, lux 24.11. 1993, 18.13S-21.45E Uhlig leg.” (1 ex. ZMHB); “Kavango: Popa Falls 18.07S-21.35E, 26.2.-3.3. 1992 Uhlig leg.” (2 exs. ZMHB, 1 ex. MZH; habitus in Fig. 411); “Okarupa ca. 17 mi E of Okahandja 4900 ft, 22.5. 1954/pools in overflow stream from dam, much weed and algae” (2 exs. BMNH); “Namib Mt. Naukluft river 24.16 S-16.15 E 10.8. 1989 shorewashing, river Endrödy & Klimaszew” (1 ex. TMSA); “Damaral. Okahandja 21.59S-16.52E / 12.9. 1974 shore washing Endrödy-Younga leg.” (1 ex. TMSA); “Kro, ca.15 mi. SE of Namutoni 30.5. 1954 / Shallow and muddy, with algae” (2 exs. BMNH, 1 ex. MZH); “Ca. 7 mi NE of Grootfontein 29.5. 1954 / Waterhole in dolomits” (2 exs. BMNH, 1 ex. MZH); “Ovamboland Namutoni 31.5. 1954 / Weedy water-hole and stream” (1 ex. TMSA). – Botswana: “Tsotsorogo Pan 17.6.-9.7. 1930 / L. taeniolatus Régimbart det. Gschwendtner” (14 exs. TMSA); same data but L. congener O-C. det. Wewalka 1981” (3 exs. OLML); “N´Kate Makarikari 6-23.8. 1930 / L. taeniolatus Régimbart det. Gschwendtner” (2 exs. TMSA); “Chobe NP, Savuti-Camp 18.33.55S-24.03.53E, 11.3. 1993 lux Uhlig leg.” (1 ex. ZMHB). – South Africa: “Trsvl., 5 mi W Warmbad 24-25.2. 1968 Spangler” (33 exs. USNM, 5 exs. MZH); “Tvl Nylstroom Donkerpoort dam 24.8. 1948” (1 ex. AMGS, possibly belongs to the type serie of L. congener but lacks paratype label); “Tvl Dam wall 28.11. JOC.” (1 ex. AMGS); “Tvl Potgietersrust 23.4. 1933 Taylor” (1 ex. AMGS); “OLF 101D 21.3. 55” (TVL Olifantsvlei) (1 ex. AMGS); “Transvaal Bronkhorstspruit Rinosterpoort b. Franz leg.” (1 ex. NMW); “C. Transvaal Roodeplaat Dam 28.37S-28.23E / 14.8.1974 shore washing leg. A. Strydom” (1 ex. TMSA, 1 ex. MZH); “Transvl Pretoria distr., Roodeplaat / UV light trap 30.10.-10.11. 1960 Dr. Neubecker” (1 ex. TMSA); “Roodeplaat Pretoria Distr. 10. 1960 Neubecker” (2 exs. TMSA); “Tv Nelshoogte Forest Station 25.50-30.50E / 2.12.1986 UV light collection Endrödy-Younga leg.” (1 ex.TMSA); “Tv Nelshoogte galery for. below St. 25.51S-30.53E / 4.12. 1987 UV light collection leg. Endrödy-Younga” (1 ex. TMSA); “Tvl Naboomspr. Torino Ranche24.37S-28.38E / 15.1. 1989 UV light, vlei edge Endrödy-Younga leg.” (1 ex. TMSA); “Tvl Rhenosterpoort N.R. 25,45S-28.55E / 15.11. 1975 at light Endrödy-Younga leg.” (1 ex. TMSA, 1 ex. MZH); “Nelspruit Pond 27.4. 2010, S25°32'13,83”, E30°59'50,35” Hidalgo-Galiano & Kleynhans leg.” (1 ex. CIR); “Transvaal April 1946” (1 ex. AMS); “Fountains 26.8. 05” (1 ex. TMSA); “Fountains 26.8. 1895(?)” (1 ex. TMSA); “Plat River 6-18.4. 05 / Waterberg Distr. Swierstra / L. taeniolatus Régimbart Gschwendtner det.” (7 exs. TMSA); “W Cape Arniston 7. 1946” (1 ex. BMNH); “ECPr. Matatiele 4-5.5. 1956 JOC.” (3 exs. AMGS); “ECPr. Matatiele 5.5. 1956 JOC.” (1 ex. AMGS);” “ECPr. Matatiele 5.5. 1956 JOC. / L. congener O-C. det. J. Omer-Cooper” (5 exs. AMGS); “ECPr. Mt Currie 6.5. 1956 JOC.” (2 exs. AMGS); “EC., Hwy 352, 3 km S Tsomo, in river 22.5. 2005 Challet” (1 ex. CGC); “ECPr., Qumbu 2.5. 1956 J.O-C.” (1 ex. AMGS); “Transkei Port St Jones Silaka 31.33S-29.30E/2.12. 1987 UV light collection leg. Endrödy-Younga” (1 ex. TMSA); Nat. -Drakensbg. Cathedral Peak 28.57S-29.12E / 15.3.1976 at light leg. Endrödy-Younga” (3 exs. TMSA); “Natal, TUG 77 Q38 (Tugela River system, Colenso 25.7. 1954” (1 ex. AMGS); “Natal Zululand Mtuba-tuba 23.9. 1947 JOC.” (1 ex, AMGS); “Natal 1942” (1 ex. AMGS); “Caffraria / Wahlb.” (1 ex. NHRS). – Swaziland: ”9 mi. from Mbabane 6.12. 1948 JOC” (1 ex. AMGS). – Lesotho: “Hensley’s Dam 8 mi. SW Leribe 30.3. 1951 / Brinck-Rudebeck” (6 exs. MZLU).

Comments on synonymy

The holotypes of L. taeniolatus and L. congener have been examined and compared with each other. No diagnostic differences can be discerned in shape of male genitalia. Externally both species are similar and accordingly they both belong to the same species, and L. congener is hence a junior subjective synonym of L. taeniolatus.

Diagnosis

Laccophilus taeniolatus is a widely distributed, but still reasonably uniform species. It resembles externally most of L. propinquus, L. simplicistriatus and L. complicatus. From the two latter species L. taeniolatus can in most cases be separated by study of the elytral colour pattern: dark irrorations are clearly reduced frontally on each side of the suture and an irregular, pale area is formed on base of each elytron. Laccophilus propinquus and L. taeniolatus requires dissection of male genitalia for correct identification; apex of penis exhibits a small difference in location of the small, frontal process. Additionally penis is distinctly shorter in L. propinquus.

Description

Body length 3.8–4.2 mm, width 1.9–2.3 mm. Colour pattern of dorsal aspect of body distinct; exhibits limited variation (Figs 410–412).

Head: Pale ferrugineous, posteriorly darker, ferrugineous to dark ferrugineous. Sometimes posteriorly with darker area; however vague and hardly discernible. Slightly mat; rather finely microsculptured; double reticulation indistinct; weakly developed medially on head. At eyes finely and irregularly punctate; between eyes with a few scattered, fine punctures.

Pronotum: Pale ferrugineous, frontally and posteriorly in middle with a dark brownish to dark ferrugineous marking. Posterior margin narrowly dark. Frontal and basal dark areas sometimes connected by a vague, ferrugineous area. Finely microsculptured, submat. Reticulation indistinctly double; large meshes (when distinguishable) contain 2–4 small meshes. At margins with very fine, sparse and irregularly distributed punctures.

Elytra: Pale ferrugineous, with dense, dark ferrugineous to blackish irrorations, which are somewhat unevenly distributed on elytra. Frontally, along dark suture, dark irrorations sparser, in part absent. Longitudinal, dark irrorations can in part (especially laterally) simply be “hollow” and consist only of dark outlines (Figs 410–412). Submat, finely microsculptured; reticulation indistinctly double. Kind of meshes, in general, difficult to classify according to size. Weakly developed rudiments of large meshes can be discerned. Each elytron with three (discal, dorsolateral and lateral), longitudinal areas with fine and sparse punctures.

Ventral aspect: Pale ferrugineous to ferrugineous. No distinct colour pattern. Rather shiny, very finely microsculptured; in part microsculpture indistinct. Abdomen basally very finely and sparsely striated. Metacoxal plates with fine, shallow, transversely located furrows. Apex of prosternal process slender, pointed. Apical sternite lacks asymmetric knob; provided with curved, sublateral impressions (of variable kind; one more pronounced) (Fig. 56).

Legs: Pro- and mesotarsus somewhat enlarged, rather long. Provided with suckers.

Male genitalia: Penis in lateral aspect quite long and evenly curved; minor frontal process protuding (Fig. 253).

Female: Apical sternite (Fig. 57).

Distribution

Gambia, Guinea-Bissau, Guinea, Burkina Faso, Chad, Central African Republic, Sudan, Sierra Leone, Liberia, the Ivory Coast, Togo, Benin, Nigeria, Cameroon, Gabon, Congo, Zaire, Kenya, Tanzania, Angola, Zambia, Malawi, Zimbabwe, Mozambique, Namibia, Botswana, South Africa, Swaziland, Lesotho (Fig. 539). Because of extensive taxonomic confusion we have only accepted personally verified records.

Collecting circumstances

A tolerant species, which occurs in a great variety of water bodies, both in stagnant and running waters. It is also collected from temporary pools. Laccophilus taeniolatus is a good flier and has often been captured at light collection. It also occurs in water bodies created or highly influenced by man, as in agricultural irrigation ditches. It dwells in forests as well as in open habitats as steppes and savannas (e.g. Guignot 1959a). Regarding collecting localities, see also Bilardo and Rocchi (2013).

Laccophilus propinquus Omer-Cooper, 1958

Figs 57–58, 254, 413, 540

Laccophilus propinquus Omer-Cooper 1958b: 37, 43, 45, 46 (original description, faunistics); Omer-Cooper 1965: 77, 82 (description, faunistics); Omer-Cooper 1970: 288, 289, 290 (discussion, description, faunistics); Pederzani 1988: 107 (faunistics, biology); Nilsson 2001: 249 (catalogue, faunistics); Nilsson 2015: 216 (catalogue, faunistics).

Type locality

Malawi: Mwanza.

Type material studied

(14 exs.). Holotype: male: “Type / male, female (symbols) Types / River near portuguese border, near Mwanza 9.II. (11 ?) 1948 / Brit. Mus. 1957-660 / L. propinquus O-C.”(BMNH; habitus in Fig. 413). – Paratype: female: Pinned together with holotype but on separate label (1 ex. BMNH); “Paratype / Nyasaland Zomba Plateau Reservoir 7.11. 1948” (1 ex. AMGS); “Paratype / Nyasaland Reservoir Mwanza 9.11. 1948” (2exs. AMGS); “Paratype / Nyasaland stream longer Lilongwe rd. 20 miles from Dedza 30.9. 1948” (1 ex. AMGS); “Paratype / Nyasaland dam on lower Lilongwe rd. 29.9. 1948” (2exs. AMGS); “Paratype / Nyasaland Cisaiti R. nr. Dedza 28.9. 1948” (1 ex. AMGS); “S. Rhodesia Inganyi River 17.IX. 1948 / L. propinquus sp. n. det. J. Omer-Cooper” (4 exs. AMGS); Nyasaland Dambo below Livingstonea Lake shore 21.9. 1945 / Paratype / L. propinquus O.-C., O. Cooper det.” (1 ex. IRSNB).

Additional material studied

(20 exx.). Tanzania: “Ruvu North Forest Reserve, waterholes, 3 km SE of Base 6°37'20” S, 38°55'00” E alt. 250 ft 1.11. 1992 / Hynd Collection” (4 exs. BMNH, 1 ex. MZH); “Ruvu North Forest Reserve, Base Camp6°37'40” S, 38°51'14” E alt. 200 ft 30.10. 1992 / Hynd Collection” (1 ex. MZH); “Zanzibar Pemba Sept. 1955 Fowler” (3 exs. AMGS); “Zanzibar Mangapwani Rd. Sept. 1955 JOC.” (5 exs. AMGS). – Malawi: “Dam, Dedza on lower Lilongwe rd. 30.9. 1948” (2exs. AMGS); “Mtiti R. 1.10. 1948” (3exs. AMGS).

Diagnosis

Laccophilus propinquus is very closely related to L. taeniolatus and externally very similar to this species. Small but distinct differences in shape of penis allow confident separation of the two species; penis short and apical process vague in L. propinquus (see also diagnosis of L. taeniolatus).

Description

Body length 3.8–3.9 mm, width 1.9–2.0 mm. Pale ferrugineous; dorsal, dark ferrugineous colour pattern of body quite distinct (Fig. 413).

Head: Pale ferrugineous, posteriorly with vague dark ferrugineous area. Submat, finely microsculptured. Reticulation almost of one kind, simple; double reticulation indistinct, weakly developed and difficult to distinguish. At eyes with irregularly distributed, sparse punctures.

Pronotum: Pale ferrugineous, anteriorly and at base with rather narrow, vague blackish ferrugineous to dark ferrugineous markings which are medially connected by a vague ferrugineous area. Submat, finely microsculptured; reticulation indistinctly double but size classes difficult to separate. Laterally and anteriorly with indistinct and sparse punctures.

Elytra: Pale ferrugineous, with rather distinct dark ferrugineous markings formed as irrorations (Fig. 413). Rather shiny, although finely microsculptured. Reticulation indistinctly double; clear size-classes difficult to discern. Discal, dorsolateral and lateral rows of fine and irregular punctures discernible but weakly developed.

Ventral aspect: Pale ferrugineous. Rather shiny and very finely (partly indistinctly) microsculptured. Abdomen with fine to very fine striae. Metacoxal plates with fine and shallow transverse furrows. Almost impunctate. Prosternal process slender, pointed. Apical ventrite almost symmetric, lacks lateral knob; apex more angle-shaped than in female (Fig. 57).

Legs: Pro- and mesotarsus long, slender, and with suckers.

Male genitalia: Penis In lateral aspect comparatively short, evenly curved and apical process not prominent (Fig. 254).

Female: Externally as male but apex of apical ventrite broader and more rounded (Fig. 58).

Distribution

Malawi, Zimbabwe, Tanzania and Zanzibar (Fig. 540). Omer-Cooper (1965) adds Mozambique.

Collecting circumstances

The species has been collected in streams e.g. with rocks, sand and some vegetation. It is also recorded from standing water in a swamp and a reservoir.

Laccophilus complicatus Sharp, 1882

Figs 59–60, 255, 414, 541

Laccophilus complicatus Sharp 1882: 308, 309 (original description, faunistics, discussion); Kolbe1883: 402 (description, faunistics); v. d. Branden 1885: 20 (catalogue, faunistics); Régimbart 1895: 138 (description, faunistics); Zimmermann 1919: 122 (faunistics); Zimmermann 1920a:17 (catalogue, faunistics); Bertrand 1928a: 184, 185 (juvenile faunistics); Bertrand 1928c: 365 (juvenile description); Bertrand 1948: 12: (juvenile faunistics); Bertrand 1954: 284 (juvenile discussion, faunistics); Guignot 1946c: 279, 281, 283 (discussion, description, faunistics); Guignot 1959a: 570, 573, 574 (description, faunistics); Bertrand 1963: 411 (discussion, faunistics); Omer-Cooper 1970: 290, 291 (description); Bertrand and Legros 1971: 245 (faunistics, biology); Bameul 1984: 94 (faunistics); Rocchi 1991: 80, 86 (faunistics); Nilsson 2001: 242 (catalogue, faunistics); Pederzani and Rocchi 2009: 95 (faunistics, list); Nilsson 2015: 210 (catalogue, faunistics).

Type locality

Madagascar.

Type material studied

(2 exs.). Laccophilus complicatus: Lectotype (by present designation) male: “Type / Madagascar / Sharp Coll. 1905-313 / Madag. / Laccophilus rivulosus Klug /Type 574 L. complicatus sp. n. Madagascar” (BMNH). – Paralectotype: female: ”574 / Cotype / Madagascar / Sharp Coll. 1905-313 / Madagascar / Laccophilus complicatus Shp Co-type” (1 ex. BMNH).

Additional material studied

(390 exs.). Madagascar: “E-Mad., Morarano, N. Morananga 900 m asl, 13.1. 1995 Janák” (9 exs. NMW; habitus in Fig. 414); “Moramanga env. 10-18.12. 1997 Pacholátko” (1 ex. NHMB); “E-Mad. Ampamoho nr Andilamena 1200-1300 m asl 18-20.1. 1995 Dunay & Janák” (20 exs. NMW); “Ambatombe nr Andilamena 900 m asl, 17.1. 1995 Dunay & Janák” (10 exs. NMW, 10 exs. MZH, 1 ex. NHRS); “E-Mad. Andranokobaka, N Moramanga 800 m asl 13.1. 1995 Dunay & Janák” (21 exs. NMW); “W-Mad. Manindaray W Sakahara 700-800 m asl 30.1. 1995 Dunay & Janák” (1 ex. NMW); ”5 km S Ampamoho 950-1000 m asl 18-20.1. 1995 Dunay & Janák” (3 exs. NMW); “E Sakahara 30.1. 1995 Manindray 700-800 m Dunay & Janák” (1 ex. NMW); “Mad. Centr. Antananarive 18-19.1. 1993 Janák” (1 ex. NMW); “Envir. de Tananarive 7. 1970 Pederzani” (1 ex. AMGS, 2 exs. NHMB); “Prov. Tananarive, env. de Arivonimamo 22.7. 1970 Pederzani” (1 ex. AMGS); “Tan. Manjakatompo 3.1. 1958 Keiser” (1 ex. NHMB); “Mandrare Bas., Loc. prés Andaza, affl. non nommé, Riv., 46°34'05"E, 24°03'16"S, Alt. 315 m 26.4. 1995 Elouard & Pilaka” (1 ex. NMW); “Prov. Tamatava, 3.3 km N Ambabasoratra 31.8. 1962 Cashatt” (1 ex. USNM); “Mad-est 1100-1200 m, NP Ranomafana / Vohiparara 21-24.1. 1993 Janák” (2 exs. NMW); “Prov. Fianarantsoa 7 km W Ranomafana, 1100 m 8-21-10. 1988 Steiner W.E. / At black light in montane rainforest” (3 exs. USNM, 1 ex. MZH); “Prov. Fianarantsoa 7 km W Ranomafana, 1000 m 23-28.2. 1990 Steiner W.E.” (1 ex. USNM); “Ankaratra (Antananarivo) Res. Manjakatompo 6.10. 2001 / Gerecke et Goldsmith leg. / Helocrene at left border of affluent to Lac Froid, 1700 m asl” (2 exs. BMNH); “Manjakatompo 5.10. 1989 Bartolozzi & Taiti leg. / m 1700 Stazione pescicoltura / L. complicatus Sharp det. Rocchi 1989”(3 exs. CSR); “Antranovy (Antananarive) 13 km W from Arivonimana, 13.7. 2001 helocrene in rice field exp. N / 1480 m asl / Gerecke et Goldsmith leg.” (9 exs. BMNH, 2 exs. MZH); “Tsimelahy (Tuelar) 5.9. 2001, Riv. Antarantsa downstr. piceine naturelle / 200 m asl / Gerecke et Goldsmith leg.” (1 ex. BMNH); “Foret de l’Est Perinet-Anosibe 11-12. 1959” (1 ex. BMNH); “Tan. Madag, Ampefy, Lac Kavitaha wi 25.III. 58 Keiser” (1 ex. NHMB); “Tam. Perinet 3.12. 1957 Keiser” (1 ex. NHMB); “La Mandraka, ex. coll. Breuning” (1 ex. MRAC); “Suberbieville, ex. coll. Breuning” (5 exs. MRAC); “Tananarive 22-29.i. 1972 Hecq” (1 ex. MRAC); “Tananarive, at light 12.12. 1955 E. Mac (?) Callan” (1 ex. AMGS); “Tananarivo Friedrichs / L. complicatus Sharp det. Brancucci” (1 ex. ZMHB); “Fianarantsoa Prov., Foret d’Antsirakanbiaty, 7.6 km 285˚ WNW Itremo, elev. 1550 m 22-26.1.2003 / 20°35'36"S 046°33'48"E, collected at light in montane rainforest, Fischer, Griswold & al. leg.” (2 exs. CAS); “Fianarantsoa Ranomafana National Park, Talatakely res. lab. area, black light mercury vapour / light, elev. 940 m 21°14'53,5"S, 47°25'35,9"E, 31.10-20.11.1998 Lee & Ribardo leg.” (1 ex. CAS); “Fian Isalo Menamaty R., degraded river with lots of vegetation used by women to wash clothes in, 11.5. 2006, N-22°33.001, E45°24.074, 757 m Bergsten et al.” (12 exs. NHRS, 5 exs. MZH); “Fian Andringitra Zomandao R. bridge on road to the camp belamba, vegetation rich edges along the river N-22°6.225, E46°55.244, 8-9.5.2006, 1421 m Bergsten et al.” (13 exs. NHRS); “Fian. Col. des Tapias Rte Tana-Fianarantsoa, vegetation- rich pond with fish 6.5. 2006 N-20°46.376, E47°10.749, 1718 m Bergsten et al.” (9 exs. NHRS); “Mangoky Bas, Loc. Andringitra, Camp B, Zomandao River 46°53'46"E/22°07'12"S 1600 m asl 30.11. 1993 leg. ORSTOM” (1 ex. NHRS); “Andringitra Rambavy R. (Cascade) N-22.153, E 46.9, 1979 m, 8.5. 2006 Bergsten et al. / BMNH(E) <794244> DNA Voucher” (1 ex. NHRS); “Ft Dauphin, rice paddies N-24.824, E 46.866, 34.44 m 19.5. 2006 Bergsten at al/BMNH(E)(E) <794247> DNA voucher” (1 ex. NHRS); “TOLI Zombitse, N-22 64 Andramomena R. pool, N-22.64, E44.864: 577 m, 15.5.2006 leg. Bergsten” (1 ex. NHRS); “Mangoro Bas Loc. Ankirihitra Tsarantanana River 47°17'37"E/19°23'00"S, 11.1. 1997 Elouard leg.” (1 ex. NHRS); “Mad-est. 1100-1200 m P.N. Ranomafana / Vohipara 21-24.1. 1993 Dunay & Janák leg.” (1 ex. NHRS); “Mah/Tol. Melaky/Menabe Ambojihanahary NP, S18.26849. E045.46346, 906 m.a.o., 19.12. 2009 water net, field Bergsten et al. / NHRS-JLKB 000000723” (1 ex. NHRS); “Madagascar” (1 ex. USNM, 2 exs. ZMHB).

Specimen with uncertain locality

(2 exs.). “Dirjo 24” (1 ex. NMW); “Mexique” (1 ex. MHNG).

Diagnosis

Laccophilus complicatus, distributed solely on Madagascar, resembles most of three species on mainland of Africa, viz. L. simplicistriatus, L. taeniolatus and L. propinquus. In general L. complicatus is somewhat larger than the three other species. Elytral colour pattern in L. complicatus is uniform and more evenly distributed than in L. taeniolatus and in L. propinquus, in which a vague and irregular, dark, longitudinal marking can be discerned discally on each elytron. Elytral colour pattern in L. simplicistriatus is fairly uniform and exhibits only slight variation. In L. complicatus penis in lateral view is not expanded posterior to apex while in L. simplicistriatus corresponding feature is clearly expanded on each side. From L. taeniolatus the species is distinguished by the tip of penis, which is clearly curved ”upwards” while tip of L. complicatus penis is almost straight. Regarding L. propinquus, L. complicatus is separated from it as L. taeniolatus is but tip of penis vague.

