Laccophilus modestus Regimbart , 1895

Bistroem, Olof, Nilsson, Anders N. & Bergsten, Johannes, 2015, Taxonomic revision of Afrotropical Laccophilus Leach, 1815 (Coleoptera, Dytiscidae), ZooKeys 542, pp. 1-379 : 98-102

Taxon classification Animalia Coleoptera Dytiscidae

Laccophilus modestus Regimbart, 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 Laccophilus 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 / Laccophilus 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 Reg. 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 / Laccophilus 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 / Laccophilus 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 / Laccophilus 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 / Laccophilus 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 / Laccophilus modestus Régb. det. Wewalka 2005" (7 exs. CGW). - Burkina Faso (= Ht Volta): "Ouagadougou X. 1926" (1 ex. NHMB); "Haute Volta Bobo Dioulasso / Mu seum Paris 12 - 1930 - IV 1931 Ch. Alluaud & P. Chappuis / male symbol / Laccophilus 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 / Laccophilus 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 / Laccophilus 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 / Laccophilus 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 / Laccophilus 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. / Laccophilus livens Régb. det. Régimbart 1891 / Ex Types" (1 ex. IRSNB; type material of Laccophilus 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 Laccophilus modestus and holotype of Laccophilus 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 Laccophilus modestus var. tostus and Laccophilus 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 Laccophilus 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 Laccophilus 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 Laccophilus 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.