Review of the Haliplidae of Madagascar, with Descriptions of Two New Species (Coleoptera) Bernhard Van Vondel & Johannes Bergsten

Tijdschrift voor Entomologie 155 (2012) 57–66 brill.nl/tve

Review of the Haliplidae of Madagascar, with descriptions of two new species (Coleoptera) Bernhard van Vondel & Johannes Bergsten

The Haliplidae of Madagascar are reviewed. Six species are known, including two species described here as new: Haliplus gamma sp. n. and Haliplus madagascariensis sp. n. Haliplus aspilus Guignot, 1957 is established as junior synonym of Haliplus incrassatus Régimbart, 1900. A lectotype is designated for Haliplus alluaudi Régimbart, 1903. Haliplus insularis Guignot, 1960 is redescribed. A key to the species of Madagascar is included. Some ecosystem conservation issues related to the occurrence of Haliplidae on Madagascar are discussed. Bernhard van Vondel*, Natuurhistorisch Museum Rotterdam, p/o Roestuin 78, 3343 CV Hendrik-Ido-Ambacht, The Netherlands. [email protected] Johannes Bergsten, Department of Entomology, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden. [email protected]

Introduction The examined material originates from the follow- ing institutions: Recently the Afrotropical species of the aquatic beetle family Haliplidae were revised (van Vondel 2010). MNHN Muséum National d’Histoire Naturelle, Paris, France (A. Mantilleri) Of three species from Madagascar the type material NMPC Museum of Natural History, Prague, could then not be found. Recently the junior au- Czech Republic (J. Hájek) thor discovered the missing types in the Muséum NMW Naturhistorisches Museum Wien, Vi- National d’Histoire Naturelle in Paris, together with enna, Austria (M. Jäch) an undescribed species. Examination of the type NHRS Swedish Museum of Natural History, material together with the discovery of another new Stockholm, Sweden (J. Bergsten) BMNH The Natural History Museum, London, species by the senior author justiﬁes a review of the UK (C. Taylor) Haliplidae from Madagascar. Based on this study six species are now known from Madagascar, of which two are here described as new. Distribution For all species the known distribution is illustrated and presented on provincial level, based on the Material and methods revision of the Afrotropical Haliplidae (van Von- del 2010) and the recently examined specimens. This study of forty specimens is an extension of the Madagascar is divided, from north to south, in six revision of the Afrotropical Haliplidae (van Vondel provinces: Antsiranana, Mahajanga, Toamasina, An- 2010) and methods used are the same. tananarivo, Fianarantsoa and Toliara.

Tijdschrift voor Entomologie 155: 57–66, Figs 1–22. [ISSN 0040-7496]. brill.nl/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 1 August 2012. DOI 10.1163/221194312X651382