Description

Body length 3.9–4.6 mm, width 2.2–2.6 mm. Pale ferrugineous, with blackish ferrugineous to dark ferrugineous, dense irrorations on elytra (Fig. 414). Dorsal colour pattern of body quite stable; rarely elytral irrorations laterally and at base slightly reduced.

Head: Pale ferrugineous, posteriorly, narrowly dark ferrugineous. Submat, reticulation in part double; large meshes often weakly developed and hardly visible. At eyes, with fine and irregular punctures, extending medially towards midhead.

Pronotum: Pale ferrugineous, anteriorly and posteriorly in middle with vague, darkened areas. Reticulation quite dense, double. Large meshes weakly developed; one mesh contains between three and six fine meshes. At margins with fine, irregular punctures, which are rather indistinct at posterior margin.

Elytra: Pale ferrugineous, with dense blackish ferrugineous to dark ferrugineous irrorations (Fig. 414). Submat, with quite dense reticulation. Reticulation indistinctly double; indistinct fragments of larger meshes extensively discernible. Each elytron provided with three, longitudinal, rather vague areas of fine and sparse punctures.

Ventral aspect: Pale ferrugineous to ferrugineous, abdomen in posterior part blackish to dark ferrugineus. Rather shiny to submat, very finely reticulated. Almost impunctate; fine punctures discernible on apical ventrite. Apical ventrite as in Fig. 59. Basal ventrites provided with fine, slightly curved, striae. Metacoxal plate with some very shallow, almost transversely located furrows. Prosternal process, slender, extended and pointed.

Legs: Pro- and mesotarsus somewhat enlarged; provided with suckers.

Male genitalia: Penis in lateral aspect evenly curved, broad; extreme apex curved upwards and sharp (Fig. 255).

Female: Pro- and mesotarsus slender. Apical ventrite as in Fig. 60. Rarely microsculpture of body is strongly developed and such a specimen is clearly matter than ordinary female specimens.

Distribution

Madagascar (Fig. 541).

Collecting circumstances

In literature, detailed documentation is not available. From collecting labels appear that L. complicatus has been sampled in areas between 700-1700 m a.s.l. At least once collected in a degraded river with lots of vegetation, used to wash clothes in. Obviously it is attracted by light and sampled in a montane rain forest.

Laccophilus irroratus Aubé, 1838

Figs 61–62, 256, 415, 535

Laccophilus irroratus Aubé 1838: 427 (original description, faunistics); Sharp 1882: 309 (description, faunistics); Kolbe 1883: 426 (description, faunistics); Régimbart 1895: 138, 141 (description, faunistics, discussion); Alluaud 1897: 212 (faunistics); Peschet 1917: 23, 24, 55 (description, faunistics); Zimmermann 1920a:20 (catalogue, faunistics); Guignot 1946c: 268 (description, faunistics); Vinson 1956: 29 (faunistics, list, biology); Guignot 1957a: 98 (faunistics); Guignot 1959a: 557, 560, 562 (description, faunistics); Guignot 1961a: 930 (faunistics); Vinson 1967: 314 (faunistics, list); Wewalka 1980: 729, 730 (faunistics); Bameul 1984: 93, 102 (faunistics); Nilsson 2001: 245 (catalogue, faunistics); Pederzani and Reintjes 2002: 40 (faunistics); Nilsson 2015: 213 (catalogue, faunistics).

Type locality

Réunion: Ile de France et Bourbon.

Type material studied

(1 ex.): Holotype, male: “Data in NHRS JLKB 000030278 / Laccophilus irroratus mihi h. in ile de France D. Latereille / Ex Musaeo Dejean / D. Sharp Monogr. / irroratus / Dr. Régimbart 1893 / Coll. Oberthur” (MNHN).

Additional material studied

(21 exs.). Mauritius: “I. Maurice Montrésor Ch. Alluaud 1893 / male symbol / Museum Paris coll. Maurice Régimbart” (1 ex. MNHN); “I. Maurice Mon Désert P. Carie Fév. 1903 / Laccophilus irroratus Aubé / Museum Paris coll. 1945 R. Peschet” (1 ex. MNHN); “Ile Maurice / de Borre” (3 exs. MHNG); “Ins. Mauritius Westw.” (1 ex. ZMHB). – Reunion: “La Reunion Palmistes Ch. Alluaud 1893 / Museum Paris coll. Maurice Régimbart 1908 / irroratus Aubé” (2 exs. MNHN; habitus in Fig. 415); “S St. Benoit NW Cambourg, ca. 250 m 2-3.1. 1999 Wewalka / L. irroratus Aubé Wewalka det. 99” (2 exs. CGW, 1 ex. MZH); “E St. Joseph Riv. Langevin 250 m 28.12. 1998 leg. Wewalka / L. irroratus Aubé Wewalka det. 99” (4 exs. CGW, 4 exs. MZH); “Plaine des Cafres 3.3. 1935 Vinson” (1 ex. MNHN); “Ins. I. Fr. Dufr.” (1 ex. ZMHB).

Diagnosis

Laccophilus irroratus is characterized by large body size, peculiar elytral colour pattern and penis, which is different from all other African Laccophilus species; penis in lateral aspect quite delicate, distinctly curved and simple, exhibiting minor modifications.

Description

Body length 4.0–4.3 mm, breadth 2.4–2.6 mm. Dorsal, colour pattern of body stable, exhibits only minor variation (Fig. 415).

Head: Pale ferruginous, posteriorly close to pronotum with narrow but distinct, dark ferrugineous area. Rather shiny, although distinctly microsculptured. Reticulation double, large meshes contain 3–5 small meshes. In middle of head small meshes in part reduced. At eyes with scattered, fine punctures; area with punctures extends towards middle of head.

Pronotum: Pale ferrugineous to ferrugineous. Anteriorly, at level of eyes with a broad, black to dark ferrugineous area. At base with a quite narrow, black to dark ferrugineous area, which is medially somewhat enlarged. Delimitation of darker areas is somewhat vague. Rather shiny, although distinctly microsculptured; reticulation double. Large meshes distinct while small meshes, especially medially, are fine, in part hardly visible or totally absent. Punctures absent, except in frontal part and laterally where puncture very fine and scattered.

Elytra: Pale ferrugineous, with distinct, only slightly variable, dark ferrugineous irrorations (Fig. 415). One specimen with quite broad, transverse, pale area frontally. Rather shiny, although distinctly microsculptured; reticulation double. Large meshes contain, when discernible, 3–5 fine meshes. Fine meshes sometimes obliterated. Fine scattered punctures form a discal row of punctures. Scattered, irregular punctures indicate presence of dorsolateral and lateral rows of punctures. Laterally, with a pre-apical furrow.

Ventral aspect: Blackish ferrugineous to dark ferrugineous, no distinct colour pattern but three basal ventrites are somewhat paler than apical ones. Rather shiny, in part with very fine microsculpture. Almost impunctate, except apical ventrite, with scattered fine punctures, and frontally on metathorax, with fine, fairly dense punctures. Abdominal ventrites with fine, curved striae. Metacoxal plates with about 10 shallow and in part slightly indistinct furrows. Metacoxal process not modified. Apical ventrite, symmetric (no knob discernible) (Fig. 61). Prosternal process rather narrow, apex only slightly extended, pointed.

Legs: Pro- and mesotarsus slender, somewhat extended. Segments provided with suckers.

Male genitalia: Penis delicate, in lateral aspect distinctly curved; apex simple and exhibits hardly any modifications (Fig. 256).

Female: Pro- and mesotarsus slender. Apical ventrite (Fig. 62).

Distribution

Mascarene Islands; Réunion, Mauritius (Fig. 535). Guignot (1957a) gives also Rodriguez. Laccophilus irroratus is an endemic species of Mascarene Islands (Guignot 1961a).

Collecting circumstances

Almost unknown. Reported in rock pools of slow stream (Vinson 1956).

Laccophilus rivulosus Klug, 1833

Figs 63–64, 257, 416, 542

Laccophilus rivulosus Klug 1833: 48 (original description, faunistics); Aubé 1838: 4 (description, faunistics); Sharp 1882: 287, 821 (description, faunistics); Kolbe 1883: 401 (description, faunistics); v. d. Branden 1885: 24 (catalogue, faunistics); Régimbart 1895: 136, 140 (discussion, description, faunistics); Régimbart 1903: 14 (faunistics); Zimmermann 1919: 122 (faunistics); Zimmermann 1920a: 25 (catalogue, faunistics); Guignot 1937: 141 (discussion); Guignot 1959a: 544 (description, faunistics); Bertrand and Legros 1971: 245 (faunistics, biology); Rocchi 1991: 86: (faunistics, list); Nilsson 2001: 250 (catalogue, faunistics); Nilsson 2015: 217 (catalogue, faunistics).

Type locality

Madagascar.

Type material studied

(4 exs.): Lectotype (by present designation): male: “9982 / Typus / L. rivulosus Kl. Madag, Goudot / Hist-Coll. (Coleoptera) Nr. 9982 Laccophilus rivulosus Kl. Madagascar Goudot Zool. Mus. Berlin” (ZMHB). – Paralectotypes: Madagascar Goud. Nr. 9982 / Typus / Hist-Coll. (Coleoptera) Nr. 9982 Laccophilus rivulosus Kl. Madagascar Goudot Zool. Mus. Berlin” (3 exs. ZMHB).

Additional material, studied

(38 exs.). Madagascar: “Antakotako II 1936” (1 ex. MNHN; habitus in Fig. 416); “Mad. Sud, Ft Dauphin Alluaud 1900” (3 exs. MNHN); “Mad. Sud, Ft Dauphin Alluaud” (2 exs. MNHN); “Tamatave Perrot” (1 ex. Z MHB, 2 exs. MNHN); “Antsianaka Perrot Freres, 1er Semestre 1892” (1 ex. MNHN, 1 ex. NHMB, 6 exs. SAMC); “Antsianaka / L. rivulosus Kl. det. M. Brancucci” (1 ex. MNB); “Suberbieville” (1 ex. MNHN); “Mt. d’Ambre / Mai” (1 ex. ZMHB); “St. Marie Moaroay / Mad. Kaudern / L. rivulosus Kl. det. Zimmermann” (4 exs. NHRS); “Mahajanga Melaky, btw Bekopaka-Antsalova, S18.91556, E044.55546, 47 m.a.o. 16.12. 2009 water net, field Bergsten et al.” (2 exs. NHRS); same data but add “NHRS-JLKB 000000726” (1 ex. NHRS); “Mahajanga Melaky, btw Antsalova-Maintirano S18.30233, E044.18071, 37 m.a.o., 18.12. 2009, water net, field, Bergsten et al. /NHRS-JLBK 000000728” (1 ex. NHRS); “Mahajanga Boeny, Ankarafantsika NP, S16.30341, E046.81073, 74 m.a.o., 29.11. 2009, 22 W black light, field, Bergsten et al.” (1 ex. NHRS); same data but “S16.31215, E046.81523, 76 m.a.o., water net, field” (2 exs. NHRS); same data but “S16.31215, E046.81523, 76 m.a.o., water net, field/NHRS-JLKB 000000724” (1 ex. NHRS); “Mahajanga Boeny, Mahavavy Kinkony RS, S16.06651, E045.77672, 24 m.a.o., 5.12. 2009, water net, field, Bergsten et al./NHRS-JLKB 000000727” (1 ex. NHRS); “Toliara Menabe, Kirindy R. S., S20.07641, E044.674708, 65 m.a.o., 11.12. 2009 water net, field, leg. Bergsten et al” (2 exs. NHRS); “FIAN, Isalo, Piscine Noir, hygropetric 12.5. 2006 Bergsten et al. /BMNH(E) <794159> DNA voucher / L. rivulosus Kl. det. Bergsten” (1 ex. NHRS); “Madagascar Fairmaire / L. rivulosus Kl. Madg.” (1 ex. NMW); “Madag. Perrier” (2 exs. SAMC).

Diagnosis

Laccophilus rivulosus is characterized by large body, peculiar elytral colour pattern and shape of penis. Dark longitudinal markings of elytra are quite broad and distinct. Pale irrorations can generally be discerned within dark, longitudinal marking. Penis resembles much of penis of L. posticus but it is somewhat larger and extreme apex more extended. Additionally large body size and clear differences in elytral colour pattern easily separates L. rivulosus from L. posticus.

Description

Body length 4.8-5.3 mm, width 2.7-3.0 mm. Dorsal colour pattern of body generally distinct and uniform (Fig. 416); rarely slightly variable.

Head: Pale ferrugineous, posteriorly between eyes with a distinctly delimited blackish ferrugineous to dark ferrugineous area. Rarely dark area reduced and only in part visible because hidden beneath frontal part of pronotum. Almost impunctate. At eyes in shallow depression with fine, irregularly distributed punctures. Submat, rather distinctly microsculptured. Reticulation double. Coarser meshes only slightly stronger in comparison with fine meshes. In part, kinds of meshes difficult to separate. When discernible coarse meshes contain 3-5 fine meshes.

Pronotum: Pale ferrugineous, frontally with a distinct, dark ferrugineous area. Posteriorly in middle with an, often, bilobed, dark ferruginoeus, narrow spot. Almost impunctate. No distinct punctation discernible. Submat, finely microsculptured. Reticulation double. Coarse meshes well developed, contain 2-5 fine meshes.

Elytra: Pale ferrugineous, with distinct, longitudinal, dark ferrugineous markings. Dark markings somewhat undulate and merged into pairs, forming pale, inner, irrorations (Fig. 416). Rarely undulation in anterior half of elytra indistinct because of expanded dark areas. Almost impunctate; an indistinct, discal row of punctures discernible. Rather shiny, although distinctly microsculptured. Reticulation double. Coarse meshes contain 2-5 fine meshes. Narrowly at suture elytra slightly elevated.

Ventral aspect: Pale ferrugineous. Almost impuntate; abdomen with a few scattered punctures. Rather shiny, although extensively, finely microsculptured. In part microsculpture almost obliterated or totally obliterated. Abdomen laterally with fine, curved striae. Apical ventrite as in Fig. 63. Prosternal process rather slender, apex pointed but not strongly extended posteriorly. Metacoxal plates with about 10 furrows, of which 2–3 closest to metasternal wing are distinct while others rather indistinct to indistinct.

Legs: Pro- and mesotarsus slightly enlarged, somewhat extended and provided with some protruding suckers.

Male genitalia: Penis quite robust, in lateral aspect distinctly curved and apex extended and sharp (Fig. 257).

Female: Apical ventrite as in Fig. 64. Pro- and mesotarsus rather slender, extended.

Distribution

Endemic for Madagascar (Fig. 542).

Collecting circumstances

Information on biology is almost totally lacking. Bertrand and Legros (1971) reports the species from a small swamp and from pools of a temporary river.

Species group 8 (L. immundus group)

Diagnosis. Quite large species with body length 4.3–4.5 mm, width 2.3–2.5 mm.

Body shape oval-oblong, dorsoventrally flattened (Fig. 417). Body colour ferrugineous to dark ferrugineous. Lacks distinct colour pattern (Fig. 417). Body microsculpture double, distinctly of two kinds.

Prosternal process slender, slightly extended, apically pointed. Apical ventrite modified; posteriorly on each side somewhat excavated; medially, posteriorly extended, lacks asymmetrical knob on one side of ventrite (Fig. 65). No stridulatory apparatus on metacoxal plates. Metacoxal process, posteriorly not extended (Fig. 6).

Paramere apically narrow, basally enlarged; moderately modified (Fig. 258). Apical part of penis straight to almost straight; apex resembles a harpoon (Fig. 258).

Species composition and distribution. One species is recognized and distributed in South Africa.

Laccophilus immundus Sharp, 1882

Figs 65–66, 258, 417, 543

Laccophilus immundus Sharp 1882: 304 (original description, faunistics, discussion); v. d. Branden 1885: 21 (catalogue, faunistics); Régimbart 1895: 131 (description, faunistics); Zimmermann 1920a: 20 (catalogue, faunistics); Guignot 1959a: 579 (description, faunistics); Omer-Cooper 1962: 294, 296 (faunistics, discussion); Omer-Cooper 1965: 76, 81 (description, faunistics); Nilsson 2001: 244 (catalogue, faunistics); Nilsson 2015: 212 (catalogue, faunistics).

Laccophilus spadix Omer-Cooper 1953: 23 (original description, faunistics, biology); Omer-Cooper 1965: 81 (list, synonymy); Nilsson 2001: 244 (catalogue, list, synonymy); Nilsson 2015: 212 (catalogue, faunistics, list, synonymy). Confirmed synonym.

Type localities

Laccophilus immundus: South Africa: Cape Town.

Laccophilus spadix: South Africa: Cape Province, Caledon district.

Type material studied

(4 exs.). Laccophilus immundus: Holotype: female: “Type / S. Africa / Sharp Coll. 1905-313 / Type 595 Laccophilus immundus sp. n. Capetown” (BMNH). – Laccophilus spadix: Holotype: male: “Type / W Cape P., stream with pools in pine wood, Caleda dist. 18.XI. 1947 J. Omer-Cooper / Brit. Mus. 1957-660 / L. spadix O-C. det. J. Omer-Cooper” (BMNH). – Paratypes: female: “Type / Cape Peninsula, small vlei nr Cape Town 18.XI. 1947 J. Omer-Cooper / Brit. Mus. 1957-660 / L. spadix O-C. det. J. Omer-Cooper female allotype” (1 ex. BMNH); “W. Cape Pr., stream with pools in pine wood Caledon distr. 18.XI. 1947 JOC. / L. spadix O-C.” (1 ex. AMGS).

Additional material studied

(945 exs.). South Africa: “Western C. Pr. stream with pools in pine wood Caledeon Dist. 18.XI. 1947 Omer-Cooper” (1 ex. AMGS); “Pools in Caledon 11. 1947” (14 exs. AMGS); “WC. Prov. Princess vlei Cape Town 13.IV. 1947”(1 ex. AMGS); “Cap. b. sp. De Vylder / L. immundus Shp det. A. Zimmermann” (5 exs. NHRS); “Cape Prov., Cape Flats, Varden Vlei 2 mi E Ottery, 2.2. 1951 / L. spadix = L. immundus det. J. Omer-Cooper” (3 exs. MZLU; habitus in Fig. 417); “Cape prov., Silversand, 8 mi W Kleinmond 19.12. 1950” (1 ex. MZLU); “Cape Good Hope Nature Reserve 7-10.3. 1968 Spangler” (422 exs. USNM, 20 exs. MZH); “CPr., Cape of Good Hope 8.1. 1994 Wewalka” (2 exs. CGC); “WC, Cape point Res., Cape peninsula 28.8. 2007, dam, Pryke leg., 34.30947S, 18.44977E” (2 exs. CCT); “WC, Cape of Good Hope 20.3. 2001, pond 2 km N field museum, Ribera et Cleslak” (1 ex. CIR); “WC, Cape of Good Hope Reserve, pond, 26.2. 1997 Turner” (60 exs. CCT, 6 exs. MZH); same data but “16.2. 1997” (100 exs. CCT); “WC, Cape Point Reserve, S341436, E182306, 30.8. 2003, seasonal pool with fibrous vegetation, Turner, Mann & Reavell” (46 exs. CCT, 6 exs. MZH); “WC, Cape Point Res., seasonal pool, 3418S, 182631E, alt. 87 m, Turner, Mann & Reavell” (25 exs. CCT); “WC, Cape Town, Cape of Good Hope Res., pond 26.2. 1997 Turner” (166 exs. CCT); same data but “Reservoir on roadside to Bordjlesri 15.2. 1997 Turner” (55 exs. CCT); “WC, Kromrivier at roadsite, Cape G.H. Res. 15.2. 1997 Turner” (1 ex. CCT); “WC, Schusters Kraal, stream, netted between footbridge and dam, 341223S, 182237E, 13.9. 2003, Turner, Mann & Reavell” (8 exs. CCT).

Comments on synonymy

Holotypes of both involved taxa have been examined and compared. No diagnostically important differences detected and accordingly, earlier introduced synonymy is confirmed. Laccophilus immundus, being older than L. spadix, is the valid name of the species.

Diagnosis

A peculiar species, which is especially characterized by shape of penis apex (harpoon-like) and by body, being almost one-coloured, piceous to dark brownish or dark ferrugineous.

Description

Body length 4.3–4.5 mm, width 2.3–2.5 mm. Dark ferrugineous to ferrugineous, dorsal colour pattern of body is vague, reduced and rather indistinct (Fig. 417).

Head: Pale ferrugineous to ferrugineous, posteriorly and at eyes dark ferrugineous; delimitation of colour pattern vague. Rather finely to finely and densely punctate. Close to eyes with a few coarser punctures. Submat, distinctly microsculptured. Meshes double; larger meshes include 2-5 fine meshes.

Pronotum: Ferrugineous, basally in middle with a dark ferrugineous marking. Punctures fine to very fine, rather dense and slightly irregularly distributed. At margins with a partly irregular row of punctures. (In part, row frontally replaced by rather narrow area of fine punctures.) Microsculpture distinct, dense and double: coarser meshes include 2-6 finer meshes.

Elytra: Dark ferrugineous, vague darker markings sometimes discernible but form no distinct colour pattern (Fig. 417). Finely and densely punctate. Very sparse (indistinct) longitudinal rows of slightly coarser punctures discernible. Laterally in posterior half along the edge of elytron punctures form a narrow furrow. Submat, microsculpture double; coarse meshes include 2-6 fine meshes.

Ventral aspect: Pale ferrugineous to ferrugineous. Almost impunctate. In part, very finely microsculptured. Metacoxal plates, especially in frontal part with rather distinct furrows. Apical ventrite lacks “one-side” asymmetric knob (Fig. 65). Ventrites with curved striae. Prosternal process slender and pointed.

Legs: Pro-and mesotarsus slightly enlarged, provided with suckers.

Male genitalia: Penis quite long and slender, slightly sinuate and apex “harpoon-like” (Fig. 258).

Female: As male but apical ventrite slightly different in shape (Fig. 66). Pro- and mesotarsus slender.

Distribution

South Africa (Fig. 543).

Collecting circumstances

Poorly documented. Label data gives various water bodies as pond, stream with pools in pinewood as a collecting sites, but available information is generally quite superficial.

Species group 9 (L. pellucidus group)

Diagnosis. Large species; body length 5.3–6.0 mm, width 3.0–3.4 mm.

Shape of body oval, dorsoventrally distinctly flattened (Fig. 418). Dorsal colour pattern diffuse, sometimes almost absent. Rarely epipleura of female distinctly expanded posterior to middle (Fig. 8). Often elytra exhibit rather dense, dark ferrugineous to ferrugineous irrorations, which laterally in part become indistinct. A few pale spots generally present at base and apicolaterally on elytra (Fig. 418). Dorsal microsculpture double, of two different kinds; in part reticulation is obscured and indistinct.

Prosternal process quite broad, rather short, apex pointed. Apical ventrite somewhat modified; posterior end of ventrite excavated on both sides and medially ventrite somewhat extended. Asymmetric knob on one side of ventrite lacking (Fig. 67). Stridulatory apparatus absent. Metacoxal process not extended posteriorly (Fig. 6).

Parameres moderately modified. Penis is exceptionally large with apex strongly modified; clearly different from all other African species (Fig. 259).

Species composition and distribution. One species recognized in this species group, distributed in Africa South of Sahara excluding Madagascar.