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Account of Haliplidae known from 15. Metaventral process impressed in the mid- Madagascar dle...... 16 Checklist – Metaventral process with two impressions Haliplus (Liaphlus) alluaudi Régimbart, 1903 ...... 15a Haliplus (Liaphlus) gamma sp. n. 15a. Elytral maculation connected to suture . . . Haliplus (Liaphlus) incrassatus Régimbart, 1900 ...... Haliplus figuratus Haliplus (Liaphlus) aspilus Guignot, 1957 syn. n. – Elytral maculation not connected to suture Haliplus (Liaphlus) insularis Guignot, 1960 (Fig.5)...... Haliplus gamma sp. n. Haliplus (Liaphlus) madagascariensis sp. n. 21. Metaventral process with punctured Peltodytes (Peltodytes) quadratus Régimbart, 1895 grooves along lateral margins, flat in the middle...... Haliplus alluaudi Identification – Metaventral process with one or two pits or All species known from Madagascar are endemic, ovalimpressions...... 22 with the possible exception of Haliplus alluaudi, 22. Elytral maculation not connected to suture which is also reported from Mauritius (Guignot ...... 23 1959), so a key to the six species would be sufficient – Elytral maculation connected to suture . . . . 24 for identification. Haliplidae, however, are able to 23. Lateral margins of pronotum convex. Pro- spreadeasilyanditcannotbeexcludedthatspecies sternal process about parallel, metaventral of the African continent may reach Madagascar. process with separate impressions (Fig. 3) Therefore the key for all Afrotropical species is also ...... Haliplus insularis updated and the necessary changes are given below. – Lateral margins of pronotum concave. Prosternal process strongly narrowed be- Key to the Haliplidae of Madagascar fore coxae, metaventral process with two 1. Last segment of palpi longer than penulti- pits close together in a common impression mate segment. Elytral suture with fine line on ...... Haliplus tanzanianus posteriorhalf...... Peltodytes quadratus 24. Pronotumimmaculate...... 24a – Last segment of palpi shorter than penulti- – Pronotummaculate...... 25 mate segment. Elytral suture without fine line ...... 2 24a. Elytra only with discal dark mark in posteriorhalf(Fig.6)...... 2. Elytraimmaculate...... Haliplus incrassatus ...... Haliplus madagscariensis sp. n. – Elytramaculate...... 3 – Elytra with maculation on disc, along base 3. Elytral maculation on disk connected to su- andonintervals...... Haliplus nigerianus turaldarkening...... 4 – Elytral darkening not connected to sutural darkening...... 5 Haliplus (Liaphlus) alluaudi Régimbart 4. Elytral maculation restricted to the disk, nar- Fig. 1 rowingtowardsapex...... Haliplus alluaudi Régimbart, 1903: 1. Lectotype ...... Haliplus madagascariensis sp. n. (here designated): “Data in NHRS, JLKB- – Elytral maculation not only along suture and 000030267; Madagascar, Centre-Sud, Alluaud on disk, but also with marks on intervals . . . 1901: 66; Alluaudi Rég. n. sp. Typos; TYPE; Lec- ...... Haliplus alluaudi totype Haliplus alluaudi Régimbart 1903, des. 5. Elytral maculation in apical half with long Vondel & Bergsten 2012” (MNHN) [examined]. marks on even intervals ...... Haliplus insularis – Elytral maculation along base and suture, Remarks marks like Greek characters on elytra...... Van Vondel (2010) could not find type material in ...... Haliplus gamma sp. n. the collections of MNHN. Recently however, the junior author discovered two male and one female Corrections to the key of the Afrotropical syntypes, of which one is here selected as lecto- Haliplidae type. To include the species here described, Haliplus Additional material examined. 2 paralectotypes: 1 , gamma and H. madagascariensis, and the redescribed 1 , “Data in NHRS, JLKB-00003271 + 00003272, H. insularis, in the key of van Vondel (2010), the fol- Madagascar Sud, Fort Dauphin, Alluaud 1900, 8; lowing adaptations are necessary: Alluaudi Rég. n. sp.; Paralectotypes, left ,right,

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Figs 1–4. Haliplidae, dorsal view. – 1, Haliplus alluaudi;2,H. incrassatus;3,H. insularis;4,Peltodytes quadratus (photographs J. Bergsten). Scale line: 1 mm.

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Figs 5, 6. Haliplidae, dorsal view. – 5, Haliplus gamma sp. n.; 6, H. madagascariensis sp. n. (photographs: 5, J. Bergsten; 6, B. v. Vondel). Scale line: 1 mm.

Haliplus alluaudi Régimbart 1903”. 1 , Soalala, leg. Haliplus (Liaphlus) gamma sp. n. Perrier. 1 , Soalala, leg. Perrier. 1 , Ambongo, river Figs 5, 7, 8, 13, 16 Ambarizanahary [?], 1927, leg. S. Petit (MNHN). Type material. Holotype : Madagascar, Mahajanga, 1 , Madagascar: Antananarivo: Vakinankaratra: “Data in NHRS, JLKB-000030269; Soalála, Perrier; Manjakatompo Stn. Forestière 500 m E. Lac Froid Museum Paris Madagascar, coll. Perrier de la Bathie; by the road: S19.34485 E047.33381, 1620 m, n. sp.?; Haliplus sp. n. Det. J. Bergsten, 2011; Holo- 04.xi.2011, leg. J. Bergsten, R. Bukontaite, T. Ra- type Haliplus gamma Vondel & Bergsten 2012” narilalatiana & J.H. Randriamihaja (NHRS). 1 , (MNHN). 2 , Madagascar: Toamasina: Antsinanana: RN2, Ambodivoanio 41 km N Toamasina: S17.87002 Remarks E049.46249, 20 m, 16.xi.2011, leg. J. Bergsten, R. Although only one female specimen is available, Bukontaite, T. Ranarilalatiana & J.H. Randriamihaja itsverydistinctcharactersclearlydeﬁnethisasa (NHRS). 1 , Madagascar: Mahajanga; Melaky; Ts- separate species. ingy de Bemaraha NP S19.14114, E044.81245, 45 mao, 14.xii.2009, leg. J. Bergsten, N. Jönsson, T. Ra- Diagnosis narilalatiana, H.J. Randriamihaja (NHRS). The very peculiar elytral marks (resembling Greek characters) distinguish it from related species.