Laccophilus pellucidus Sharp, 1882

(Figs 8, 67–68, 259–260, 418, 560)

Laccophilus pellucidus Sharp 1882: 304 (original description, faunistics); v. d. Branden 1885: 23 (catalogue, faunistics); Régimbart 1895: 131 (description, faunistics); Zimmermann 1920a: 24 (catalogue, faunistics); Guignot 1959a: 579, 584 (description, faunistics); Omer-Cooper 1962: 295 (faunistics); Omer-Cooper 1965: 76, 87 (description, discussion, faunistics, synonymy L. ampliatus Régimbart and L. pellucidus); Omer-Cooper 1967: 60 (discussion); Biström 1979: 22 (faunistics); Pederzani 1988: 107 (faunistics, biology); Nilsson and Persson 1993: 82, 94 (faunistics, biology); Rocchi 2000: 25 (discussion, description); Nilsson 2001: 248 (catalogue, faunistics); Nilsson 2015: 215 (catalogue, faunistics).

Laccophilus ampliatus Régimbart 1895: 130 (original description, faunistics); Régimbart 1906: 248 (faunistics); Zimmermann 1920a:16 (catalogue, faunistics); Omer-Cooper 1958b: 37, 48, 49, 51 (discussion, description, faunistics, biology, synonymy L. ampliatus and L. pilitarsis Régimbart); Omer-Cooper 1965: 87 (description, faunistics, synonymy L. ampliatus and L. pellucidus Sharp); Omer-Cooper 1967: 60 (discussion, list, synonymy); Nilsson and Persson 1993: 82 (list, synonymy); Nilsson 2001: 24 (catalogue, faunistics, list, synonymy); Nilsson 2015: 215 (catalogue, faunistics, list, synonymy). Confirmed synonym.

Laccophilus pilitarsis Régimbart 1906: 247 (original description, faunistics); Régimbart 1908: 5 (faunistics); Zimmermann 1920a: 24 (catalogue, faunistics); Peschet 1921: 5, pl. 1 fig. 5 (discussion, description, faunistics); Omer-Cooper 1931: 756 (description, biology, faunistics); Guignot 1946c: 282, 283, 284, 312 (description, faunistics); Guignot 1954: 29 (faunistics); Omer-Cooper 1957: 21, 90 (faunistics); Omer-Cooper 1958a: 59 (faunistics); Omer-Cooper 1958b: 37 (synonymy L. ampliatus Régimbart); Guignot 1959a: 576, 581 (description, faunistics); Guignot 1959b: 355 (faunistics); Guignot 1959d: 162 (faunistics); Bertrand and Legros 1975: 672 (discussion); Nilsson and Persson 1993: 82 (list, synonymy L. pellucidus Sharp); Nilsson 2001: 248 (catalogue, faunistics, list, synonymy); Nilsson 2015: 215 (catalogue, faunistics, list, synonymy). Confirmed synonym.

Type localities

Laccophilus pellucidus: South Africa: Natal, Bedford District.

Laccophilus ampliatus: South Africa: Natal.

Laccophilus pilitarsis: Kenya: Taveta.

Type material studied

(6 exs.). Laccophilus pellucidus: Holotype (type unique): male: Laccophilus pellucidus Type D.S. Bedford District, Caffraria 175 Laccophilus / Type H.T. / Sharp Coll. 1905-313 (BMNH).

Laccophilus ampliatus: Lectotype (by present designation): female: “Natal / Museum Paris Coll. Maurice Régimbart 1908 / Type / ampliatus Rég. type” (MNHN). – Paralectotype: female: “98 / Laccophilus ampliatus Rég. type unique / SAM Type acc. no. 838” (1 ex. SAMC). [Comment: note that Régimbart, when describing L. ampliatus, indicated presence of both sexes in type material – both are, however, females.]

Laccophilus pilitarsis: Lectotype (by present designation): male: “Afrique Orle Anglaise Taveta Ch. Alluaud I-IV. 1904 / Museum Paris coll. Alluaud / Type” (MNHN). – Paralectotypes: “Afrique Orle Anglaise Nairobi (Wa Kikuyu et Masai) Ch. Alluaud 2. sem. 1903 / Museum Paris coll. Alluaud / Type” (2 exs. MNHN).

Additional material studied

(413 exs.). Ethiopia: “Abyssinia Hora Bishoftu 7000 ft. 23.XII. 1926 JOC.” (3 exs. AMGS); “Hora Horeso 7000 ft. 1.12. 1926 JOC.” (12 exs. AMGS); “Small pond Hora Abjata 5000 ft. 18.XI. 1927 JOC.” (2 exs. AMGS). – Sudan: “S. Sudan R. Yei at Amadi 28.1. 1954 JJOC.” (1 ex. AMGS); “S. Sudan stream from hot springs Nyangwara 30,5E 4,39N 29.1. 1954 JJOC.” (6 exs. AMGS); “S. Sudan R. Yel at Amadi 28.1. 1954 JJOC.” (1 ex. AMGS); “S Sudan, sandy river 50 mi NW Juba 29.1. 1954 J. & J. Omer-Cooper” (5 exs. AMGS); “Equatoria Mundri-Lalyo 25-26.2. 1963 Linnavuori” (1 ex. MZH); “Blue Nile Ingessana Mts. 17-22.11. 1962 Linnavuori” (2 exs. MZH); “Equatoria Lalyo-Juba 26-27.2. 1963 Linnavuori” (7 exs. MZH). – Uganda: “Madi V. 1927 Carpenter” (1 ex. AMGS). – Kenya: “Selengai Riv. 21.6. 1970 E.S. Brown” (2 exs. BMNH); “River Athi Bushwackers´ Camp 29.3. 1964 E.S. Brown” (1 ex. BMNH); “Athi River Machakos District 14.9. 1976 M. Holmen / L. pilitarsis Rég. det. M. Holmen” (1 ex. ZMUC); “Nairobi Natnl Park 22.3. 1953 Hippo pools E.S. Brown” (1 ex. BMNH); “Kibwezi River Machakos District 13.9. 1976 Holmen M. / L. pilitarsis Rég. det. Holmen 1976” (8 exs. ZMUC); “Cha Shimba R., Kwale Kwale District 18.9. 1976 M. Holmen” (3 exs. ZMUC); “Mwatsuma R. Mariakani Kilifi District 16.9. 1976 M. Holmen” (2 exs. ZMUC); “Manjewa Riv Mariakani Kilifi / Kwale District 16.9. 1976 M. Holmen” (5 exs. ZMUC); “Maji ya Chumwi River Kwale District 16.9.1976 M. Holmen / L. pilitarsis Rég. det. M. Holmen 1976” (3 exs. ZMUC) “Afrique Orle Anglaise Voi Ch. Alluaud 1909 / Septembre” (2 exs. MNHN); “Afrique Orle Anglaise Voi Ch. Alluaud 1909 / Septembre / Det. Dr. Guignot Laccophilus pilitarsis Rég.” (1 ex. MNHN); “Fort Hall Br. O. A.” (1 ex. NHMB, 1 ex. ZMHB); “Mulango Br. O. A.” (1 ex. ZMHB);” Br. O. A. Kibwezi Scheffler” (1 ex. ZMHB). – Tanzania: “Mkulumuzi Riv. Paramba, Tanga District 26.9. 1976 M. Holmen / L. pilitarsis Rég. det. Holmen det.” (1 ex. ZMUC); “Sigi Riv. Ralley Estate Tanga District 26.9. 1976 M. Holmen / L. pilitarsis Rég. det. M. Holmen” (1 ex. ZMUC); “Kombe Stream Doda Tanga District 23.9. 1976 M. Holmen / L. pilitarsis Rég. det. Holmen” (2 exs. ZMUC); “TPC S Moshi canals 28.9. 1976 / L. pilitarsis Rég. det. Holmen” (3 exs. ZMUC); “Stream of Hegongo Tanga District 22.9. 1976 M. Holmen / L. pilitarsis Régb. det. Holmen” (1 ex. ZMUC); “Uluguru Mts. Kimboza Forest 250 m 18.7. 1981 Stoltze & Scharff leg. / L. pilitarsis Rég. det. Holmen 1981” (1 ex. ZMUC); “Kilimandj. Sjöstedt / Kibonoto 1-1200 m” (4 exs. NHRS); same data but “1000-1300 m” (1 ex. ZMHB); “? einem Tümpel nahe am Myanwaya Fluss 24.5. 1899” (1 ex. NHMB). – Zaire: “C.B. PNU, Kamusanga affl. g. Lufira / f. mt. Sombwe (750 m) 12.VII. 1949 Mis. De Witte 2776a” (1 ex. MNHN). – Zambia: “Muchinga Escarpment ca. 47 km ENE Rufunsa 14°57'S, 30°04'E lux Göllner leg. 25.3. 1993” (1 ex. ZMHB); “Mountain stream crossing road Kafue-Chirundu 9.8. 1986 Pederzani / L. pellucidus Sharp det. Rocchi 1990” (1 ex. CSR). – Malawi: “7 km W Golomoti SE14,34Bc 11.12. 1983” (2 exs. TMSA); “Nyasaland Ft. Hill Yambe Stream 17.10. 1948 JOC.” (1 ex. AMGS); “Nyasaland 40 mi. from Njakwa on Ft Hill rd 18.10. 1948” (1 ex. AMGS); “Nyasaland stream 20 miles from Dedza on Lower Lilongwe rd 30.9. 1948” (1 ex. AMGS); “Nyasaland Mtiti River 1.X. 1948 JOC.” (2 exs. AMGS); “Selima env. 80 km E Lilongwe 5-6.1. 2002 Kantner” (3 exs. NHMB); “Mulanje mnts env. 22-26.12. 2001 Kantner” (2 exs. NHMB). – Mozambique: “Mkura Stream on Chikukwa Camp, N-19.5516, E33.06916, 24.9. 2002 Bills” (1 ex. AMGS). – Zimbabwe: “S. Rhod., stream with lilies betw. Salisbury & Bromley 12.IX. 1948” (2 exs. AMGS); “Rhodesia Sebakwe” (2 exs. SAMC); “Shangani R. / 13.9. 1948 JOC. / L. pilitarsis Rég. det. J. Omer-Cooper” (2 exs. AMGS, 1 ex. TMSA); “Wankie Nat. Pk. Main Camp nr Pan S. / MV light trap 10.11. 1961 J.S. Weir” (1 ex. BMNH, 1 ex. MZH; habitus in Fig. 418); “S. Rhod. Wankie reserve Masumu dam 4.IX. 1948” (2 exs. AMGS). – Swaziland: “Little Usutu River nr. Mbabane 5.12. 1948 JOC.” (1 ex. AMGS). – South Africa: “Tv. Nelshoogte gallery forest below St. 25.51S-30.53E / 4.12. 1986 UV light collection Endrödy-Younga 2354” (2 exs. TMSA, 1 ex. MZH); “Kruger Nat. Pk Letaba Riv. bel. dam 23.46S-31.30E / shorewashing 1.3. 1995 Endrödy-Younga 3122” (4 exs. TMSA, 1 ex. MZH); “Kruger Nat. Pk Levuvu Riv.22.27S-31.10E / 12.2. 1994 shorewashing Endrödy-Younga 2998” (1 ex. TMSA); “Trsvl KNP, Gudzani, N-24.260, E31.840, 30.6. 1960” (1 ex. AMGS); “Trsvl, Wit R at Plaston, N-25.350, E31.070, 25.6. 1960” (3 exs. AMGS); “Trsvl, Polluted Pan, S of Olifants Gorge, KNP, N-24.010, E31.740 28.6. 1960” (1 ex. AMGS); “Trsvl, Satara KNP, Windmill dam, N-24.400, E31.770, 30.6. 1960” (2 exs. AMGS); “KNP survey Shingwedzi 19-20.11. 1961 Vári & Rork” (1 ex. TMSA); “Trsvl Warmbad 24-25.2. 1968 Spangler” (21 exs. USNM, 4 exs. MZH); “Trsvl Bundu Inn 25.28S-28.55E / 24.3. 1974 shore washing Endrödy-Younga 304” (1 ex. TMSA); “Fountains Pta 5.11. 1932 G. van Son” (7 exs. TMSA); “Transvaal Koop R. Barberton 5. Dec. 1948” (1 ex. AMGS); “OFS Parys” (1 ex. SAM); “Zululand Hluhluwe Game Res. 28.05S-32.04E / 27.11. 1992 shorewashing, shade, E-Y 2861” (2 exs. TMSA); “Natal Hluhluwe Game Reserve 18.4. 1951/Brinck-Rudebeck” (2 exs. MZLU); “Natal R. Natal National Park, the Hostel 5.4. 1951/Brinck-Rudebeck” (8 exs. MZLU); “Zululd Mtubatuba 28.22S-32.19E / 4.4. 1974 muddy shore washing, Endrödy-Younga 319” (3 exs. TMSA, 1 ex. MZH); “Natal Richmond 4.7. 1947 J.O.C.” (3 exs. AMGS, 8 exs. USNM, 1 ex. MZH); “Natal Little Bushmans River 15.4. 54 Oliffe” (2 exs. AMGS); “Umzikulu 6.4. 1947 J.O.C.” (3 exs. AMGS); “Natal Celenso (?) 12.11. 1953 / Tug 36Q3 12.11.53” (1 ex. AMGS); “Kw. Natal, Mhlatuze R above weir, Felixton N-28.840, E31.910, 7.3. 1962” (7 exs. AMGS); “Kw. Natal, below Masonite Effluent. Estcourt, Little Bushmans R., N-29.000, E29.880, 15.4. 1954 Oliff” (1 ex. AMGS); “Kw. Natal, Lufafa R confluence, N-30.001, E30.1825, 5.5. 1996 Dickens & de Moor” (2 exs. AMGS); “Kw. Natal, Mooi R, at Hornet Corner, N-28.950, E30.380, 15.3. 1995 Dickens” (3 exs. AMGS); “Kw. Natal, Mooi R, at Glenfem Bridge, N-29.390, E29.810, 15.3. 1995 Dickens” (1 ex. AMGS); “Kw. Natal, Klein Mooi R, at Durleigh Farm, N-29.230, E29.900, 15.3. 1995 Dickens” (1 ex. AMGS); “Kw. Natal, Betw. Aberfoyle, Impofana Farms, Mpofana R. N-29.400, E30.070, 3.1. 1995 Dickens” (1 ex. AMGS); “Kw. Natal, Mooi R., above Rosetta N-29.320, E29.970, 15.3. 1995 Dickens” (26 exs. AMGS); “Kw. Natal, Lions R, at Weltwreden Farm, N-29.440, Dickens” (1 ex. AMGS); “NW Pr., twin streams in Siyai, 10.3.1981 Reawell” (5 exs. AMGS); “Kw. Natal, Richards Bay, Umhlatuze floodplain 7.6. 1985 Reavell” (4 exs. AMGS); “Kw. Natal, Volkrust Road Bridge, Ncandu R., N-27.750, E29.930, 4.12. 1973 Metz” (6 exs. AMGS); same data but “24.9. 1974” (1 ex. AMGS); same data but “20.5. 1974” (1 ex. AMGS); same data but “8.1. 1974” (2 exs. AMGS); same data but “19.3. 1974” (2 exs. AMGS); Kw. Natal, Ncandu R. St. 7, 27.8. 1974 Metz” (2 exs. AMGS); “Kw. Natal, Ngagane R., below Newcastle Sewage works, N-27.720, E30.020, 5.12. 1973 Metz” (1 ex. AMGS); same data but “20.3. 1974” (1 ex. AMGS); “Kw. Natal, Ngagane R., Mandini-Iscor Rd. Bridge, N-27.720, E30.060, 24.9. 1974 Metz” (1 ex. AMGS); “Kw. Natal, Ngagane R., Steildrift rd., N-27.770, E30.020, 19.6. 1974 Metz” (1 ex. AMGS); “Kw. Natal, Injambili R, inland S coast rd., N-30.620, E30.520, 7.6. 1972 Chutter” (20 exs. AMGS); “Kw. Natal, Amahlongwa R, S Coast Rd. N-30.250, E30.720,9.6. 1972 Chutter” (12 exs. AMGS); “Kw. Natal, Izotsha R., Inland Coast Rd. 5.6. 1972 Chutter” (10 exs. AMGS); “Kw. Natal, Buffalo R, at Retreat, N-27.720, E30.180, 7.11. 1973 Metz” (1 ex. AMGS); “Kw. Natal, Little Amazimtoti R. N-30.060, E30.820, 15.6. 1984 Pretorius” (5 exs. AMGS); “Kw. Natal, Stn 10 at Colenso, Tugela R., N-28.730, E29.820, 12.11. 1953 Oliff” (2 exs. AMGS); “Kw. Natal, Ingane R., old S Coast Rd., N-30.170, E30.780, 15.6. 1964 Pretorius” (17 exs. AMGS); “Coward’s Bus Dam 26.6. 1993 Reavell” (9 exs. AMGS); “Nyebo Stream Transkei 5.4. 1947 J.O.C.” (2 exs. AMGS); “Butterworth Riv. 16.4. 1947 J.O.C.” (4 exs. AMGS); “Freddy van Zyl Bridge, Oorslas Spruit 25.2. 1947 J.O.C.” (1 ex. AMGS); “Modder River” (1 ex. SAMC); “Z.A. Barbeton Distr. Suid Kaap R. / Humus Oct. 1961 / Leleup” (1 ex. TMSA); “East London Fort Jackson pond by railway 14.III. 1955” (3 ex. AMGS); “E.C.Pr. Plutos Vale 20 July 1946 J.O-C.” (1 ex. AMGS); “ECPr., close to entrance of Dwesa Nature Reserve, stone bottomed river edge, S32°15.602, E28.46.368, Alt. 27 m, 23.1.1905 Bergsten leg.” (2 exs. NHRS); “Albany Blouwkrantz 1939” (1 ex. AMGS); “WPr., Mouth Swart R entering Hartebeesport Dam, in Eichhornia beed 30.5. 1971 Reavell (1 ex. AMGS); “CPr., Swart Kei R, nr Tylden, N-32.110, E27.020, 4.2. 1973 Stuart & Greig” (3 exs. AMGS); “ECPr., Bloukrans R. GHT Pt Alfred Road, N-33.375, E26.705, 9.9. 1972 Stobbs” (1 ex. AMGS); “ECPr., Mncotsho R., N-32.54.48, E27.36.52, 11.8. 2003, De Moor & Barber-James” (1 ex. AMGS); “ECPr., Mncotsho R, Trib. Buffalo R., N-32.54.48, E27.36.52, 11.2. 2003, De Moor & Barber” (1 ex. AMGS); same data but “N-32.54.43, E27.36.48, 18.5. 2004” (1 ex. AMGS); same data but “N-32.54.43, E27.36.48, 30.8. 2001 De Moor & De Moor” (2 exs. AMGS); “ECPr., Xolo R, Trib. carrying sewage works discharge, N-32.50.11, E27.37.49, 11.12. 2003 De Moor & Barber-James” (1 ex. AMGS); same data but “19.5. 2004” (3 exs. AMGS); same data but “2.10. 2002” (1 ex. AMGS); same data but “20.2. 2002” (1 ex. AMGS); “ECPr., Xolo R., upstream of confluence with trib., N-32.50.15, E27.37.48, 8.6. 2000 De Moor & Barber-James” (2 exs. AMGS); same data but “30.8. 2000” (1 ex. AMGS); “ECPr., Xolo R. Dam at Lily Stone Farm, N-32.52.10, E27.38.34, 16.5. 2001, De Moor & Barber-James” (1 ex. AMGS); “ECPr., Xolo R. Tributary, N-32.50.11, E27.37.49,1.12. 2004 De Moor & Barber-James” (1 ex. AMGS); “ECPr., Xolo R. Dam on Sidney Hill Farm, N-32.52.00, E27.37.09, 1.4. 2000, De Moor & Barber-James” (2 exs. AMGS); “ECpr., Rwantsa R., Dam, N-32.53.20, E27.37.55, 30.8. 2000, De Moor & Barber James” (1 ex. AMGS); “ECpr., Rwantsa R. at Farm Wolsley, N-32.54.03, E27.41.51, 2.10. 2002 De Moor & Barber-James” (1 ex. AMGS); same data but “10.12. 2003” (4 exs. AMGS); “ECpr., Rwantsa R. dam at Farm Mistrey, N-32.53.20, E27.37.55, 12.10. 2002 De Moor & Barber-James” (1 ex. AMGS); same data but “1.10. 2002” (2 exs. AMGS); same data but “15.5. 2001” (1 ex. AMGS); same data but “11.12. 2003” (5 exs. AMGS); “ECPr., Nahoon River at Witchkranz, N-32.51.10, E27.39.08, 19.5. 2004 De Moore & Barber-James” (2 exs. AMGS); same data but “4.5. 2000” (1 ex. AMGS); same data but “Nahoon R system 13.8. 2003” (2 exs. AMGS); “ECPr. Nahoon R. ca. 100 m above Dabadaba R confluence, N-32.50.28, E27.39.21, 10.12. 2003, De Moor & Barber-James” (1 ex. AMGS); same data but “19.2. 2002” (3 exs. AMGS); “ECPr., Maclear Munic. Dam, Mooi R. trib., N-31.0561, E28.31472, 27.3. 1993 Scott & al.” (3 exs. AMGS); “C.Pr., Upper Gatberg River, at Madun N-31.270, E28.170, 24.3. 1991 Barber-James & De Moor” (1 ex. AMGS)

Comments on synonymy

Earlier introduced synonymy of L. pellucidus, L. ampliatus and L. pilitarsis is confirmed. L. pellucidus, being the oldest, is the valid name of the species.

Diagnosis

Laccophilus pellucidus is especially characterized by large body combined with very peculiarly shaped penis, different from all other African Laccophilus species; penis voluminous with twisted, apical extension.

Description

Body length 5.3–6.0 mm, width 3.0–3.4 mm. Dorsal, aspect of body pale ferrugineous, without distinct colour pattern (Fig. 418). Elytra sometimes provided with pale, vague spots. Additionally, vague irrorations may be discerned on elytra.

Head: Pale ferrugineous. Slightly mat and distinctly microsculptured; reticulation double; large meshes contain 2–6 finer meshes. At eyes with an irregular row of fine punctures.

Pronotum: Pale ferrugineous, laterally with vague paler areas. Slightly mat, with distinct microsculpture. Reticulation generally of two different kinds; larger meshes contain 2–7 small meshes. At margins, with scattered, fine and irregular punctures.

Elytra: Pale ferrugineous to ferrugineous, without distinct colour pattern; sometimes with vague paler areas/spots (Fig. 418). Epipleuron posterolaterally not expanded. Weakly developed, fine irrorations often discernible. Rather shiny although finely reticulated. Reticulation double; large meshes contain 2-6 finer meshes. Posterolaterally double reticulation obscure and in part indistinct. Discal, dorsolateral and lateral, rather fine and somewhat irregular row of punctures discerned. Elytron posterolaterally with a row of fine and dense, quite long hairs.

Ventral aspect: Almost impunctate, except base of metathorax and two sclerites covered by apical ventrite; with distinct punctures. Rather shiny, extensive and very fine microsculpture discernible. Metacoxal plates and base of abdomen with striae. Prosternal process pointed, comparatively short and broad. Apical ventrite lacks distinct knob (Fig. 67).

Legs: Pro- and mesotarsus enlarged, somewhat extended and provided with distinct suckers.

Male genitalia: Penis almost straight, voluminous with twisted, apical extension (Figs 259–260).

Female: Rarely elytron laterally, between middle and apex, with a distinct lateral expansion (Fig. 8). Apical ventrite (Fig. 68). Pro- and mesotarsus slender.

Distribution

Sudan, Ethiopia, Uganda, Kenya, Tanzania, Zaire, Zambia, Malawi, Mozambique, Zimbabwe, Swaziland, South Africa (Fig. 560). Additional country record is Lesotho (Omer-Cooper 1965)

Collecting circumstances

Some biological information is available in Omer-Cooper (1958b); in most cases sampled in various kinds of streams but also in water holes. In Ethiopia sampled at high altitudes (between 5000-7500 feet) under the name L. pilitarsis (Omer-Cooper 1931) and as L. pellucidus at altitudes of 1500-2300 m (Nilsson and Persson 1993). See also Pederzani (1988).

Species group 10 (L. adspersus group)

Diagnosis. Small to large species; body length 2.9–5.1 mm, width 1.6–2.9 mm.

Shape of body oval to oblong, dorsoventrally flattened (Figs 419–420). Dorsal colour pattern variable; sometimes unicoloured without any pattern, sometimes with vague, dense irroration which in part can be reduced and finally species with distinct, often patchy colour pattern (Figs 426, 438, 443, 445). Dorsal microsculpture double; divided into two size-classes i.e. large and small meshes. Sometimes either kind can be reduced and at least in part indistinct or absent.

Prosternal process slender, posteriorly extended, apically pointed. (Prosternal process broken in unique specimen of L. amicus and state accordingly unknown). Apical ventrites with posterior end modified; excavated on each side of midline; at midline posteriorly extended (Figs 69–70), in males apical ventrite provided with an asymmetric knob on one side (Figs 71, 73). Stridulatory apparatus lacking. Metacoxal process not extended posteriorly (Fig. 6).

Paramere generally slightly enlarged, quite simple and exhibits generally no or moderate modifications (Fig. 268). Penis rather slender, always curved or angled; almost all species with a distinct apex which is variously modified (hooked, bifid, curved etc.) (Figs 263, 270, 291, 299).

Species composition and distribution. 24 species are recognized in this species group, which most probably is artificial and can be further split. No synapomorphous character for the group detected.

Comments. Laccophilus amicus, of which only female is known is characterized by small body (length 3.3–3.4 mm, width 1.8) and by peculiar elytral colour pattern (Fig. 442). L. amicus is most probably closely related to L. bellus, on the basis of external similarity.

To observe that present key is tentative and in determination both external features and male genitalia should be checked.

Key to species (males)

1 Elytra unicoloured, dark to pale ferrugineous; lack distinct colour pattern (Figs 444–446); small species (length 2.9–3.3 mm) 2
Elytra with variable colour pattern; generally larger species 3
2 Elytra pale ferrugineous to ferrugineous (Fig. 444) L. septicola (p. 145)
Elytra blackish ferrugineous (Fig. 445) L. pullatus (p. 147)
3 Elytra blackish ferrugineous with small, pale ferrugineous spots (Fig. 446); penis as in Figs 293–294; small species (length 2.9 mm) L. luteosignatus (p. 148)
Elytral colour pattern different; penis shape different; larger species (length min. 3.1 mm) 4
4 Penis strongly angled in lateral aspect (about 75–90°) (Figs 295, 301); large species, length 4.3–4.9 mm 5
Penis generally straighter (angle more than 100°); many species less than 4 mm in length 6
5 Elytral colour pattern distinct; with two transverse, pale, areas where irrorations reduced (Fig. 452); penis as in Fig. 301 L. guignoti (p. 157)
Elytral colour pattern vague, indistinct, no transverse pale areas (Fig. 447); penis as in Fig. 295 L. benoiti (p. 150)
6 Penis, lateral aspect, with small, sharp knob located close to inner curvature (Fig. 266) 7
Penis lacks corresponding knob (Fig. 276) 9
[Comment: note that some specimens of L. adspersus have a resembling knob on penis, which is less sharp and pronounced]
7 Colour pattern evenly distinct in basal half of elytra (Fig. 424); penis as in Fig. 268 L. nodieri (p. 113)
Scutellar region with reduced, indistinct elytral colour pattern (dark ferrugineous irrorations vague) (Fig. 422); penis different 8
8 Penis, lateral aspect, with apex outline rounded (Fig. 266) L. modestus (p. 107)
Penis, lateral aspect, with apex outline angled (Fig. 267) L. cryptos (p. 111)
9 Penis apex distinct, almost bifid (Fig. 299); large species (4.6–5.1 mm) L. vermiculosus (p. 154)
Penis apex different; generally smaller species 10
10 Penis apex simple, exhibits no modifications (Figs 297–298) (Madagascar) L. addendus (p. 151)
Penis apex variously modified (Mainland Africa, Madagascar) 11
11 Penis robust, apex large and outline evenly curved (Fig. 270); elytral colour pattern variable Figs 426–429 (sp. complex?) L. remex (p. 118)
Penis less robust to quite delicate, apex smaller, often differently shaped 12
12 Penis, lateral aspect, external outline angled (curvature smooth) (Figs 275–276) 13
- Penis, lateral aspect, external outline almost evenly curved (Figs 277, 283) 15
13 Penis apex large, distinct (Fig. 275) L. turbatus (p. 123)
Penis apex, small, hardly discernible (Fig. 276) 14
14 Elytral irrorations distinct (Fig. 433); penis apex broad (Fig. 276) L. pallescens (p. 125)
Elytral irrorations diffuse (Fig. 435); penis apex narrow (Fig. 278) L. mediocris (p. 132)
15 Extreme penis apex projects forwards (Fig. 277); elytra anteriorly at suture with distinct, quite long, narrow pale area without irroration (Fig. 434) L. trilineola (p. 130)
Extreme penis apex curved (Fig. 286); elytral colour pattern different 16
16 Penis, lateral aspect, inner outline with medial expansion (Fig. 285) L. enigmaticus (p. 138)
Penis, lateral aspect, inner outline lacks medial expansion (Fig. 283) 17
17 Penis apex broad, truncate, and turned upwards, appears in lateral aspect, narrow (Fig. 280) 18
Penis apex broad, truncate but not turned upwards, appears in lateral aspect broad (Fig. 269), or shape of penis apex different (Fig. 265) 19
18 Elytral colour pattern distinct (Figs 438–439; penis as in Fig. 283 L. saegeri (p. 136)
Elytral colour pattern diffuse (Fig. 436); penis as in Fig. 280 L. epinephes (p. 134)
19 Penis apex broad, hooked (Fig. 269); elytral colour pattern generally distinct (Fig. 425) 20
Penis apex different (e.g. Fig. 265); elytral colour pattern rather diffuse (irrorations less pronounced) (e.g. Fig. 419) 21
20 Large species (body length 4.2–4.6 mm); male genitalia (Fig. 269) L. flaveolus (p. 115)
Small species (body length 3.1–3.3 mm); male genitalia (Fig. 288) L. bellus (p. 144)
21 Penis, inner outline almost evenly curved from base to apex (Fig. 286) L. restrictus (p. 140)
Penis, inner outline angled, not evenly curved from base to apex (Fig. 265) 22
22 Penis apex (Fig. 265); elytral colour pattern (Fig. 420) (Madagascar) L. olsoufieffi (p. 105)
Penis apex variable (Figs 261–264); elytral colour pattern (Fig. 419) (Mainland Africa) L. adspersus (p. 97)

Laccophilus adspersus Boheman, 1848

Figs 69–70, 261–264, 419, 544

Laccophilus adspersus Boheman 1848: 246 (original description, faunistics); Sharp 1882: 287, 819 (description, faunistics); v. d. Branden 1885: 20 (catalogue, faunistics); Régimbart 1894: 237 (description, faunistics); Régimbart 1895: 135 (discussion, description, faunistics); Régimbart 1905: 208 (faunistics); Régimbart 1906: 248 (faunistics); Régimbart 1908: 5 (faunistics); Zimmermann 1920a:16 (catalogue, faunistics); Peschet 1925: 31 (faunistics); Omer-Cooper 1931: 756 (description, biology, faunistics); Gschwendtner 1932a: 12 (faunistics); Gschwendtner 1935a: 15 (faunistics); Guignot 1946c: 270, 273, 274, 276, 278, 312 (description, faunistics, discussion); Guignot 1953c: 145 (faunistics); Omer-Cooper 1956: 21 (faunistics, biology); Omer-Cooper 1957: 16, 18, 19, 90 (description, discussion, faunistics); Omer-Cooper 1958a: 59 (faunistics); Omer-Cooper 1958b: 37, 47 (discussion, description, faunistics); Guignot 1959a: 562, 566 (description, discussion, faunistics); Omer-Cooper 1962: 295 (faunistics); Bertrand 1963: 411 (discussion, faunistics); Omer-Cooper 1965: 77, 85 (description, faunistics); Bertrand and Legros 1967: 862 (faunistics); Omer-Cooper 1970: 290, 291, 292, 293, 294 (description, discussion, faunistics); Medler 1980: 155 (faunistics, list.); Bilardo 1982b: 251 (faunistics); Bilardo and Rocchi 1987: 104: (faunistics, biology); Pederzani 1988: 107 (faunistics, biology); Bilardo and Rocchi 1990: 162, 177 (faunistics, biology); Curtis 1991: 186 (faunistics); Nilsson and Persson 1993: 80 (discussion); Rocchi 2000: 24 (faunistics); Nilsson 2001: 240 (catalogue, faunistics); Bilardo and Rocchi 2002: 173 (list, faunistics); Pederzani and Reintjes 2002: 38 (faunistics); Reintjes 2004: 66 (faunistics); van Vondel 2005: 130 (faunistics, biology); Nilsson 2015: 208 (catalogue, faunistics).

Laccophilus livens Severin 1892: 472 (nomen nudum, discussion); Régimbart 1895: 135 (original description, faunistics); Zimmermann 1920a:21 (catalogue, faunistics); Zimmermann 1926: 24 (faunistics, discussion); Gschwendtner 1930: 88 (faunistics); Gschwendtner 1931: 180 (faunistics); Gschwendtner 1935a: 15: (faunistics); Gschwendtner 1938a: 5 (faunistics); Gschwendtner 1938b: 337 (faunistics); Guignot 1943: 99 (faunistics); Guignot 1946c: 269, 273, 276, 277, 312 (description, faunistics, discussion); Guignot 1953b: 234 (faunistics); Legros 1954: 268 (faunistics); Omer-Cooper 1957: 16 (synonym L. adspersus Boh.); Legros 1958: 211 (faunistics); Guignot 1959a: 558, 562, 565, 566 (description, discussion, faunistics); Guignot 1959d: 161, 162 (discussion, faunistics); Omer-Cooper 1965: 85 (list, synonymy); Omer-Cooper 1970a: 290, 291, 292, 29 (list, synonymy); Hernando 1990: 177, 178: (discussion, description); Nilsson 2001: 240 (catalogue faunistics, list, synonymy); Pederzani and Reintjes 2002: 40 (faunistics); Nilsson 2015: 208 (catalogue, faunistics). Confirmed synonym.

Laccophilus vitshumbii Guignot 1959d: 161 (original description, faunistics); Guignot 1961b: 238 (faunistics, discussion); Omer-Cooper 1970: 290, 293, 294 (discussion, description); Nilsson and Persson 1993: 58, 80, 94 (faunistics, biology); Nilsson 2001: 253 (catalogue, faunistics); Pederzani and Reintjes 2002: 40 (faunistics); Nilsson 2015: 219 (catalogue, faunistics). New synonym.

Laccophilus adspersus nigeriensis Omer-Cooper 1970: 291, 292, 293 (original description, faunistics); Medler 1980: 155 (catalogue, faunistics); Nilsson 2001: 240 (catalogue, faunistics); Pederzani and Reintjes 2002: 38 (faunistics); Nilsson 2015: 208 (catalogue, faunistics). New synonym.

Laccophilus adspersus sudanensis Omer-Cooper 1970: 292, 293 (original description, faunistics); Nilsson 2001: 240 (catalogue, faunistics); Pederzani and Reintjes 2002: 38 (faunistics); Nilsson 2015: 208 (catalogue, faunistics). New synonym.

Type localities

Laccophilus adspersus: South Africa: Caffraria interior.

Laccophilus livens: Zaire: Boma.

Laccophilus vitshumbii: Zaire: Lake Edouard, Vitshumbi.

Laccophilus adsperus nigeriensis: Nigeria: Jos.

Laccophilus adspersus sudanensis: Sudan: S of Rumbek near Wulu.

Type material studied

(56 exs.). Lectotype (by present designation): male: “Caffraria. / J. Wahlb. / Paratype / 3465 E91 / Naturhistoriska Riksmuseet Stockholm Loan no 1261/05” (NHRS). [Commments: no holotype was chosen in original description and neither has any lectotype thus far been designated. One male specimen is provided with Typus label but unfortunately its dissected genitalia are missing. We have dissected another male belonging to the type series (labelled paratype). Male genitalia is preserved in this specimen and we have chosen it therefore to be lectotype of L. adspersus Boheman.] – Paralectotypes: Same data as lectotype, but labelled as “Typus / Allotypus / Paratypus” (4 exs. NHRS).

Laccophilus livens: Lectotype (by present designation) male: Label with “male” symbol / Matadi Congo / Type / L. livens Rég.” (MNHN). – Paralectotypes: “Banana-Boma M. Tschoffen 91. Det. Régimb. 91 / 11174 / Régimbart det. 1891: Laccophilus livens Rég. / Ex. Typis” (7 exs. IRSNB). [Comment: one additional paralectotype in IRSNB with same data belongs to another species (= L. modestus Régb.)]; “Paratype / Banana-Boma M. Tschoffen 91. Det. Régimb. 91 / 11174 / Type Laccophilus livens Régt.“Type” (1 ex. BMNH); “Cotype / Congo / Laccophilus livens Régt.“Co-type”” (1 ex. BMNH); “Severin Banana Africa / Banana Boma M. Tshoffen 91 Dét. Régimb. / Laccophilus livens Rég. Type / Type” (1 ex. RMNH); same, but “Afr. occ.” (2 exs. RMNH); “Matadi M. Tshoffen / Laccophilus livens Rég. Types / SAM Type Acc. no. 839” (2 exs. SAMC); same data and “Cotype” (1 ex. IRSNB).

Laccophilus vitshumbii: Holotype: male: “Lac Édouard, Vitshumbi, 3043, mare I à Juissiaea M.T. 13-14.VI. 1953” (not studied; in IRSNB according to original description). – Paratypes: studied: “Congo Belge Lac. Edouard Vitshumbi, mare I + Juisseau MT 13-14.VI. 1953, 3043 / male symbol / Paratype” (2 exs. MNHN); “Congo Belge, Lac Edouaurd, Ishango d’Semliki (sur Graminées) 5.II. 1954, 3118a / Paratype” (1 ex. MNHN); “Male symbol / Congo Belge, Lac Edouaurd, Vitshumbi, mare II + Lemna 14.VI. 1953, 3042 / Paratype / L. vitshumbii (1 ex. MNHN).

Laccophilus adspersus nigeriensis: Holotype: male: “Type, male symbol / L. adspersus nigeriensis O-C. / Nigeria, Reservoir, stream, Jos 10.IV. 1963 J.O-C.” (AMGS). – Paratypes: “L. spp. / adspersus ? / Nigeria, stream & reservoir Jos 10.IV. 1963 J. O-C.” (4 exs. AMGS); “Nigeria, stream near Zaria 4.IV. 1963 J. O-C.” (4 exs. AMGS); “Nigeria (15A), stream Kaduna-Kontagora rd. 3.IV. 1963 J. O-C.” (1 ex. AMGS); “Nigeria, river, Jos-Bauchi rd. 9.IV. 1963 J. O-C.” (3 exs. AMGS); “Nigeria, river between Jos & Bauchi 9.IV. 1963 J. O-C.” (1 ex. AMGS); “Nigeria 27, detritus pond 45 miles from Jos on Bauchi rd 9.IV. 1963 J. O-C.” (14 exs. AMGS); adspersus / Nigeria, stream nr Bukuru 11.IV. 1963 J. O-C:” (1 ex. AMGS).

Laccophilus adspersus sudanensis: Holotype: male: L. adspersus subsp. sudanensis O-C. / Type / S. Sudan, rain ponds S. of Rumbek nr. Wulu 19.VII. 1954” (AMGS).

Additional material studied

(428 exs.). Sudan: “Rain ponds S of Rumbek nr Wulu 19.VII. 1954” (1 ex. AMGS); “Upper Nile Malakal 5-20.1. 1963 Linnavuori” (1 ex. MZH); “Equatoria, Mundri-Lalyo 25-26.2. 1963 Linnavuori” (1 ex. MZH); “Equatoria, Mwolo-Mundri 24.2. 1963 Linnavuori” (2 exs. MZH); “Equatoria, Lalyo-Juba 26-27.2. 1963 Linnavuori” (6 exs. MZH); “Equatoria, Loka Forest 8-10.4. 1963 Linnavuori” (1 ex. MZH); “Equatoria, Nzara 22.4. 1986 Wewalka / L. adspersus sudanensis O-C. det. Wewalka” (6 exs. CGW). – Ethiopia: “Water hole N Makki River, 6000 ft., 28.9. 1926 J. Omer-Cooper” (5 exs. BMNH); “Stream W of Zaquála 6000 ft., 27.10. 1926 J. Omer-Cooper” (1 ex. BMNH, 1 ex. MZH); “Hora Harsadi, Addas 1.12. 1926, 7000 ft, JOC” (1 ex. AMGS); “Hora Horeso 7000 ft., 1.12. 1926 JOC” (1 ex. AMGS). – Ivory Coast: “Comoé NP, N8,5° W3,5° Reintjes / 20.2. 1999 temporary creek” (1 ex. NMW). – Nigeria: “Kontagora pools in dry stream bed 3.IV. 1963 JOC.” (2 exs. AMGS); “Stream nr Zaria 4.IV. 1963 JOC.” (5 exs. AMGS); “Stream escarpment Jos-Wambe rd 13.IV. 1963 JOC” (1 ex. AMGS); “River between Jos-Bauchi 9.IV. 1963 JOC.” (4 exs. AMGS); “A stream nr Bakura 11.IV. 1963 JOC.” (1 ex. AMGS); “Stream Kaduna-Zaria rd 4.IV. 1963 JOC (10 exs. AMGS); “Kaduna-Zaria rd 4.IV. 1963 JOC” (2 exs. AMGS); “Stream & reservoir Jos 10.IV. 1963 JOC” (5 exs. AMGS); “Stream 86 miles from Makureli on Jos road 25.IV. 1963” (5 exs. AMGS); “Pools in dry stream bed Kontagora 5.IV. 1963 JOC” (1 ex. AMGS); “Pools, bridge over trib. of R. Niger, rd Kaduna-Kontagora” (1 ex AMGS); “Stream Kaduna-Kontagora rd 3.IV. 1963 JOC” (1 ex. AMGS); “Stream crossing Kaduna rd Zaria 8.IV. 1963 JOC” (1 ex. AMGS); “R. Kaduna 4.5 miles from Jos 13.IV. 1963 JOC.” (3 exs. AMGS). – Cameroon: “20 km NW Bangante Forest, savannah at river, at light 15.1. 1978/Gärdenfors, Hall & Samuelsson leg.” (1 ex. MZLU); “Maroua 26.8. 1973” (1 ex. NHMB). – Central African Republic: “Bozo 21.5. 1981/Degallier” (1 ex. MZH); same but “12. 1981” (1 ex. NHMB); same but “8. 1981” (1 ex. NHMB). – Zaire: “Longitshimo River, N-7.163, E20.880, 17.8. 2007 Graham” (1 ex. AMGS); “Lulimbi (Rutshuru) 1976 Lejeune” (2 exs. MRAC); “Parc National Garamba 29.9. 1951 De Saeger H. 2494” (2 exs. NHMB) “Parc National Garamba 28.8. 1952 De Saeger H. 3987” (1 ex. MRAC); “Parc National Garamba 27.6. 1952 De Saeger H. 3717” (1 ex. MRAC); “Parc National Garamba 5.5. 1952 De Saeger H. 3421” (1 ex. MRAC); “Parc National Garamba 19.3. 1952 De Saeger H. 3199” (1 ex. MRAC); “Parc National Garamba 6.2. 1952 De Saeger H. 3095” (1 ex. MRAC); “Parc National Garamba 3.4. 1952 De Saeger H. 3278” (6 exs. MRAC, 3 exx. MZH); “Parc National Garamba 2.4. 1952 De Saeger H. 3272” (2 exs. MRAC); “Parc National Garamba 31.8. 1952 De Saeger H. 3870” (2 exs. MRAC); “Parc National Garamba 1.8. 1952 De Saeger H. 3871” (1 ex. MRAC); “Parc National Garamba 4.4. 1952 De Saeger H. 3290” (1 ex. MRAC); “Parc National Garamba 1.9. 1952 De Saeger H. 4035” (5 exs. MRAC); “Katanga, Mwadingusha 21.5. 1965 Verheyen leg.” (1 ex, MRAC). – Uganda: “Kampala Hoima Rd 16.4. 1929 G.L.R. Hancock” (1 ex. AMGS); “Kampala 30.1. 1927 H. Hargreaves” (2 exs. AMGS). – Kenya: “Mombasa 25 km Nord palude de Kikalibala presso strada 14.7. 1968 Pederzani” (1 ex. AMGS); “Mombasa 30 km Nord Kikalibala swamp 14.7. 1968 Pederzani” (1 ex. AMGS); “Lambwe Valley, on light 11.6. 1974 van Etten” (2 exs. RMNH); “Manjewa R. Mariakani Kilifi / Kwale district 16.4. 1976 Holmen 4281” (1 ex. MZH); “Pond NE of Mariakani, Kilifi Distr. 16.9. 1976 Holmen 6076” (1 ex. ZMUC); Momb. Kilifi district 17.9. 1976 Holmen EF 8057” (1 ex. ZMUC); “Dam N of Gotani, Kilifi District 15.9. 1976 Holmen 5987” (1 ex. ZMUC); “Arabuko Sokoke Forest (30 km S Malindi) 8-24.6. 1998 Bartolozzi & Sforzi leg. alla luce” (1 ex. CSR); “Arabuko Sokoke Forest Res., Kilifi Distr., 20 km S Malindi/21.5.-7.6. 1994 Bartolozzi et al” (2 exs. CSR); “Thika 7.12. 1989 Jäch leg.” (1 ex. NMW); “Nairobi 3.11. 1967 / Reichart leg.” (1 ex. USNM); “Rabur 20.11. 1967 / Reichart leg.” (1 ex. USNM); “Kiserian 26.10. 1967/Reichart leg.” (1 ex. USNM, 1 ex. MZH); same data but “30.10. 1967” (1 ex. USNM); “Kibwezi Scheffler” (1 ex. ZMHB); “Meru Distr., Mourglia/Matiri (Mituguu) 8.11. 1983 800 m” (3 exs. NHMB); “Wa Kikuyu Bassin de l’Athi, Alluaud N. 1908” (1 ex. NHMB); “Br. O. A. Fort Hall” (1 ex. NHMB). – Tanzania: “Petukiza, ponds Tanga district 23.9. 1976 Holmen 1772” (1 ex. ZMUC); “Lukoka Pond, Tanga District 22.9.1976 Holmen 7230” (1 ex. ZMUC); “Tanganyika Ukerewe VII. 1933” (3 exs. OLML); “Kilimandjaro Sjöstedt 1905-1906/Kibonoto 1000-1300 m/21 Sept.” (1 ex. NHMB, 6 exs. NHRS, 3 exs. ZMHB); “Wembäre Steppe 6. 1911” (1 ex. ZMHB); “Pr. Shinyanga 60 km E Kahama 22.12. 2006 1150 m Kudrna Jr. lgt.” (1 ex. CFP); “Pond in stream bed, 107 mi from Dodoma 15.2. 1954” (1 ex. AMGS). – Angola: “Ca 10 mls W of Cainde, c. 3500 ft 15.4. 1954 / stagnant water hole, nitellid algae and muddy silt” (10 exs. BMNH, 2 exx. MZH); “Namakunda 6. 1948 16.15E. 18.50S C. Koch leg.”(1 ex. BMNH). – Zambia: “Kasempa env. 16-18.11. 2006, Z. Jindra leg.” (1 ex. NMPC); “Chinganganka 17.3. 1993 lux 15°53’ / 28°11'E, lux, hills, Uhlig leg.” (1 ex. ZMHB); “Kafue NP, Chunga Camp 26-29.3. 1993, 15°02'S / 26°00'E, lux, Göllner leg.” (1 ex. ZMHB); “Kafue NP, Chunga Camp 27.3. 1993, 15°02'35"S/26°00'09"E, lux, Uhlig leg.” (2 exs. ZMHB); “Africa Copperbelt Pr. Muekera 23.1. 1982 Selander / rain pond” (1 ex. MZH). – Malawi: “R Mtiti N of Lilongwe 1.X. 1948 JOC.” (1 ex. AMGS); “R Diedma Lilongwe rd. 30.IX. 1948 JOC.” (2 exs. AMGS); “River nr Dedza 28.IX. 1948 JOC.” (1 ex. AMGS); “Stream 20 miles from Dedza on lower Lilongwe rd 30.IX. 1948” (4 exs. AMGS); “Dedza dam on lower Lilongwe rd 29.9. 1948” (2 exs. AMGS); “Dedza env. 6-13.1. 2002 Bezdek leg.” (1 ex. NMPC); “Balaka env. 19-20.7. 2001 J. Bezdek leg.” (3 exs. NMPC, 1 ex. MZH); same but “5-6.1. 2002” (4 exs. NMPC, 1 ex. MZH); “Balaka env., 19.12. 2002 180 km SE Lilongwe Kantner” (1 ex. NHMB); “Stream (?) N of R. Mtiti X. 1948 / Paratype / L. simulator sp. n. det. J. O. Cooper” (1 ex. IRSNB; paratype L. simulator O-C.). – Zimbabwe: “Wankie Game Res. JOC. Waterholes / L. adspersus Boh. Det. JOC. (2 exs. AMGS); “Wankie Game Res. 5 Sept. 1948 JOC. Ponds at Robins Restcamp / L. adspersus Boh. Det. JOC.” (2 exs. AMGS); “Wankie Game Res. September 1948 JOC., waterhole / L. adspersus Boh. Det. JOC.” (4 exs. AMGS); “Wankie Game Res. 2.IX. 1948 JOC. / L. adspersus Boh. Det. JOC.” (2 exs. AMGS); ”5 mi SE Wankie 7.4. 1968 Spangler” (2 exs. USNM); “Gokwe Sengwa W.L.R.I. 28.12. 1982 -4.1. 1983 Bell / blacklight” (1 ex. NHMB); “Shangani R. 13.IX. 1948 J. O-C. / L. adspersus Boh. Det. JOC.” (2 exs. AMGS); “Stream Halfway hotel Gatooma Salisbury 14.IX. 1948 JOC.” (3 exs. AMGS); “Stream Halfway hotel Gatooma-Salisbury 14.IX. 1948 / L. adspersus Boh. Det. JOC.” (1 ex. AMGS); “Pool Lundi 22.N. 1948 J.O-C.” (5 exs. AMGS); “Sinkukwe 30 Dec. 1948 JOC.” (21 exs. AMGS, 3 exs. USNM, 1 ex. MZH); “Salisbury Mashonaland 1893 Marshall” (2 exs. SAM); “Nuanetsi River, Majinji Pan 4-5. 1961” (8 exs. BMNH, 1 ex. MZH); “Matopos NP 28.11-1.12. 1993, 20°33'S/28°30'E lux Uhlig leg.” (1 exs. ZMHB, 1 ex. MZH); “Gwai River 3.4. 1968 Spangler” (1 ex. USNM); “Ngezi N.P. env., 1.12. 1998 Kantner” (1 ex. NHMB); “Birkennough Bridge 24.1. 1998 Kantner” (1 ex. NHMB); “Mushandike Sanct. 10.12.1998 Kantner” (2 exs. NHMB, 1 ex. MZH); “Kariba env. 20.12. 1998 Kantner” (1 ex. NHMB); “Pond 26 mi. from Fort Victoria, Beit Bridge Rd.13.11.1948 J.O-C.” (2 exs. AMGS). – Namibia: “Omapapurawe Guard Post, 200 m from campsite, Cunene R., N-17.218, E13.645, pool, 15.11. 1997 Bethune et al. (1 ex. AMGS); “Kaokoveld, Sanitatas abt 85 mi WSW Ohopoho 14-16.6. 1951 / L. adspersus Boh. det. J. Omer-Cooper” (4 exs. MZLU); “Kaokoveld, Kowares 90 mi SE Ohopoho 3.6. 1951” (1 ex. MZLU). – Botswana: “Metsimaklaba 7-12.3. 1930 / L. livens Régb. det. Gschwendtner” (1 ex. OLML, 6 exs. TMSA); “N’Kate Makarikari 6-23.8. 1930 / L. adspersus Boh. det. Gschwendtner” (1 ex. TMSA); “Tsotsorogo Pan 17.6.-9.7. 30 / L. livens Régimbart det. Gschwendtner” (1 ex. OLML, 1 ex. TMSA); “Kasane 25-28.7. 1930 / L. addendus Shp det. Gschwendtner” (2 exs. TMSA). – South Africa: “Transvaal Sand R. 16.XII. 1953” (1 ex. AMGS); “Tshakoma Zpbg N. 1931 van Son / L. adspersus Boh. det. Omer-Cooper” (1 ex. TMSA); same but ”det. Gschwendtner” (1 ex. TMSA); “Valdesia Zpbg N. 1931 van Son / L. adspersus Boh. det. Gschwendtner” (1 ex. TMSA); “Trsvl Koring Spruit / Waterberg Dist. 20.8. 1948 J.O.C. / L. livens Rég. det. J. Balfour-Browne” (1 ex. TMSA); “Transvaal Kruger Park 1.VII. 1960” (2 exs. AMGS); “Kruger N.P. Skukuza, 12 km S, 25.04S, 31.37E / 6.3. 1996 UV light, Endrödy-Younga” (5 exs. TMSA, 1 ex. MZH); “Kruger N.P. Skukuza Res. camp, 24.59S, 31.36E / 25.2. 1995 UV-light & trap Endrödy-Younga” (4 exs. TMSA; habitus in Fig. 419); “Kruger N.P. Levuvu River, 22.27E, 31.10E / 12.2. 1994 shorewashing Endrödy-Younga” (2 exs. TMSA); “Kruger N.P. Letaba Riv. bel. dam 23.46S-31.30E / 1.3. 1995 shorewashing Endrödy-Younga” (2 exs. TMSA); “Trsvl, K.N.P., Pan 24 km S Satara Camp, N-24.610, E31.800, 18.6. 1960” (1 ex. AMGS); “Transvaal R. Nyl at Num Num 23.VIII 1948” (2 exs. AMGS); “Transvaal R. Nyl at Num Num 23.VIII 1948 / L. livens Reg. J. Balfour-Browne det.” (1 ex. AMGS); “RSA N. Prov. near Nylstroom 20.11. 2004 Werner & Smrz” (1 ex. NHRS); “Transvaal L. adspersus Boh. Det. J. Omer-Cooper” (2 exs. AMGS); “Trsvl Naboomspruit Torino Ranche 24.37S-28.38E / 15.1. 1989 UV light, vlei edge Endrödy-Younga” (3 exs. TMSA); “Trsvl Pretoria distr. Roodeplaat / 25-26.10. 1960 UV-light Neubecker” (2 exs. TMSA); “Pretoria 6.11. 1959 Janse / L. adspersus det. Gschwendter” (1 ex, TMSA); “Trsvl 5 mi W Warmbad 24-25.2. 1968 Spangler” (16 exs. USNM, 3 exs. MZH); “Trsvl Bundu Inn 25.28S-28.55E/24.3. 1974 at merc. vap. light Endrödy-Younga” (1 ex. TMSA); “Trsvl Swartspruit Mouth, Hartebeespoortdm. N-25.750, E27.900, 11.2. 1972 Reavell” (1 ex. AMGS); “Trsvl, stream in Magaliesberg Mts., mountain stream 11.9. 1972 Reavell” (1 ex. AMGS); “Plat R. 6-18.4. 05/Waterberg Distr. Swierstra” (2 exs. TMSA); “Frere Natal 1893 Marshall” (3 exs. SAMC); “Nat. -Drakensbg, Cathedral Peak, 28.57S, 29.12E/14.3. 1976 UV light station Endrödy-Younga” (2 exs. TMSA); “Kw. Natal Port Shepstone 20 km W 2.2. 2000 Halada” (2 exs. NMW); “Kw. Natal, McLeod’s Farm nr Dargle, Umgeni R., 4.2. 1989 Reavell” (1 ex. AMGS); “Kw. Natal Lundy’s Hill N-29.741, E29.872, marginal vegetation, stones, 30.4. 1996 de Moor et al.” (1 ex. AMGS); “Kw. Natal Gravesend estate N-30.170, E30.736, 12.10. 1996 Dickens et al.” (2 exs. AMGS); “Kw. Natal, Umlazi R., Tali Area, N-29.800, E30.520, 11.2. 1954 Oliff” (1 ex. AMGS); “Kw. Natal, Felixton, main drain from mill, N-28.840, E31.880, 3.7. 1962” (1 ex. AMGS); “Kw. Natal, Pond btw. Unizul and Mtunzini N-28.930, E31.750, marginal vegetation 20.9. 1995 Reavell” (1 ex. AMGS); “Kw. Natal Volkrust Road Bridge, Ncandu R., N-27.750, E29.930, 4.12. 1973 Metz” (1 ex. AMGS); “Kw. Natal, Below Newcastle Sewage Works, Ngagane R. N-27.720, E30.020, 19.6. 1994 Metz” (1 ex. AMGS); “Kw. Natal, Klip R. Stn. 3, 11.9. 1975 Sibbald & Brown” (1 ex. AMGS); “Kw. Natal, Ngogo R,. N-28.21.23, E29.43.25., 3.4. 1975 Metz” (2 exs. AMGS); “Kw. Natal, Izotsha R. Inland S Coast Rd N-30.780, E30.400, 5.6. 1972 Chutter” (1 ex. AMGS); “Kw. Natal, Little Amanzimtoti R, N-30.060, E30.820, 15.6. 1964 Pretorius” (1 ex. AMGS); “Natal Ladysmith 1000 m 29.12. 1993 Wewalka / L. adspersus Boh. det. Wewalka 1994” (3 exs. CGW); “Natal roadside puddles ca 2 km S Mbazwana to Hluhluwe nr Sodwana 5.3. 1997 Turner” (1 ex. NHMB); “Gauteng Tswaing 25.24S, 28.06E / 16.2. 2003 light trap” (1 ex. TMSA, 1 ex. MZH); “ECPr., close to Dwesa Nature Reserve, vegetation rich pond S32°17.027, E28°47.506, alt. 188 m 23.1. 2005 Bergsten” (1 ex. NHRS); “ECPr., close to Dwesa Nature Reserve, muddy pond with vegetation edges S32°18.582, E28°49.002, alt. 76 m 24-25.1. 2005 Bergsten” (1 ex. NHRS); “ECPr. Lusikisiki 19.III. 1956 JOC.” (1 ex. AMGS); “ECPr. Komgha quarry pond 20.III. 1955” (1 ex. AMGS); “ECPr. Quanbu 2.V. 1956” (1 ex. AMGS); “ECPr. St Johns 10.II. 1956” (3 exs. AMGS); “ECPr. Umzikulu 14.III. 1956 JOC.” (1 ex. AMGS); “Pirie Forest II. 1944 JOC.” (2 exs. AMGS); “ECPr. Mncotsho R., Trib. Buffalo Riv. N-32.54.43, E27.36.48, 18.2. 2002 de Moor” (5 exs. AMGS); ECpr. Nahoon R. at Witch Kranz, site NO, N-32.502, E27.392, 22.5. 2002, de Moor & Barber-James” (1 ex. AMGS); “ECpr., Dam on Rwantsa R, N-32.53.20, E27.37,55, 30.8. 2000 de Moor & Barber-James” (3 exs. AMGS); “ECPr. Rwantsa R. dam on Farm Mistley, N-32.53.20, E27.37.55, 10.11.2000 de Moor & Barber-James” (1 ex. AMSG); “ECPr. Rwantsa R. at Witchkranz, N-32.52.25, E27.38.34, 1.9. 2000 De Moor & Barber-James” (1 ex. AMGS); “ECPr. Rwantsa R. at Witchkranz, N-32.52.25, E27.38.34, 7.6. 2000 de Moor & Barber-James” (1 ex. AMGS); “ECPr., Rwantsa R. at Farm Wolsley, N-32.54.02, E27.51.95, 18.5. 2004 de Moor & Barber-James” (1 ex. AMGS); “ECPr., Rwantsa R. at Farm Wolsley, N-32.54.02, E27.41.51, 10.12. 2003 de Moor & Barber-James” (1 ex. AMGS); “ECPr., Rwantsa R. at Farm Wolsley, N-32.54.03, E27.41.51, 18.5. 2004 de Moor & Barber-James” (1 ex. AMGS); “ECPr. Rwantsa R. at Farm Sebastepol, N-32.53.00, E27.40.45, 7.5. 2000 De Moor & Barber-James” (3 exs. AMGS); “ECPr., Xolo R. trib. carrying sewage discharge, N-32.50.11, E27.37.49, 19.2. 2004 de Moor &Barber-James” (1 ex. AMGS); “ECPr., Mncotsho R., N-32.54.48, E27.36.52, 11.8.2003 de Moor & Barber-James” (1 ex. AMGS); “ECPr., Mncotsho R., N-32.54.48, E27.36.52, de Moor 15.5. 2001” (1 ex. AMGS); “ECPr., Mncotsho R., trib. Buffalo R., N-32.54.43, E27.36.48, 8.11. 2000 de Moor& Barber-James” (1 ex. AMGS); “ECPr., Mncotsho R., trib. Buffalo R., N-32.54.43, E27.36.48, 30.8. 2000 de Moor & Barber-James” (17 exs. AMGS); “ECPr., Mncotsho R., trib. Buffalo R., N-32.54.43, E27.36.48, 18.5. 2000 De Moor & Barber-James” (2 exs. AMGS); “ECPr., Nahoon R. at Witchkranz N-32,51,10, E27.39.08, 19.5. 2004, de Moor & Barber-James” (1 ex. AMGS); “ECPr., Nahoon R. at Witch Kranz N-32,50,28, E27.39.21, 22.5. 2002, De Moor & Barber-James” (1 ex. AMGS); “ECPr. Port, St Johns 15.2. 1956 / L. adspersus Boh. Det. JOC” (2 exs. AMGS); “ECPr. Mt Frere 8.V. 1956 JOC. / L. adspersus Boh. Det. JOC.” (1 ex. AMGS); “ECpr. 9.8. 1990 N-33.348, E26.678, Old Quarry Site, Manley Flats, dam, Baber-James & de Moor (2 exs. AMGS); “E Cape 9.3. 1997 Amatola Mts. 20 km NNE Aice 32°47'S 26°50'E Hess & Heckes” (1 ex. NMW); “EC., Hwy 352, 3 km S Tsomo, in river 22.5. 2005 Challet leg.” (1 ex. CGC); “North West 50 km S Kimberley Ritchie 12.1. 2000 Halada leg.” (1 ex. NMW). – Swaziland: “Little Usutu R nr. Bremersdorp 5.12. 1948” (1 ex. AMGS); “Bremersdorp 4.12. 1948 stream with muddy ponds JOC.” (2 exs. AMGS).

Comments on synonymy

The type material of all five taxa involved have been studied and compared (except holotype of L. vitshumbii; not found). Minor variation in shape of penis apex and dorsal colour pattern can be recognized. There are, however, a series of transitional morphs between the extremes both regarding genitalia and external appearance. Distribution covers extensive areas of Africa south of Sahara which justifies occurrence of minor morphological variation within a species. No clear morphological evidence and distributional pattern are thus present which would merit separation of species or subspecies. Accordingly earlier synonymy of L. adspersus and L. livens is confirmed. Furthermore L. vitshumbii, L. adspersus nigeriensis and L. adspersus sudanensis are all considered new synonyms of L. adsperus. Laccophilus adsperus being the oldest available name is the valid name of the species.

Diagnosis

Despite slight variation in shape of penis apex and elytra colour pattern in L. adspersus these features are still the best way of separation the species from other Laccophilus species. Laccophilus adspersus resembles most of L. olsoufieffi. Further study may show that they are also conspecific (see diagnosis of L. olsoufieffi on p. 106).

Description

Body length 3.6–4.2 mm, width 1.9–2.2 mm. Body almost unicoloured pale ferrugineous to ferrugineous; elytra with slightly indistinct irrorations. At base irrorations slightly sparser and often slightly reduced. Some specimens exhibit a variable pale spot with reduced irrorations posteriorly on each elytron (Fig. 419).

Head: Pale ferrugineous. Slightly mat to rather shiny, finely microsculptured; reticulation indistinctly double. Large meshes contain 2–6 fine meshes. Almost impunctate. At eyes with some scattered, fine punctures.

Pronotum: Pale ferrugineous. Rather shiny, although finely microsculptured; reticulation double. Finer meshes sometimes indistinct and hardly discernible. When discernible large meshes contain 2–8 finer meshes. Almost impunctate; punctures indistinct and hardly visible. Scattered punctures may be discerned laterally and at anterior margin.

Elytra: Pale ferrugineous. With ferrugineous to dark ferrugineous, slightly obsolete irrorations. Sometimes each elytron posterior to middle with a pale spot where irrorations reduced (Fig. 419). Rather shiny, although densely microsculptured. Reticulation double; laterally and posteriorly double reticulation becomes indistinct. Large meshes contain generally 3–8 fine meshes. Fine meshes in part weakly developed and difficult to discern. Discal, dorsolateral and lateral rows of punctures, irregular, very fine and in part hardly visible. Pre-apical furrow fine, sparsely pubescent.

Ventral aspect: Pale ferrugineous to ferrugineous. Almost impunctate. Rather shiny although very finely microsculptured. Metacoxal plates with in part reduced transverse furrows. Abdomen basally with sparse, somewhat curved striae. Apex of prosternal process slender and pointed. Apical ventrite with a small knob on one side (Fig. 69).

Legs: Pro- and mesotarsus slender, somewhat extended, provided with suckers.

Male genitalia: Note variation in shape of penis apex; extreme apex exhibits a gradual change from pointing straight forwards to, being somewhat curved and blunt (Figs 261–264).

Female: Externally as male but apical ventrite lacks asymmetric knob (Fig. 70). Additionally pro- and mesotarsus slender.

Distribution

Sudan, Ethiopia, Ivory Coast, Nigeria, Cameroon, Central African Republic, Zaire, Uganda, Kenya, Tanzania, Angola, Zambia, Malawi, Zimbabwe, Namibia, Botswana, South Africa, Swaziland (Fig. 544). Confusion in species delimitation has been common during the years and accordingly only personally verified records are accepted in the map.

Collecting circumstances

Insufficiently documented. Van Vondel (2005) reports the species to be sampled in a pond surrounded by tree, bottom covered with plant remains and water lily growths. Furthermore collected from pools with water lily and Pistia stratiotes and in stagnant remain of brooklet. Nilsson and Persson (1993) report L. vitshumbii collected at light, in temporary ponds in almost dry stream (1450–2350 m a.s.l.). Omer-Cooper (1931) recorded the species in high altitudes (5500–7500 ft). Additional information may be gathered from the literature, e.g. Bilardo and Rocchi (1987) and Pederzani (1988). Often collected at light and with light traps.

Laccophilus olsoufieffi Guignot, 1937

Figs 71–72, 265, 420–421, 545

Laccophilus olsoufieffi Guignot 1937: 141 (original description, faunistics); Gschwendtner 1938a: 5 (faunistics); Guignot 1941: 36 (description, discussion); Guignot 1946c: 269, 273, 274, 275, 276, 278, 312 (description, faunistics); Guignot 1959a: 558, 562, 565, 566 (description, discussion, faunistics); Bameul 1984: 94 (faunistics); Hernando 1990: 177, 178 (discussion, description); Rocchi 1991: 86 (faunistics, list); Nilsson 2001: 248 (catalogue, faunistics); Pederzani and Reintjes 2002: 40 (faunistics); Nilsson 2015: 215 (catalogue, faunistics).

Type locality

Madagascar: Maroansétra.

Type material studied

(8 exs.). Holotype: male: “Maroansétra, Madagascar X. 1936 / male symbol / Type” (MNHN). – Paratypes: males and females: “Madagascar Maroansetra X 1936 / Paratype” (1 ex. IRSNB); “Antakotako Madagascar II 1936 / female-symbol / Paratype” (1 ex. IRSNB); same data but “male symbol” (1 ex. MNHN; habitus in Fig. 416); same data, but with “n. spec. det. Gschwendt”, (1 ex. MNHN); “Male / Madagascar Vatomandry VIII. 1934 Vadon / Paratype” (1 ex. AMGS); “Madagascar Antakotako 11. 1936 / female-mark / Paratype” (2 exs. AMGS).

Additional material studied

(12 exs.). Madagascar: “Tananarive 7. 1934 Vadon / Lac Tzimbamzaza / male symbol / Type / ab. fuscinus” (1 ex. MNHN). [Comment: the specimen has no status as type material being associated with the name ab. fuscinus, which is infrasubspecific.] – “E-Mad. Ampamoho nr Andilamena 1200-1300 m asl.18-20.1. 1995 Dunay & Janak” (7 exs. NMW, 2 exs. MZH; habitus in Fig. 420); “Ese 5 km S Ampamoho pr. Andimalena 1. 1995 G. Dunay & J. Janak leg.” (1 ex. NMPC); “Toliara Menabe, Kirindy RS, S20.07655, E044.67532, 57 m.a.o., 12.12. 2009, water net, field, Bergsten et al. / 000000470 NHRS-JLKB” (1 ex. NHRS).

Diagnosis

Resembles most of and probably closely related to L. adspersus from which L. olsoufieffi can generally be distinguished by study of the penis. Minor difference can be recognized in bending of the penis. Moreover, body of L. olsoufieffi seems to be slightly more robust than L. adspersus in general. In L. olsoufieffi irroration covers often almost entire elytron but sometimes there is posterior to middle a patch with sparse irroration or irroration is totally absent. Further study may reveal that the two species are synonymous.

Description

Body length 3.8–4.3 mm, width 2.1–2.4 mm. Specimens regarded as aberration “fuscinus” are slightly larger; length 3.9–4.4 mm, width 2.2–2.5 mm. Additionally “fuscinus” lacks pale area (irroration absent or strongly reduced) posterior to middle of elytron. Habitus and dorsal colour pattern (Figs 420–421).

Head: Pale ferrugineous. Slightly dull, rather finely reticulated. Reticulation double; large meshes contain 3–4, often indistinct small meshes. Almost impunctate, except at eyes; fine and scattered punctures may be discerned.

Pronotum: Pale ferrugineous to ferrugineous; no distinct colour pattern. Submat, reticulated; reticulation quite distinct and double. Large meshes may contain 4–7 small meshes. Anteriorly and laterally with fine, in part indistinct, scattered punctures.

Elytra: Pale ferrugineous, extensively provided with ferrugineous irrorations (Figs 420–421). Somewhat posterior to middle with a vague transverse area where irrorations extensively absent (forming a vague transverse pale marking interrupted by suture). Rarely pale area lacking; “ab. fuscinus”. Submat, finely and quite distinctly reticulated. Reticulation distinctly double; large meshes contain generally 3–6 smaller meshes. Laterally, sublaterally and discally with sparse and irregular punctures (forming longitudinal areas with scattered puncture). Lateral, pre-apical furrow fine, finely pubescent.

Ventral aspect: Pale ferrugineous to ferrugineous, distinct colour pattern lacking. Rather shiny to submat; extensively with fine reticulation, which in part is rather indistinct. Basal ventrites with rather distinct, curved striae. Almost impunctate. Apex of prosternal process slender, slightly extended and pointed. Metacoxal plates in anterior half with fine, transversely located, shallow furrows; in posterior half furrows absent. Apical ventrite asymmetric, with knob on one side (Fig. 71).

Legs: Pro- and mesotarsi slightly enlarged, extended, with suckers.

Male genitalia: Penis long, bended and extreme apex points forwards (Fig. 265).

Female: Apical ventrite symmetric, lacks knob (Fig. 72). Protarsus slender; claws slightly extended and moderately curved.

Distribution

Madagascar (Fig. 545). Records outside Madagascar are to be considered uncertain.

Collecting circumstances

Not documented.

Laccophilus modestus Régimbart, 1895

Figs 73–74, 266, 422, 546

Laccophilus modestus Régimbart 1895: 133 (original description, faunistics); Régimbart 1906: 248 (faunistics, disussion.); Zimmermann 1920a: 23 (catalogue, faunistics); Guignot 1946c: 270, 273, 276, 278, 312 (discussion, description, faunistics); Guignot 1952c: 521 (faunistics); Capra 1952: 6 (faunistics); Guignot 1953b: 234 (faunistics); Guignot 1955b: 1096 (faunistics); Guignot 1956a: 88 (faunistics); Omer-Cooper 1958b: 37, 47, 48, 49 (discussion, description, faunistics); Guignot 1958: 8 (discussion); Guignot 1959a: 562, 566, 568 (description, discussion, faunistics); Guignot 1959d: 162 (discussion, faunistics); Guignot 1961b: 238 (faunistics); Omer-Cooper 1965: 77, 87 (description, faunistics); Omer-Cooper 1970: 293 (description); Legros 1972: 466 (faunistics); Bilardo and Pederzani 1978: 119 (faunistics, description); Medler 1980: 155 (faunistics, list); Bilardo and Rocchi 1990: 177 (faunistics); Nilsson and Persson 1993: 81, 94 (faunistics, biology); Nilsson et al.1995: 505 (faunistics); Rocchi 2000: 24 (faunistics); Nilsson 2001: 247 (catalogue, faunistics); Bilardo and Rocchi 2002: 174 (list, faunistics); Pederzani and Reintjes 2002: 40 (faunistics); Reintjes 2004: 68 (faunistics); Nilsson 2015: 214 (catalogue, faunistics).

Laccophilus modestus v. tostus Régimbart 1895: 133, 134 (original description, faunistics); Zimmermann 1920a: 23 (catalogue, faunistics); Guignot 1946c: 270 (description); Guignot 1959a: 568 (female description, faunistics); Guignot 1959d: 162 (discussion, faunistics); Bilardo and Pederzani 1978: 119 (faunistics, description); Nilsson 2001: 247 (catalogue, list, synonymy L. modestus Régimbart); Nilsson 2015: 214 (catalogue, faunistics, list, synonymy). Confirmed synonym.

Laccophilus espanyoli Hernando 1990: 177 (original description, faunistics); Nilsson 2001: 243 (catalogue); Pederzani and Reintjes 2002: 40 (faunistics); Nilsson 2015: 211 (catalogue, faunistics). New synonym.

Type localities

Laccophilus modestus: Mali: Badoumbé (Ht. Senegal).

Laccophilus modestus var. tostus: Gabon: Cap Lopez.

Laccophilus espanyoli: Senegal: Oussaduye.

Type material studied

(9 exs.). Laccophilus modestus: Lectotype (by present desgination): male: “Ht. Sénégal Badoumbé Dr. Nodier I à V – 1882 / male symbol / co-type / L. modestus Rég.” (MNHN). [Comment: Guignot (1959a) indicates existence of a lectotype but survey of collections in Paris museum reveals that no such specimen can be distinguished; see also Nilsson (2001).] – Paralectotypes: Same data as lectotype but, labelled with female symbol (1 ex. MNHN); “Badoumbé / Museum Paris coll. Maurice Régimbart 1908 / modestus Rég. Koppi Wehncke” (1 ex. MNHN).

Laccophilus modestus var. tostus: Cotype: female: “Gabon Mocquerys / female symbol / Cotype” (MNHN). Additionally, three specimens mounted together and labelled “Gabon Mocquerys/Museum Paris Coll. Maurice Régimbart 1908 / modestus Rég. v. tostus Rég.” probably also belong to the type material but have no type indication (3 exs. MNHN).

Laccophilus espanyoli: Holotype: male: “Holotip / Senegal Oussaduye 13-XI-65 Sala leg. / Laccophilus espanyoli sp. n. C. Hernande det. / 78-0572 MZB” (MZBS). – Paratype: female: Same data as holotypus but “Paratypus / 78-0752 MZB” (1 ex. MZBS).

Additional material studied

(371 exs.). Gambia: “Abuko Nat. Res., at light at the Bamboo Pool 18.30-20.30, 18.11. 1977 UTM 28PCK2181 / Cederholm et al. N. 1977” (3 exs. MZLU); “Tendema Camp, at light in semiarid veg near river Gambia 18.30-20.30, 14.11. 1977, UTM 28POK1285, loc. 12A / Cederholm et al. N. 1977” (1 ex. MZH); “Riv. Tanji 3 km SW Brufut. At light 19.00-21.00, 28.2 1977, UTM 28PCK087773 / Cederholm et al. Febr.-March 1977” (2 exs. MZLU, 1 ex. NHMB); “Outside Abuko Nat. Res., at waterworks. At light 19.00-22.00 UTM 28PCK214812 / Cederholm et al. Febr-March 1977” (2 exs. MZLU); “Bathurst Jan. 68 Palm / L. modestus Régb.det. Persson” (5 exs. MZLU); “Bathurst Januari 1968 Leiler” (3 exs. NHRS); “Kuntaur NW Georgetown 21.11. 2003 Vondel” (1 ex. CSR). – Gambia/S. Senegal: “Stream N of Selety 13°10'N, 16°36'W, 19.2. 1976 Holmen” (2 exs. ZMUC). – Senegal: “Riv. Cazamance Carabane Dr. Collin / Museum Paris coll. Maurice Régimbart 1908” (2 exs. MNHN); “3 km SSW Toubakouta, 10 km S Ziguinchor, 4.3. 1977, at light 19.00-22.00, Loc. No. 16, UTM 28PCJ585782 / Cederholm et al. Febr-March 1977 / L. modestus Rég. det. M. Brancucci” (3 exs. MZLU, 6 exs. NHMB); “Swamps ca 3 km SW Ziguinchor 8.3. 1977, UTM 28 PJC59-89- / Cederholm et al. Febr- March 1977” (5 exs. MZLU); “2.5 km ESE Ziguinchor in cultivated area, at light 20-21.30, 11.11. 1997 / Cederholm & al. (1 ex. NHMB); “In forest, 1 km NE Djibelor, ca 7,5 km SW Ziguinchor, at light 19-21. 9.11. 1977 / Cederholm & al.” (1 ex. NHMB); “Ht Senegal Badoumbé 1-5. 1882 Nodier” (1 ex. SAMC); “60 km S Velingara Pakour 27.6. 2004 leg. Marek Halada” (2 exs. NMPC); “1 km NW Bignona 26 km N Ziguinchor, at light 19.15-20.30, 3.3. 1977, UTM 28PCK654170 / Cederholm & al.” (1 ex. NHMB). – Guinea Bissau: “Cachheu 12 km E Varela, earth pit pond, 9.4. 1993 Persson” (12 exs. MZLU). – Guinea: “Seredou 4.4. 1975, lux Zott leg.” (2 exs. ZMHB). – Mali: “Haut Sénégal Khayes Dr. Nodier 11-12 1881 / female symbol / Co-type / L. modestus Rég.” (2 exs. MNHN; not type material due to deviating label data); “Kogoni 10. 1966 Schmitz” (1 ex. MRAC, 1 ex. MZH); “K. Macina 10.11. 1973 Reynolds” (18 exs. BMNH, 4 exs. MZH); “San, Bani river 13°18'N, 4°54'W, 22.2. 2000 Komarek & Meyer / L. modestus Régb. det. Wewalka 2001” (1 ex. NMW); “Kéniéroba 70 km SW Bamako 12°06’ N 8°20'W, 1. 2011 Kravchenko” (1 ex. TAU). – Niger: “Niamey 9. 1988, at light, Jongema / L. modestus Régb. det. Wewalka 2005” (7 exs. CGW). – Burkina Faso (= Ht Volta): “Ouagadougou X. 1926” (1 ex. NHMB); “Haute Volta Bobo Dioulasso / Museum Paris 12 – 1930 – IV 1931 Ch. Alluaud & P. Chappuis / male symbol / L. modestus Rég.” (1 ex. MNHN; habitus in Fig. 422); “Nadiagow MV August 2005 Moretto” (3 exs. NHMB). – Chad: “nr Bongor 27.5. 1973 Linnavuori” (2 exs. MZH). – Sudan: “Upper Nile, Malakal 5-20.1. 1963 Linnavuori” (12 exs. MZH); “Sudan Malakal 1963 Linnavuori” (1 ex. MZH); “Dahr el Ghazal, Wau 19.2. 1963 Linnavuori” (2 exs. MZH); same but “R. Malmul 21.2. 1963” (1 ex. MZH); Equatoria Lalyo-Juba 26-27. 2. 1963 Linnavuori” (1 ex. MZH); “Gilo water tank (pumped up from stream) 20.3. 1980 Armstrong” (1 ex. USNM); “Kinyetti River at Imeila 19.3. 1980 Armstrong” (2 exs. USNM); “Senaar a. bl. Nil, lux 21.10. 1979 Hieke” (1 ex. NHMB); “Nyangwara, stream from hot springs, N4.39, E30.5, 29.1. 1954 J. Omer-Cooper” (1 ex. AMGS); “Sandy River 50 mi NW Juba 29.1. 1954J. Omer-Cooper” (1 ex. AMGS); “Moya Sawu 45 mi from Amadi-Juba rd 29.1. 1954” (1 ex. AMGS). – Liberia: “Suakoko 8.4. 1953” (3 exs. USNM); “Suakoko 18-19.3. 1952 / Blickenstaff Light trap” (6 exs. USNM, 2 exs. MZH); “Suakoko 27.2. 1952 Blickenstaff” (1 ex. USNM); “Suakoko 22.25.2. 1952 Blickenstaff” (2 exs. USNM); “Suakoko 14.3. 1952 / 6-9 pm light trap” (2 exs. USNM). – Sierra Leone: “Makeni 12°03'W, 8°53'N, 27.11. 1993, light trap / Cederholm-Danielsson-Hall / L. modestus Régb. det. Nilsson 94” (10 exs. MZLU; 2 exs. MZH); same as but “det. Persson” (4 exs. MZLU); “Kalangba 8.11. 1980 D. Jump leg.” (1 ex. USNM). – Ivory Coast: “Bingerville 1-12.3. 1962 Decelle” (1 ex. MRAC); same but “7. 1962” (1 ex. MRAC); “Comoé NP, N8,5° - W3,5°, Reintjes / 9.1. 1999” (2 exs. NMW); same but “28.2. 1999” (1 ex. NMW); same but “21.3. 1999” (1 ex. NMW); “Touba, à la lumière 4. 2002 Moretto / L. modestus Régb. det. Rocchi 2002” (1 ex. CSR). – Ghana: “Upper East Pr., Navrongo env. 11-13.6. 2006 Pokorny S.” (2 exs. NMPC); “N Reg. Nyankpala 15 km W von Tamale leg. Endrödi / Lichtfalle 1-30.4. 1970” (1 ex. CGW); “N Reg. Damongo Mole game res. 220 m, N9°04’ – W1°48’ Endrödy-Younga / on light 12.8. 1971” (1 ex. CGW); “Ashanti Reg. Kumasi Nhiasu 330 m, N 6.43-W 1.36 Endrödy-Younga / at light 12.6. 1967” (1 ex. CGW); “Volta Reg., Volta Riv. at Kpong 28.11. 1993 light trap T. Andersen leg.” (1 ex. MZH). – Ethiopia: “Bahar Dar 8.10. 1968 Harde leg.” (2 exs. CGW). – Benin: “Zagnanado Dahomey” (1 ex. MNHN); “Dep. Atlantique, Allada, Glotomè (village) 31.1.2006 leg. Goergen et al. / 06°41'06,8"N, 02°02'36,8"E, 17 m asl, slowly running stream” (3 exs. NMW, 1 ex. MZH); same but “Awoute / 06°39'54,9"N, 02°09'34,1"E, 25 m asl, small ponds” (1 ex. NMW); “Dep. Zou, Zogbodomé, Lokoli (forest), Hlanzoun riv. 3, 6.2. 2006 leg. Goergen et al. / 0.7°03’ N, 02°15’ E, muddy stream” (6 exs. NMW, 2 exs. MZH);“Dep. du Zou, commune de Zogbodomé 29.1. 2006 Goergen / Lokoli forest 07°03'N, 02°15'E, 17 m asl, light trap” (4 exs. NMW, 1 ex. MZH); “Calav i IITA, light trap: fallow 20.6. 2004 Goergen” (1 ex. NMW). – Nigeria: “Ibadan ca. Jan-Juni 1954 Stenholt Clausen / L. mediocris G. det. J. Balfour-Browne” (1 ex. ZMUC); “Ibadan, at light 27.11. 1955” (1 ex. BMNh); same but “26.9. 1956” (1 ex. BMNH); “NW St. Badeggi rice fields 8-9.8. 1973 Linnavuori” (133 exs. MZH); “W. St. Ife 7-8.7. 1973 Linnavuori” (1 ex. MZH); “EC St. Norcap near Abakaliki 29.6. 1973 Linnavuori“ (2 exs. MZH); “Zaria 1969 à la lumière Roberts” (1 ex. MRAC); “Zaria pr. Zaria 5-6.3. 1949 Malkin” (1 ex. BMNH); “Zaria 1969” (1 ex. NHMB); “Samaru Endrödy Younga / Light trap 20.10. 1969” (1 ex. CGW); “River, Bauchi rd 21 mi from Jos 9.4. 1963 J. Omer-Cooper” (1 ex. AMGS); “Katsina-Dawyra rd, Marsh, 6.4. 1963 JOC” (4 exs. AMGS). – Cameroon: “Akonolinga, moist secondary forest and plantation, at light, 7.1. 1978 / L. modestus Régb. det. Nilsson” (2 exs. MZLU); “Yaounde Bor to Kosti by boat 3-14.3. 1978 Perkins leg.” (8 exs. USNM, 1 ex. MZH); “Maroua 26.8. 1973” (1 ex. NHMB); “Maroua, Nayo Tsanaga 26.10. 1977” (1 ex. NHMB); “Emana Obala 16.5. 1970” (1 ex. NHMB). – Central African Republic: “Bozo, lum. 11. 1981 / Degallier” (1 ex. NHMB). – Congo: “Parc Nat. d’Odzala, Mboko-Lango 21.8. 2002 Bilardo” (4 exs. CSR). – Zaire: “Tshuapa, Mbandaka ca. 0°03'N - 18°28'E, 8-9.3. 1963 a.l., Stam” (1 ex. RMNH); same but “15-16.3. 1963” (1 ex. RMNH); “Banana Boma M. Tschoffen 91 Det. Régimb. / L. livens Régb. det. Régimbart 1891 / Ex Types” (1 ex. IRSNB; type material of L. livens); “Boma M. Tschoffen” (1 ex. SAMC); “PNG I/c/2”’, 25.2. 1950 Demoulin 259” (1 ex. IRSNB); “PNG I/a/2”’, 30.1. 1950 Demoulin 240” (1 ex. IRSNB); “PNG I/b/3’, 15.2. 1950, Demoulin 253” (1 ex. NHMB); “PNG I/a/4, 6.3. 1950, Demoulin 297” (1 ex. NHMB).

Specimens with uncertain determination

Tanzania: “Deforested place nr Mangula, 297 m, at light, 18.7. 2004 Sprecher” (1 ex. NHMB; single female specimen); “Mizimu Mwanihana Mnts N.P. S07.48.21,8, E36.51.09,5, 850 m, 3-6.8. 2010 light trap Smith & Takano” (1 ex. BMNH; single female specimen). – Mozambique: “Mandambuzi, Manda Wilderness Res. S12°17.697’, E34°46.260’ Watson 16.2. 2008” (1 ex. CGF; single female specimen).

Comments on synonymy

The lectotype of L. modestus and holotype of L. espanyoli have been examined and compared. Morphological features in shape of penis and external appearance of body show that the two taxa are conspecific. Laccophilus modestus, being the older name is the valid name of the species. Earlier established synonymy of L. modestus var. tostus and L. modestus is also confirmed (name given for female being dimorphous, vide below under female description).

Diagnosis

Laccophilus modestus is characterized by appearance of elytra; irrorations reduced basally, and by features exhibited by the penis; inner outline of penis provided with a minute but distinct knob. Extreme apex of penis frontally rounded (vide diagnosis of L. cryptos on p. 112).

Description

Body length 3.5–3.8 mm; width 1.9–2.1 mm. Elytral irrorations are dark ferrugineous to ferrugineous against pale ferrugineous background (Fig. 422). Most specimens have vague irrorations on elytra but basal part of elytra to a variable degree lacking darker markings.

Head: Pale ferrugineous. Submat, finely microsculptured. Reticulation double, but due to minor size difference division in two mesh-size classes difficult. Almost impunctate; at eyes small area with fine, dense and irregular punctures.

Pronotum: Pale ferrugineous, no distinct colour pattern. Submat, finely microsculptured. Reticulation double; large meshes contain 3–6 small meshes. At margins with very fine, irregular punctures. Basally punctures hardly visible; only laterally clearly discernible.

Elytra: Pale ferrugineous, with slightly vague ferrugineous to dark ferrugineous, quite dense irrorations. Sometimes, irrorations anteriorly reduced and almost absent (Fig. 422). Submat, finely microsculptured. Reticulation double, but large meshes only discernible in mediobasal area. Large meshes, when discernible, contain 3–6 small meshes. Very fine, somewhat irregular punctures form a discal, dorsolateral and lateral row, out of which the two latter rows are rather indistinct.

Ventral aspect: Pale ferrugineous to ferrugineous, no colour pattern exhibited. Almost impunctate. Scattered, single punctures may be discerned. Rather shiny, although very finely microsculptured. In part reticulation reduced or absent. Abdomen with fine, slightly curved striae. Prosternal process slender, apex extended and pointed. Metacoxal plates frontally with indistinct, shallow furrows. Apical ventrite asymmetric, with single, minute, lateral knob (Fig. 73).

Legs: Pro- and mesotarsus slightly enlarged, somewhat extended and provided with distinct suckers.

Male genitalia: Penis (Fig. 266) in lateral aspect with extreme apex rounded and not sharply angled as in L. cryptos (Fig. 267).

Female: Externally resembles male but pro- and mesotarsus slender. Apical ventrite lacks knob, almost symmetric (Fig. 74). Some female specimens have strongly developed dorsal reticulation making body dull. Female is accordingly dimorphous. This extreme morph was named var. tostus and here listed as synonym of L. modestus.

Distribution

Gambia, Senegal, Guinea Bissau, Guinea, Mali, Niger, Burkina Faso, Sudan, Ethiopia, Liberia, Sierra Leone, Ivory Coast, Ghana, Benin, Nigeria, Cameroon, Central African Republic, Gabon, Congo and Zaire (Fig. 546). A single female specimen from Mozambique and two females from Tanzania are considered uncertain. Due to widespread confusion of species-status and delimitation of it, only verified records are included in the map. Additional country records are Tanzania (Zanzibar) (Régimbart 1895), Kenya (Régimbart 1906), Somalia (Capra 1952), Malawi (Omer-Cooper 1958b) and South Africa (Omer-Cooper 1965).

Collecting circumstances

Information in literature is uncertain but can be found by checking the references above. Label data indicate that the species occur both in stagnant and running waters: collected in swamps and small ponds as well as in a slowly running stream and in a muddy stream. Also collected at light, e.g. in semiarid vegetation near a river and in moist secondary forest and in a plantation.

Laccophilus cryptos sp. n.

Figs 75–76, 267, 423, 547

Type locality

South Africa: Zululand, Mission Rock, St. Lucia (28.22S-32.35E).

Type material studied

(24 exs.). Holotype: male: “S. Afr.; Zululand St. Lucia, Mission Rock 28.22S-32.35E / 18.12. 1975; E-Y: 983, at black light leg. Endrödy-Younga” (TMSA). – Paratypes: Same data as holotype (3 exs. TMSA, 1 ex. MZH; habitus in Fig. 423); “S. Afr.; Zululd Ndumu Banzi, fresh wat. pan 26.53S-32.16E / 16.2. 1989; E-Y: 2612 shorewashing, Endrödy & Klimaszew” (2 exs. TMSA). – Zaire: “Coll. Mus. Congo, Elisabethville (à la lumière) I-1956/I-1957 Ch. Seydel” (1 ex. MRAC). – Mozambique: “Prov. Sofala 10 km NW Save 6-7.12. 2003 A. Kudrna jr. lgt.” (1 ex. CFP). – Zimbabwe: “Zimbabwe centr. 30 km S Harare 30.11. 1998 leg. F. Kantner” (1 ex. NHMB). – Namibia: “SW Africa Tondoro Okawango 20-23.1. 1975 leg. H. Roer” (1 ex. CGW); “SWA / Namibia Nyangara / Okawango 1-9.4. 1988 leg. H. Roer” (3 exs. CGW). – Botswana: “V.-L. Kal. Exp. Tsotsorogo Pan 17.6.-9.7. 1930 / L. livens Régb. det. L. Gschwendtner” (9 exs. TMSA, 1 ex. MZH).

Diagnosis

Very closely related to L. modestus. Correct determination requires examination of the penis, the apex of which exhibits distinct, species-specific features. In L. cryptos extreme tip of penis apex clearly angled while rounded in L. modestus.

Description

Body length 3.3–3.8 mm, breadth 1.7–2.0 mm. Dorsal, colour pattern slightly variable. Elytra with somewhat obscure, dark ferrugineous irrorations, which at base generally, are almost lacking (Fig. 423).

Head: Pale ferrugineous to ferrugineous; lacks distinct colour pattern. Rather shiny, although finely microsculptured; reticulation double. Large meshes only slightly more strongly developed than small meshes. Large meshes contain 2–5 small meshes. Impunctate, except at eyes; with fine, irregular punctures. Area of punctures extends towards disc-middle, still leaving a considerable impunctate area in middle of head.

Pronotum: Pale ferrugineous to ferrugineous; lacks distinct colour pattern. Rather shiny, although finely microsculptured; reticulation double. Large meshes only slightly more strongly developed than small meshes. Large meshes contain 2–5 small meshes. Smaller meshes, in part, rather indistinct. Impunctate, except frontally and laterally; with fine, irregular punctures.

Elytra: Pale ferrugineous, with somewhat vague, dark ferrugineous irrorations, which often disappear at base of elytra (Fig. 423). Rather shiny, although finely microsculptured. Large meshes only slightly more strongly developed than small meshes. Large meshes contain 2–5 small meshes. Laterally and posteriorly size-categories of microsculpture disappear. Fine, somewhat irregular punctures form a clearly discernible discal row of punctures. Dorsolateral and lateral rows of punctures indistinct; simply indicated by a few, fine and scattered punctures. Pre-apical, lateral row of punctures form a shallow by discernible, pubescent furrow.

Ventral aspect: Pale ferrugineous to ferrugineous; without distinct colour pattern. Rather shiny, although very finely microsculptured; microsculpture in part indistinct. Ventrites with very fine, slightly indistinct, curved striae. Metacoxal plates with a number of rudimentary, transverse furrows. Lateral impression of metacoxal plates moderately deep. Furrows weakly developed, in part indistinct. Impunctate, except apical ventrite, with some fine, scattered punctures and an asymmetric minute knob locate on one side of the ventrite (Fig. 75). Prosternal process slender, posteriorly somewhat extended; apex pointed.

Legs: Pale ferrugineous to ferrugineous. Pro- and mesotarsus slightly enlarged, with distinct suckers.

Male genitalia: Extreme apex of penis tip angled (Fig. 267).

Female: Pro- and mesotarsus slender. Apical ventrite lack small, asymmetric knob (Fig. 76). Body microsculpture variable; sometimes more strongly developed and denser than in male; sometimes as male.

Etymology

The species name cryptos is a Greece noun in apposition and refers to something hidden or secret. The name refers to the identity of the new species, which remained hidden until male genitalia were dissected, being externally similar to L. modestus.

Distribution

Zaire, Mozambique, Zimbabwe, Namibia, Botswana, South Africa (Fig. 547).

Collecting circumstances

Almost unknown. The species has been collected by shore washing and at light collection. Recorded also, from a fresh water pan.

Laccophilus nodieri Régimbart, 1895

Figs 77–78, 268, 424, 547

Laccophilus nodieri Régimbart 1895: 134 (original description, faunistics); Guignot 1946c: 270, 273, 277 (description, faunistics, discussion); Guignot 1959a: 562, 567 (description, faunistics); Bruneau de Miré and Legros 1963: 873, 888 (faunistics); Bilardo and Pederzani 1978: 119: (faunistics, description); Biström 1979: 22 (faunistics); Medler 1980: 155 (faunistics, list.); Nilsson 2001: 247 (catalogue, faunistics); Pederzani and Reintjes 2002: 38, 40 (discussion, faunistics); Reintjes 2004: 68 (faunistics); van Vondel 2005: 131 (faunistics); Nilsson 2015: 215 (catalogue, faunistics).

Type locality

Mali: Badoumbé.

Type material studied

(2 exs.). Lectotype (by present designation) male: “Ht. Sénégal Badoumbé Dr. Nodier I à V 1882 / male symbol /Co-Type / L. nodieri Rég. cotype” (MNHN; habitus in Fig. 424). – Paralectotype: female: “Badoumbé Ht. Senegal / Museum Paris coll. Maurice Régimbart 1908 / nodieri Reg.” (1 ex. MNHN). [Comment: most probably the female paralectotype belongs to L. modestus.]

Additional material studied

(70 exs.). Gambia: “W Div. Abuko Nat. Res. 27.11. 2003 Vondel / L. nodieri det. Rocchi” (1 ex. CSR); “Abuko Nat. Res., at light at the Bamboo pool 18.30-20.30 18.11. 1977, UTM 28PCK2181 / Cederholm et. al leg. / L. nodieri Rég. det. Brancucci 85” (1 ex. MZLU). – Senegal: “Casamance Tabor” (1 ex. MNHN); “Dakar 5. 1939” (2 exs. MNHN). – Mali: “Goundaka, Bandiagara river 14°29'N, 3°56'W, 12.2. 2000 Komarek & Mayer / L. nodieri Rég. det. Wewalka 2001” (1 ex. NMW); “W Bandiagara, pools 14°22'N, 3°41'W, 12.2. 2000 Komarek & Mayer / L. nodieri Rég. det. Wewalka 2001” (1 ex. NMW); “W50 km E Djenne 13°50'N, 4°25'W, 12.2. 2000 Komarek & Mayer / L. nodieri Rég. det. Wewalka 2001” (1 ex. NMW). – Sudan: “Aluakulak 30,5E 6,30N 14.5. 1954 / L. nodieri Reg. det. J. Omer-Cooper” (1 ex. AMGS); “Alel rock pool 30,56 E 6,11N 18.I. 1954 JJOC.” (7 exs. AMGS); “Sandy river 50 mi. NW of Juba 29.1. 1954 JJOC.“ (4 exs. AMGS); “Stream from hot springs Nyangwara 30,51E 4,39N, 29.I. 1954 JJOC.” (2 exs. AMGS); “Sandy river 50 mi. NW of Juba 29.I. 1954 JJOC.” (2 exs. AMGS). – Ivory Coast: “Comoé N8,5° - W3,5°, 22.3. 1999 Reintjes” (6 exs. NMW); same and L. nodieri det. Rocchi” (3 exs. CSR). – Burkina Faso: “Pundu, Mte Volta 1927-28 Dez-Juni Olsufiejev” (3 exs. NHRS). – Ghana: “17 mi S Palbe 1.9. 1971, filtered black light, Gruwell” (1 ex. USNM). – Nigeria: “Stream crossing Kaduna rd. nr Zaria 4.4. 1963 JOC.” (1 ex. AMGS); “Trib. of R. Gagere en route Zaria-Katsina 5.IV. 1963 JOC.” (2 exs. AMGS); “Stream 64 mi. from Bida on Jebba rd.15.IV. 1963 JOC.” (1 ex. AMGS); “Detritus pond 45 mi. from Jos on Bauchi rd.9.IV. 1963 JOC.” (1 ex. AMGS); “R. Niger, bridge Kontagora-Kaduna rd. 3.IV. 1963 JOC.” (1 ex. AMGS); “Kontagora stream 3.IV. 1963 JOC.” (1 ex. AMGS); “Stream nr. Zaria 4.IV. 1963 JOC.” (8 exs. AMGS); “Zaria Pr., Zaria 5-6.3. 1949 Malkin” (1 ex. BMNH, 1 ex. MZH); “NC St. Zaria 2-3.8. 1973 Linnavuori” (1 ex. MZH); “NW St. Yelwa 23.7. 1973 Linnavuori” (1 ex. MZH); “Kano St., Kano-Wudil 17.5. 1973 Linnavuori” (1 ex. MZH); “R. Ogun, Olokomeji nr. Ibadan 24.III. 1963 JOC.” (2 exs. AMGS); “Pond, road Dawria-Kano 6.4. 1963 JOC.” (4 exs. AMGS). – Zaire: “PNG, Utukuru 14/s, 22.7. 1952 De Saeger 3812” (2 exs. NHMB).

Diagnosis

Externally L. nodieri resembles very much of L. flaveolus. Both species have elytral irrorations in part reduced, which makes coverage of it uneven and patchy. Shape of penis, however, is peculiar with a distinct, sharp knob in inner curvature. Corresponding sharp knob lacks in L. flaveolus.

Description

Body length 4.0–4.3 mm, width 2.2–2.4 mm. Elytral colour pattern consists of rather fine, in part unevenly distributed irrorations (Fig. 424).

Head: Pale ferrugineous; no distinct colour pattern. Impunctate, except at eyes; with fine, dense and somewhat irregular punctures. Areas of punctures extend a little towards middle of head. Slightly mat, finely microsculptured. Reticulation almost simple; only indistinct fragments of large meshes discernible. Large meshes, when discerned, contain 3–6 fine meshes.

Pronotum: Pale ferrugineous; lacks colour pattern. Rather shiny, although very finely microsculptured. Reticulation double, but difference between size classes small. Large meshes contain 3–6 fine meshes. At margins with fine, scattered and irregular punctures, which basally in middle are indistinct.

Elytra: Pale ferrugineous, with distinctly reduced, somewhat vague dark ferrugineous irrorations. Irrorations are sparsest basally and in a transverse area posterior to middle (Fig. 424). Slightly mat; finely microsculptured. Reticulation indistinctly double; difference between size classes small. Large meshes, when discernible, contain 3–6 fine meshes. Mesh-organization in part slightly vague. Almost impunctate except for discal row of punctures; consist of fine, irregularly placed fine punctures. Dorsolateral and lateral row of punctures reduced to a few irregular fine punctures.

Ventral aspect: Pale ferrugineous to ferrugineous; no distinct colour pattern exhibited. Almost impunctate. Submat to rather shiny, very finely, in part indistinctly microsculptured. Metacoxal plates in frontal half with some transversely located very shallow furrows. Abdomen basally with fine curved striae. Prosternal process rather slender, apex slightly extended and pointed. Apical ventrite asymmetric; provided with a minute but sharp lateral knob (Fig. 77).

Legs: Pro- and mesotarsus slightly enlarged, somewhat extended and provided with distinct suckers.

Male genitalia: Penis in lateral aspect slightly curved, broad, provided with a sharp, minor process, approximately in middle of the inner side of penis. Extreme apex of penis hooked (Fig. 268).

Female: Externally as male but pro- and mesotarsus slender. Apical ventrite not distinctly asymmetric; lacks lateral knob (Fig. 78).

Distribution

Gambia, Senegal, Mali, Sudan, Ivory Coast, Burkina Faso, Ghana, Nigeria, Zaire (Fig. 547). Personally verified records only accepted in map. Additional country records are Chad (Bruneau de Miré and Legros 1963), Guinea (Reintjes 2004) and Benin (van Vondel 2005a).

Collecting circumstances

Sampled e.g. in pools with loamy bottom and in stagnant remain of brooklet (van Vondel 2005a). Label data give the species to be collected from various pools, at light and in a sandy river.

Laccophilus flaveolus Régimbart, 1906

Figs 79–80, 269, 425, 548

Laccophilus flaveolus Régimbart 1906: 249 (original description, faunistics); Zimmermann 1920a:18 (catalogue, faunistics); Gschwendtner 1930: 88 (faunistics, description, discussion); Guignot 1943: 98 (faunistics, discussion); Guignot 1946c: 265, 268, 272, 273, 312, 315 (discussion, description, faunistics); Guignot 1948: 15 (faunistics); Guignot 1950a: 262, 263 (faunistics); Guignot 1952e:167 (discussion); Legros 1954: 268 (discussion); Guignot 1955e: 2 (discussion); Guignot 1956b: 220 (discussion); Omer-Cooper 1958b: 37, 48, 49 (discussion, description, faunistics, biology); Guignot 1959a: 557, 561, 562 (description, faunistics); Guignot 1959d: 161 (discussion, faunistics); Omer-Cooper 1965: 77, 87 (description, faunistics); Legros 1972: 466 (faunistics); Bameul 1984: 94 (faunistics); Pederzani 1988: 107 (faunistics, biology); Rocchi 1991: 86 (faunistics, list); Nilsson 2001: 243 (catalogue, faunistics); Pederzani and Reintjes 2002: 38, 40 (discussion, faunistics); Nilsson 2015: 211 (catalogue, faunistics).

Laccophilus pampinatus Guignot 1941: 35 (original description, faunistics); Guignot 1946c: 269, 273, 274, 276, 313 (description, faunistics, discussion); Guignot 1959a: 558, 562, 564, 565, 566, 568 (redescription, discussion, faunistics); Nilsson 2001: 248 (catalogue, faunistics); Pederzani and Reintjes 2002: 38, 40 (discussion, faunistics); Nilsson 2015: 215 (catalogue, faunistics). New synonym.

Comment on identity

Confusion in determination of species has been a common problem. Accordingly, records and data from old literature must be considered carefully.

Type localities

Laccophilus flaveolus: Kenya: Baie de Kavirondo.

Laccophilus pampinatus: Uganda: Central Uganda.

Type material studied

(37 exs.). Laccophilus flaveolus: Lectotype (by present designation) male: Kenya “Lac Victoria Nyanza Baie de Kavirondo IX-X. 1903 / male symbol / Cotype” (MNHN). – Paralectotypes: Same data as lectotype (2 exs. MNHN; habitus in Fig. 425); “Cotype / Victoria Nyanza Kavirondo-Bay / Brit. Mus.1905-199 / Laccophilus flaveolus Rég. sp. n. type” (1 ex. BMNH); “Cotype / Lac Victoria Nyanza Baie de Kavirondo IX-X-1903” (1 ex. IRSNB); L. flaveolus Baie de Kavirondo Lac Victoria Nyanza Alluaud IX-X. 1903” (4 exs. IRSNB); same data and on same pin (1 ex. IRSNB; belongs to L. pallescens Régimbart); Afr. Orle Anglaise Baie de Kavirondo (Victoria Nyanza N-E.) Ch. Alluaud IX-X-1903 / Museum Paris coll. Maurice Régimbart 1908 / Laccophilus flaveolus Rég. sp. n. type / Type” (20 exs. MNHN); “Baie de Kavirondo (Alluaud) / Museum Paris coll. Maurice Régimbart 1908 / flaveolus Rég.” (3 exs. MNHN).

Laccophilus pampinatus: Holotype: male: Uganda, “Ouganda Central Alluaud I-II. 1909 / male symbol / Type / Det. Dr. Guignot Laccophilus pampinatus Guign. Type” (MNHN). – Paratypes: Uganda, Same as holotype, but labelled as “Paratype” (1 ex. MNHN). – Kenya, “Lac Victoria Baie de Kavirondo Alluaud IX. 1903 / male symbol / Laccophilus pampinatus Guignot Paratype / Paratype” (1 ex. MNHN); “Afr. Or. Angl. (Lac Victoria) Baie de kavirondo Alluaud & Jeannel Déc. 1911 – 1112 - St. 22 et 23 / female symbol / Laccophilus pampinatus Guignot Allotype female / Allotype” (1 ex. MNHN).

Additional material studied

(155 exs.). Sudan: “Kawrajena 20.3. 1947” (2 exs. ZMUC). – Zaire: “PNA 23.8. 1957 Vanschuytbroeck / Secteur Nord, rive dr. Semliki, rte Muramba, 905 m” (6 exs. MRAC, 1 ex. MZH); “PNA 26.8. 1957 Vanschuytbroeck / Secteur Nord, marais Buyansha sur r. dr. Semliki, 905 m” (6 exs. MRAC); “PNA 27.8. 1957 Vanschuytbroeck / Secteur Nord, rive Ihunga, af dr. Semliki 1300 m” (2 exs. MRAC); “PNG Ndelele K. 117/14s 19.3. 1952 H. De Saeger” (2 exs. MRAC); “PNG Ndelele/14s 1.8. 1952 H. De Saeger” (2 exs. MRAC); “PNG PpK.14/g/14s, 4.4. 1952 H. De Saeger 3290” (4 exs. MRAC, 1 ex. MZH); PNG II/gd/14, 30.7. 1952 H. De Saeger” (1 ex. MRAC); “PNG II/gd/4, 29.5.1952 De Saeger” (1 ex. IRSNB). – Uganda: “Uganda Central Alluaud I-II. 1909 / Cotype” (1 ex. MNHN; not type material); “Butiaba Flats 2.9. 1967 Brown” (1 ex. BMNH). – Kenya: “Dam at Kaloleni Mission, Kilifi Distr. 15.9. 1976 Holmen” (1 ex. ZMUC). – Tanzania: “Ukerewe I. Father Conrad” (5 exs. BMNH, 1 ex. MZH); “Tanganyika Ukerewe / L. pampinatus Guignot det. Wewalka 1979” (7 exs. OLML); “Mwanza nr Lake Victoria 1957 / Marginal pools and ditches” (2 exs. BMNH, 1 ex. MZH); “Mwanza nr Lake Victoria / Sweet potato channels” (1 ex. BMNH); “Stream, Mbeya-Tunduma rd., 18.10. 1948 JOC” (1 ex. AMGS); “Foothills of Kilimandjaro 14.2. 1954” (2 exs. AMGS); “Zanzibar Pemba, Sept, 1955 Fowler” (2 exs. AMGS). – Zambia: “S Luangwa NP, Mfuwe Crocodile Farm 23.3. 1993 13°06'03"S-31°47'32"E, 450 m, lux Uhlig leg.” (1 ex. ZMHB); “Lusaka 5.11. 1973 Lange” (1 ex. CGC). – Malawi: “River nr Portuguese border nr Mwanza 9.2. 1948” (1 ex. AMGS); “Zomba plateau, stream 6000 ft 7.3. 1948” (2 exs. AMGS). – Mozambique: “Magude16.8. 1915 C.J. Sw. / L. adspersus Boh. det. Gschwendtner” (8 exs. TMSA). – Zimbabwe: “Small stream Halfway Hotel Salisbury-Gatooma 14.9. 1948” (1 ex. AMGS); “Salisbury 14.9. 1948 J. Omer-Cooper” (2 exs. NHMB); “Wankie Game Res. Masumu Dam Sept. 1948 JOC.” (4 exs. AMGS); “Wankie Game Res. Waterhole Sept. 1948 JOC.” (1 ex. AMGS); “5 mi SE Wankie 7.4. 1968 Spangler” (25 exs. USNM, 5 exs. MZH); “Pool Lundi 22. N. 1948 JOC.” (20 exs. AMGS); “Sinkukwe 30.12. 1948 J.O.C.” (6 exs. AMGS). – Botswana: “Tsotsorogo Pan 17.6.-9.7. 1930 V.-L. Kal. Exp. / L. adspersus Boh. Gschwendtner det.” (9 exs. TMSA, 1 ex. MZH); same data but L. pampinatus Guignot det. Wewalka 1979” (1 ex. OLML); “Kabulabula Chobe River 11-24.7. 1930 V.-L. Kal. Exp. / L. adspersus Boh. Gschwendtner det.” (1 ex. TMSA); “N’Kate Makarikari 6-23.8. 1930 V.-L. Kal. Exp. / L. adspersus Boh. Gschwendtner det.” (2 exs. TMSA); “Metsimaklaba 7-12.3. 1930 V.-L. Kal. Exp. / L. adspersus Boh. Gschwendtner det.” (3 exs. TMSA). – South Africa: “Kruger Nat. Pk, Skukuza res. camp, 25.00S-31.35E/19.2.1995 UV-light & trap E-Y: 3102, Endrödy-Younga leg.” (1 ex. TMSA); “Kruger Nat. Pk, Skukuza 12 km, 25.04S-31.37E / 6.3. 1996 UV-light E-Y: 3217, Endrödy-Younga leg.” (1 ex. TMSA, 1 ex. MZH); “Trsvl Kruger National Pk, Leeu Pan NE Skukuza 1.5. 1951 / Brinck-Rudebeck / L. flaveolus Régb. det. J. Omer-Cooper” (1 ex. MZLU); “Mpumalanga, 7 km upstream from Skukuza, Sabie R. N-24.970, E31.540, 25.10. 1990 de Moor” (1 ex. AMGS); “Kruger Nat. Pk, Levuvu River 22.27S-31.10E / 12.2. 1994 E-Y: 2998 shorewashing” (1 ex. TMSA); “Trsvl., Naboomspruit Torino Ranche 24.37S-28.38E / 15.1. 1989 E-Y: 2774, UV light, vlei edge Endrödy-Younga leg.” (1 ex. TMSA). – Madagascar: “Zombitse Ankilemiletsy, muddy waterhole N- 22.868, E 44.576, 544 m 14.5. 2006 Bergsten/BMNH(E) <794187> DNA voucher / L. flaveolus Régb. det. Bergsten” (1 ex. NHRS); same data, but “<794193> DNA voucher” (1 ex. NHRS).

Comments on synonymy

The lectotype of L. flaveolus and the holotype of L. pampinatus have been examined and compared. No morphological features, which would justify separation of two species were detected. Accordingly, they are synonyms and L. flaveolus being the older name is the valid name for the species.

Diagnosis

Laccophilus flaveolus is separated from resembling species by study of elytral colour pattern in combination with peculiarly shaped penis. Externally it resembles most of L. nodieri but there is clear difference in shape of penis: Penis of L. flaveolus lacks sharp knob on inner outline. Additionally, penis long, medially somewhat bent and extreme apex hooked with extreme tip sharp (Fig. 269).

Description

Body length 4.2–4.6 mm, width 2.3–2.4 mm. Pale coloured except elytra. Elytral colour pattern is formed by quite extensive irrorations; irrorations somewhat sparse and in part reduced (Fig. 425).

Head: Pale ferrugineous. Rather shiny to submat, reticulation double but difference between size-categories small. Large meshes contain 2–10 finer meshes when they are discernible. Almost impunctate; at eyes with fine and irregularly distributed punctures. Area of punctures at each eye extends towards middle but areas are not connected.

Pronotum: Pale ferrugineous, base in middle often with narrow darkened area. Rather shiny although finely microsculptured; reticulation double. Large meshes contain 2–10 meshes; sometimes fine meshes indistinct and hardly visible. At margins with very fine, scattered punctures; pronotum discally impunctate.

Elytra: Pale ferrugineous, extensively with dark brown to dark ferrugineous, rarely slightly variable irrorations (Fig. 425). Same groundplan of irrorations discernible, although slight reduction sometimes present. Rather shiny to submat, finely microsculptured; reticulation double. Large meshes only slightly coarser than fine meshes. Large meshes contain between 2–7 fine meshes. Posteriorly fine and large meshes are mixed so that separate size categories are not discernible. Impunctate, except for three irregular longitudinal rows of very fine and scattered punctures, located discally, dorso-laterally and laterally. Lateral, pre-apical furrow rather shallow, distinctly pubescent.

Ventral aspect: Pale ferrugineous. Slightly mat, very finely microsculptured. Metacoxal plates with some very indistinct transverse furrows. Ventrites basally sparsely striated; striae curved. Almost impunctate. Apex of prosternal process slender and pointed. Apical ventrite with a minute but distinct asymmetric knob. A minor knob can also be detected on the other side (Fig. 79).

Legs: Pale ferrugineous. Pro- and mesotarsus slightly enlarged and extended, with suckers.

Male genitalia: Penis long, medially somewhat bent and extreme apex hooked with extreme tip sharp (Fig. 269).

Female: Apical ventrite as in Fig. 80. Pro- and mesotarsus slender. Epipleuron slightly enlarged posterior to middle and external edge can sometimes be detected when specimen is studied from above.

Distribution

Sudan, Zaire, Uganda, Kenya, Tanzania, Zambia, Malawi, Mozambique, Zimbabwe, Botswana, South Africa and Madagascar (Fig. 548). Confusion in the determination of species frequent. Accordingly only verified records are mapped. Additional country records, not verified, are Ivory Coast and Mali (Omer-Cooper 1965) and Senegal (Legros 1972).

Collecting circumstances

Insufficiently known. Omer-Cooper (1958b) reports the species from various minor water bodies, as streams and ponds. Label data give as collection technique UV light collection, light trap and shorewashing. Detailed information is not available.

Laccophilus remex Guignot, 1952

Possibly a species complex Figs 81–82, 270–274, 426–429, 549

Laccophilus remex Guignot 1952e: 167 (description, faunistics); Guignot 1953e: 4 (discussion); Guignot 1954: 25 (discussion); Guignot 1956b: 219 (faunistics); Omer-Cooper 1958b: 37, 47 (discussion, description, faunistics); Guignot 1958: 7 (discussion); Guignot 1959a: 535, 557, 560, 562 (description, discussion, faunistics); Omer-Cooper 1965: 77, 85 (description, faunistics); Medler 1980: 155 (faunistics, list); Pederzani and Rocchi 1982: 72 (faunistics); Nilsson and Persson 1993: 80, 94 (faunistics); Nilsson, Persson and Cuppen 1995: 505 (faunistics); Bilardo and Rocchi 1999: 232, 234 (faunistics); Nilsson 2001: 250 (catalogue, faunistics); Bilardo and Rocchi 2002: 174 (list, faunistics); Pederzani and Reintjes 2002: 38, 40 (discussion, faunistics); Bilardo and Rocchi 200a: 291 (faunistics); Bilardo and Rocchi 2006a: 130 (faunistics); Nilsson 2015: 217 (catalogue, faunistics).

Laccophilus concisus Guignot 1953e: 4 (original description, faunistics); Guignot 1954: 24 (description, faunistics); Guignot 1958: 7 (discussion); Guignot 1959a: 558, 562, 563 (description, faunistics); Guignot 1961b: 238 (faunistics); Omer-Cooper 1962: 295 (faunistics); Omer-Cooper 1965: 77, 86 (description, faunistics); Omer-Cooper 1967: 60 (discussion, synonym L. praeteritus O-C. = L. concisus Guig.); Bilardo 1982: 447 (description, faunistics; given as Laccophylus concisus); Bilardo and Rocchi 1987: 104 (faunistics, biology); Nilsson et al. 1995: 505 (faunistics); Rocchi 2000: 23 (faunistics); Nilsson 2001: 242 (catalogue, faunistics); Pederzani and Reintjes 2002: 38 (faunistics); Nilsson 2015: 210 (catalogue, faunistics). New synonym.

Laccophilus turneri Omer-Cooper 1956: 21 (no description, faunistics, biology); Omer-Cooper 1957: 17, 90 (original description, faunistics); Omer-Cooper 1958b: 47 (discussion); Omer-Cooper 1965: 86 (synonym L. remex Guignot ); Nilsson and Persson 1993: 80 (list, synonymy); Nilsson et al. 1995: 505 (list, synonymy); Nilsson 2001: 250 (catalogue, faunistics, list, synonymy); Pederzani and Reintjes 2002: 40 (faunistics); Nilsson 2015: 217 (catalogue, list, synonymy). Confirmed synonym.

Laccophilus praeteritus Omer-Cooper 1957: 18, 90 (original description, faunistics); Omer-Cooper 1958b: 37, 47, 48, 49 (discussion, description, faunistics, biology); Omer-Cooper 1965: 86 (list synonymy, L. concisus Guignot); 1967: 60 (discussion, synonymy); Nilsson 2001: 242 (catalogue, faunistics, list, synonymy); Pederzani and Reintjes 2002: 40 (faunistics, list, synonymy); Nilsson 2015: 210 (catalogue, list, synonymy). New synonym of L. remex.

Type localities

Laccophilus remex: Ivory Coast: Duékoué.

Laccophilus concisus: Zaire: PNU, Lusinga.

Laccophilus turneri: South Africa: Boekenhout, Nylstroom (River Nyl at Num Num).

Laccophilus praeteritus: South Africa: Transvaal, Ermelo.

Type material studied

(26 exs.) Laccophilus remex. Holotype: male: “Cote d’Ivoire Duékoué / Museum Paris 12-1930-IV-1931 Ch. Alluaud et P.A. Chappuis / Det. Dr. Guignot Laccophilus tschoffeni Rég. / Type / F. Guignot det. 1952 Laccophilus remex Guign. Type, male symbol” (MNHN). – Paratypes: females: “Cote d’Ivoire Duékoué / female symbol / Museum Paris 12-1930-IV-1931 Ch. Alluaud et P.A. Chappuis / Paratype” (1 ex. MNHN); same data but labelled “Allotype” (1 ex. MNHN).

Laccophilus concisus: Holotype: male: “Holotypus / Congo Belge: PNU, Lusinga (Galerié) 22-25-V-1945 G. F de Witte: 29 / Coll. Mus. Congo (ex. coll. I.P.N.C.B.) / Laccophilus concisus Guign. sp. n. Type male / F. Guignot det., 1952 Laccophilus concisus sp. n. Type male” (MRAC; habitus in Fig. 427). – Paratypes: “Congo Belge: PNU, Lusinga (Galerié) 22-25-V-1945 G. F de Witte: 29 / Paratype / F. Guignot det., 1952 Laccophilus concisus sp. n. / R.I.Sc.N.B. I.G. 24.054” (1 ex. IRSNB); same but “7-20-VI-1945” and “191” (2 exs. IRSNB); same, but “Kabwekanono (1.815 m) 18-III-1947” and “64a” (1 ex. IRSNB).

Laccophilus praeteritus: Holotype: male: “Male / Type / Transvaal Ermelo / Dec. 1948 J. Omer-Cooper / L. praeteritus Omer-Cooper/Brit. Mus. 1957-660 / Laccophilus concisus Guign. J. Balfour-Browne det. 1960” (BMNH). – Paratypes: “Female / Type / Transvaal Belfast 30.XI. 1948 J.O.C. / Female allotype / Brit. Mus. 1957-660 / L. praeteritus (1 ex. BMNH); “Paratype / Transvaal pond W. Belfast 23.11. 1948 JOC. / L. praeteritus O-C. det. J. Omer-Cooper” (1 ex. AMGS); “Paratype / Duivels Kloof Merenskydam 24.11. 1948 Omer-Cooper” (1 ex. AMGS); “Transvaal sluggish stream / nr Ermelo 1.12. 1948 J.O.C. / L. praeteritus O-C. / Paratype” (1 ex. TMSA).

Laccophilus turneri: Holotype: male: “Type / Transvaal Nylstroom R. Nyl at Num Num 23. Aug. 1948 J. Omer-Cooper / Type, male / L. turneri O-C.” (BMNH). – Paratypes: female: Principally with same data as holotype, but labelled as “female allotype” (1 ex. BMNH); Almost labelled as holotype but “paratype” (6 exs. AMGS, 1 ex. TMSA); “Paratype / Transvaal Waterberg distr. Deel Kraal 20.8. 1948 JOC.” (1 ex. AMGS); “Paratype / Transvaal Nylstroom 19 Aug. 1948 JOC. / L. adspersus Boh. J. Balfour-Browne det.” (1 ex. AMGS); “Paratype / Transvaal Duivels Kloof 24. N. 1948 JOC.” (1 ex. AMGS); “Paratype” (1 ex. AMGS; status uncertain, because locality information absent).

Additional material, studied (278 exs.): Sudan: “Dahr el Ghazal M’Boloko 23.2. 1963 Linnavuori” (1ex. MZH); “Equatoria, Nzara 22.4. 1986 Wewalka” (1 ex. CGW); “O. Sudan Adjuba I.U. Neumann” (1 ex. ZMHB). – Sierra Leone: “Musaia 16.1. 1946 / Hippo mud pan” (1 ex, BMNH). – Liberia: “Suakoko 19.12, 1951/6-9 pm light trap Blickenstaff” (9 exs. USNM, 2 exs. MZH); “Suakoko 11.12. 1951 Blickenstaff” (1 ex. USNM, 1 ex. MZH); “Suakoko 1.1. 1952 Blickenstaff” (6 exs. USNM). – Ivory Coast: “Divo 28.11. 1963 Decelle” (2 exs. MRAC). – Ghana: “Ashanti Kumasi 330 m, N6.43-W 1.36 / 15.9. 1967 at light Endrödy-Younga” (1 ex, TMSA); “Kumasi 3.6. 67 Endrödy-Younga / L. remex Guignot det. Wewalka 76” (1 ex. MHNG); same data but “6.7. 67” (1 ex. MHNG); same data but “16.7. 67” (3 exs. MHNG); same data but “24.6. 67” (2 exs. MHNG). – Nigeria: “Stream 64 mi. from Bida on Jebba rd. 15.IV. 1963 JOC.” (3 exs. AMGS); “EC St. Norcap nr Abakaliki 29.6. 1973” (1 ex. MZH); same and “ad lucem / Linnavuori” (1 ex. MZH). – Gabon: “Makoukou Riv. Oua (Ivindo) 16-19.1. 2001 Bilardo / L. remex det. Rocchi 02” (2 exs. CSR); “Belinga 5.2.-4.4. 1963 Coiffat” (62 exs. NHMB). – Congo: “Parc Nat. d’Odzala Mboko-Lango 21.8. 2002 Bilardo” (1 ex. CSR); “Odzala NP, 400 m, 10.2. 1997 Murzin leg.” (4 exs. NMPC). – Zaire: “PNG Miss. H. De Saeger II/fd/14, 18.6. 1951, 1946” (1 ex. NHMB); “PNG Miss. H. De Saeger II/hd/17, 13.10. 1951, 2595” (1 ex. NHMB); “PNG Miss. H. De Saeger II/fd/14s, 3.4. 1952, 3278” (9 exs. MRAC, 2 exs. MZH); “PNG Miss. H. De Saeger II/fd/12, 10.3. 1952, 3180” (13 exs. MRAC, 1 ex. MZH); “PNG Miss. H. De Saeger Pali/8, 22.3. 1952, 3217” (1 ex. MRAC); “PNG Miss. H. De Saeger II/fd/13, 5.5. 1952, 3421” (1 ex. MRAC); “PNG Miss. H. De Saeger II/fc/14, 17.7. 1952, 3806” (1 ex. MRAC); “PNG Miss. H. De Saeger I/a/M 7.6. 1950 Rec. G. Demoulin 584 / L. remex Guignot det. Guignot 1957” (1 ex. AMGS); “PNG Miss. H. De Saeger II/fc/14, 4.7. 1952, 3736 / Paratype of L. saegeri Guignot” (1 ex. IRSNB); “PNU Mukana, 1810 m, 24.3. 1947 / Dr. F. Guignot det., 1953 Laccophilus concisus Guign.”(2 exs. IRSNB, 2 exs. MRAC; labelled as paratypes but not mentioned in original description); “PNU Kabwekanono p.t.s. Lufwa affl. dr. Lufira (1.815 m) 12.1. 1948, 1199a” (1 ex. MRAC; labelled as paratype but not mentioned in original description); “PNU Kaswabilenga (700 m) 17.10. 1947, 845a” (1 ex. MAC; labelled as paratype but not mentioned in original description); “PNU Mubale – 1480 m, 10-13.5. 1947, 352a” (2 exs. MRAC; labelled as paratypes but not mentioned in original description); “de Luebo à Luluabourg N. 1921 Ghesquière / L. remex Guignot det. Wewalka 1979” (1 ex. OLML). – Tanzania: “Kondoa 300 m 10. 1938” (1 ex. MNHN; habitus in Fig. 426); “Mlowa R. Tunduma-Mbeya rd. 16.10. 1948” (1 ex. AMGS); “Rukwa 26.12. 1961 C. Carnegie” (3 exs. AMGS); “T.T. Rukwa Tumba 12.1. 1991 T. river Backlund / L. remex Guignot det. Nilsson -96” (1 ex. MZLU); “Ukerewe VIII., 2 / 1294 B / L. remex Guign. det. Wewalka 79” (4 exs. OLML); “Zansibar Küste Hildebr.” (1 ex. ZMHB). – Zambia: “Katambora 40 ml W Victoria Falls, April, 1962” (1 ex. BMNH); “Kapiri Mpushi env. 13.12. 2002 Kantner” (2 exs. NHMB). – Malawi: “Nyasal. Bua R. 2.10. 1948 JOC.” (2 exs. AMGS). – Mozambique: “Mandambuzi, Manda Wilderness Res. S12°17.679’, E34°46.260’, Watson 16.2. 2008” (2 exs. CGF); “Umbeluzi R. Dec. 4. 1948 JOC.” (2 exs. AMGS). – Namibia: “E Caprivi 10 km SE Katima Mulilo 17°31'S/24°25'E, Zambesi Altwasserarm, lux 6.3. 1992 Uhlig” (5 exs. ZMHB); “Kavango Popa Falls, lux 18°07'S/21°35'E, 26.2.-3.3. 1992 Uhlig leg.” (1 ex. ZMHB). – Botswana: “Kabulabula Chobe River 11-24.7. 1930” (1 ex. OLML); “Maun Thamalakane R. 10.10. 1982 Bilardo / L. concisus G. det. Bilardo” (2 exs. MSNM). – Zimbabwe: “Kyle Recr. Pk at Lake Mutirikwi 1-5.12. 1993, lux Uhlig” (1 ex. ZMHB); “Stream Rusapi 13.11. 1948” (3 exs. AMGS); “Pool Lundi 22. N. 1948 JOC.” (5 exs. AMGS); “Stream between Salisbury-Bromley 12.11. 1948” (6 exs. AMGS); “Small stream Halfway Hotel-Gatooma, Salisbury 14.9. 1948 / Laccophilus turneri O-C. det. J. Omer-Cooper” (1 ex. AMGS); “Pool Lundi 22. N. 1948 JOC.” (11 exs. AMGS); “Kariba env. 20.12. 1998 Kantner” (1 ex. NHMB). – South Africa: “Transvaal Ermelo stream 7. Dec. 1948 JOC.” (1 ex. AMGS); “Transvaal Ermelo Dec. 1946” (1 ex. AMGS); “Kruger Nat. Pk. Skukuza Res. camp 24.59S-31.35E/3.3. 1996 UV-light Endrödy-Younga” (1 ex. TMSA); “Kruger Nat. Pk. Skukuza Res. camp 25.00S-31.35E/19.2. 1995 UV-light & trap Endrödy-Younga” (1 ex. TMSA); “Trsvl Bundu Inn 25.28S-28.55E/24.3. 1974 at merc. vap. light, Endrödy-Younga” (2 exs. TMSA); “Natal Lions R. 30.3. 1960” (2 exs. AMS); “Natal Zululand Hlui-Hlui 15.9. 1947” (1 ex. AMGS); “Zululd Ndumu Nyamithi, saline pan 26.54S-32.16E/12.6. 1989 shorewashing Endrödy-Klimaszew” (1 ex. TMSA); “Kwazulu-Natal Ndumu 25.55S-32.18E / 21.11. 2002 light trap Harrison & Müller” (1 ex. TMSA; habitus in Fig. 428); “Zululd Ndumu Banzi, fresh wat. pan 26.53S-32.16E / 16.2. 1989 shorewashing Endrödy & Klimaszew” (7 exs. TMSA, 1 ex. MZH); “Blinkwater Reserve, Greytown Natal, first stream from entrance, 1100 m mist belt grassland 4.2. 1997 Turner” (2 exs. AMGS, 1 ex. CCT); “Kwazulu Natal” (1 ex. AMGS); “Gen. 766G 3.7. 1962 (= Insese River near Empangeni station 6, Tugela River)” (1 ex. AMGS); “W. C. Prov., pond on Plettenberg rd, Knysna Distr. 14.2. 1947 JOC.” (2 exs. AMGS); “WCP, Knysna 34°04.28'S, 23°04.11'E 9 m Hotový & Mateju leg.” (2 exs. NMPC); “E. C. Prov. Uitenhage 5.12. 1954 JOC.” (1 ex. AMGS); “E. C. Prov. Mount Currie 6.5. 1956 JOC.” (6 exs. AMGS); “E. C. Prov. Mount Currie 13.11. 1957 JOC.” (6 exs. AMGS); “E. C. Prov. Hogsback 2. 1942 O.C.” (3 exs. AMGS); “E. C. Prov. E. London pond nr. Mooiplaats 8.3. 1955” (1 ex. AMGS); “E. C. Prov. Maclear 9.5. 1956 JOC.” (10 exs. AMGS); “E. C. Prov. nr. Zwatrberg 1960 Chutter” (3 exs. AMGS); “E. C. Prov. Krom R. 1960” (2 exs. AMGS); “E. C. Prov. Albany stream Storm May 1939” (3 exs. AMGS); “E. C. Prov. Albany Distr. Grahamstown 10.3. 1946 JOC.” (1 ex. AMGS); “E. C. Pr. Albany distr. Grahamstown Teafontein stream (33 26 AA) 3.V8. 1939 JOC.” (1 ex. AMGS); “CPr. Knysna Main Forest Buffelsnek 18.I. 1951 / Brinck-Rudebeck / L. concisus Guig. = L. praeteritus O-C. det. JOC” (3 exs. MZLU; habitus in Fig. 429); “CP., Pond NO Knysna on Hwy R340 8.3. 1997 Challet” (4 exs. CGC, 1 ex. MZH); “Cape, Gatberg R, vlei, N-31.250, E28.120, 26.3. 1993 De Moor & al.” (1 ex. AMGS).

Specimen with uncertain determination

South Africa: “Blinkwater Reserve, Greytown Natal, bog on summit, 1100 m nist belt grassland 4.2. 1997 Turner” (1 ex. CCT).

Comments on synonymy and species status

Male holotypes of L. remex, L. concisus, L. turneri and L. praeteritus have been examined and compared. Shape of penis is almost identical in the four taxa. Considering all examined male genitalia there is variation in shape between extremes, which makes a separation of different species difficult or almost impossible. Colour pattern of elytra is quite variable but exhibit similar ground-plan with transitional morphs between extremes. Accordingly, the four taxa are for the time being regarded, conspecific and synonymies are introduced as follows: Earlier synonymy of L. remex and L. turneri is confirmed as well as earlier synonymy of L. concisus and L. praeteritus. Laccophilus concisus is a new synonymy of L. remex. Laccophilus remex, being the oldest available name is the valid name of the species. An alternative, plausible, interpretation is that L. remex is in fact a complex of very closely related species. With present knowledge, the delimitation of the different species remains, however, an open question. Further study is definitely needed.

Diagnosis

Although L. remex, as delimited here, exhibits considerable variation in appearance of elytral colour pattern this feature can often be used for recognition of the species. The species is also characterized by the robust penis, which often exhibits some variation. Laccophilus remex resembles very much of L. turbatus but L. remex is almost always larger (body length exceeds in most cases 4 mm). Additionally external outline of penis is rounded when it is angled in L. turbatus.

Description

Body length 4.0–4.7 mm; width 2.2–2.6 mm. Elytral colour pattern variable; elytra covered with dense irrorations, which basally often are to a variable degree reduced. Basally often with quite extensive pale areas where irrorations are absent. Additionally, elytra sometimes with irrorations being sparser posterior to middle (Figs 426–429).

Head: Pale ferrugineous. Posteriorly sometimes with dark area. Submat, finely microsculptured. Reticulation double; large meshes contain 2–7 smaller meshes. At eyes with fine and irregular punctures, which extend towards middle of head.

Pronotum: Pale ferrugineous. At frontal margin and medially at posterior margin with a distinct dark to blackish ferrugineous area. Basal dark area sometimes reduced. Frontal, dark marking sometimes totally absent. Submat, finely microsculptured; reticulation double: larger meshes contain 2–7 smaller meshes. Frontally and laterally with scattered, fine punctures.

Elytra: Pale ferrugineous, with dense, dark ferrugineous to blackish irrorations. At base, irrorations often sparser, in part reduced, sometimes forming variable pale areas lacking irroration (Figs 426–429). Submat, finely microsculptured; reticulation double: large meshes contain 2–6 smaller meshes. Posteriorly two kinds of microsculpture become mixed and difficult to separate. Laterally and posteriorly double reticulation becomes obscure and indistinct. Elytron with discal, dorsolateral and lateral rows of fine and irregular punctures. Pre-apical, lateral furrow with fine hairs.

Ventral aspect: Pale ferrugineous to ferrugineous. Almost impunctate. Rather shiny, with very fine microsculpture. Metacoxal plates with a few, rather vague, transverse furrows. Ventrites with fine, somewhat curved striae. Apical ventrite of male with fine knob on one side (Fig. 81). Apex of prosternal process narrow, pointed.

Legs: Pro- and mesotarsus slightly enlarged and extended, with suckers.

Male genitalia: Penis in lateral aspect quite broad, medially distinctly bended and extreme apex hooked, outline of it, however, rounded (Figs 270–274).

Female: Pro- and mesotarsus rather slender. Apical ventrite symmetric (Fig. 82).

Distribution

Sudan, Sierra Leone, Liberia, Ivory Coast, Ghana, Nigeria, Gabon, Congo, Zaire, Tanzania, Malawi, Zambia, Namibia, Botswana, Zimbabwe, South Africa and Swaziland (Fig. 549). Unverified country records are Senegal and Guinea (Guignot 1956b), Mozambique (Omer-Cooper 1965), Ethiopia (Nilsson and Persson 1993) and Guinea Bissau (Nilsson et al. 1995). Additionally, under the name L. concisus, Bilardo (1982) gives Cameroon.

Collecting circumstances

Information in literature is superficial and sparse. Omer-Cooper (1958b) reports the species from a river with clear water and with water lilies, reed beds and patches of swamp. Also collected from pools e.g. in a tributary. Label data indicate that the species is often caught with light traps or at light. Moreover the species has been collected in various kinds of pools and ponds but also from running water as streams.

Laccophilus turbatus Guignot, 1958

Figs 83–84, 275, 430–431, 550

Laccophilus turbatus Guignot 1958: 8 (original description, faunistics); Omer-Cooper 1965: 86 (list, synonymy L. concisus Guignot?); Nilsson 2001: 242 (catalogue, faunistics, list, synonymy, L. concisus Guignot); Pederzani and Reintjes 2002: 40 (faunistics, list, synonymy, L. concisus Guign.); Nilsson 2015: 210 (catalogue, faunistics, list, synonymy, L. concisus Guignot. Restored species.

Type locality

Zaire: Parc National Garamba.

Type material studied

(8 exs.): Holotype, male: “Holotypus / Congo belge P.N.G. Miss. H. De Saeger II/hd/17, 13-X-1951 Réc. H. De Saeger. 2595 / Guignot det., 1957Laccophilus turbatus sp. n. Holotype” (MRAC). – Paratypes: “Congo Belge P.N.G. Miss. H. De Saeger/13-X-1951 Réc. H. De Saeger. 2595 / female / Paratype” (1 ex. IRSNB, 4 exs. MNHN); same data but “II/fc/14, 17.7. 1952, 3608” (1 ex. NHMB); same data but “II/gd/14s, 25.8. 1952, 3984” (1 ex. NHMB).

Additional material studied

(50 exs.): Sudan: “S. Sudan stream from hot springs, Nyangwara 30,5E, 4,39N, 29.1. 1954 JJOC.” (2 exs. AMGS). – Liberia: “Suakoko 18.3. 1952 / Blickenstaff Light trap” (1 ex. USNM, 1 ex. MZH); “Suakoko 20.2. 1952 Blickenstaff” (1 ex. USNM). – Nigeria: “R. Kaduna 4.5 mi. from Jos 13.4.1963 JOC.” (4 exs. AMGS);“R. Kaduna rd. 13.5 mi. from Jos 13.IV. 1963 JOC.” (1 ex. AMGS); “Zaria 1969 Brancucci” (1 ex, CSR); “stream Kaduna-Zaria rd. 4.IV. 1963 JOC.” (1 ex. AMGS); “river, rd. to Enugo about 79 mi. from Makurdi 24.4. 1963” (1 ex. AMGS); “stream 64 mi. from Bida on Jebba rd.12.4. 1963 JOC.” (2 exs. AMGS); “Kontagora pools in dry stream 3.IV. 1963 JOC.” (2 exs. AMGS). – Zaire: Same data as paratype, but not labelled as belonging to type material (1 ex. IRSNB); “PNG Miss. H. De Saeger II/fd/12, 10.3. 1952, 3180” (2 exs. MRAC, 1 ex. MZH); same but “II/fd/13, 5.5. 1952, 3421” (3 exs. MRAC); same but “II/fd/14s, 3.4. 1