Description Distribution (Fig. 22) Length 2.7 mm, width 1.7 mm. Body oval. Known from all six provinces of Madagascar. Guig- Head: Yellow-brown, moderately punctured. Dis- not (1959) also mentions Mauritius but we have not tance between eyes 1.6× width of one eye. Antennae been able to conﬁrm this record. and palpi yellow to yellow-brown (Figs 5, 13).

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Figs 7–21. Details of Haliplus species. – H. gamma: 7, prosternal and metaventral process in ventral view; 8, prosternal process in lateral view; 13, antenna; 16, hind leg in dorsal view. – H. insularis: 9, prosternal and metaventral process in ventral view; 10, prosternal process in lateral view; 14, antenna; 17, hind leg in dorsal view. – H. madagascariensis: 11, prosternal and metaventral process in ventral view; 12, prosternal process in lateral view; 15, antenna; 18, hind leg in dorsal view. – H. incrassatus: 19, left paramere; 20, penis; 21, right paramere. Scale lines: 0.2 mm. Downloaded from Brill.com10/02/2021 02:55:32AM via free access 62 Tijdschrift voor Entomologie, volume 155, 2012

Fig. 22. Distribution map of Haliplus alluaudi, H. gamma sp. n., H. incrassatus, H. insularis, H. madagascariensis sp. n., Peltodytes quadratus.

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Pronotum: Yellow-brown without any dark mark. NOSSI-BÉ, H. PIERRON 1885; 178, 85; MU- Lateral borders strongly convex, lateral margin con- SEUM PARIS, COLL MAURICE REGIM- sisting of a row of punctures, slightly serrate. Ante- BART 1908; TYPE; LECTOTYPE , Haliplus rior edge hooked in the middle. Densely punctured incrassatus Rég., des. Vondel 2009” (MNHN) in anterior and posterior part (Fig. 5). [designation by van Vondel, 2010]. Elytra: Yellow-brown with a peculiar dark macu- Haliplus aspilus Guignot, 1957: 71. Holotype : lation: narrow line along base and anterior one third “Data in NHRS, JLKB-000030006; Type; IN- of suture, to first primary puncture row along poste- STITUT SCIENTIFIQUE MADAGASCAR; rior two third of suture, three undulated separated Sakahara, Lambomakandro, P.Griveaud; F.Guig- marks between third and ninth primary puncture not det. 1956, Haliplus (Liaphlus) aspilus n. sp., row on basal, middle and apical part of elytra respec- Type ; coll. IRSM, Guignot types; Holotype” tively, basal mark like the Greek character gamma, (MNHN) [examined]. Syn. n. apical mark running to apex (Fig. 5). Primary punctures moderately strong and dense, about 30 punc- Remarks tures in first row, row nine strong and dense and im- Van Vondel (2010) could not find type material of pressed in the middle. Secondary punctures strong and dense along suture, sparse on other intervals, Haliplus aspilus in the collections of MNHN, but row 2 and 6 with only few basal punctures. Punc- suggested the possibility that it is a junior synonym tures outside maculation not darkened. Completely of Haliplus incrassatus. Recently, the junior author margined, shoulder weakly serrate, apex smooth. discovered the female holotype of H. aspilus and also Ventral side: Body yellow-brown to brown. Ely- amaletypeofH. incrassatus. After examination of tral epipleura yellow-brown, reaching sixth sternite, the type material we consider both species conspe- strongly punctured in anterior part. Prosternal pro- cific. cess about parallel in anterior half, slightly widened The citation of van Vondel (2010) for Haliplus in anterior half, on each side a weak groove with incrassatus mentioned 1899, but this is corrected here strong punctures in posterior half, densely punctured to 1900. in anterior part (Figs 7, 8). Mesepimeron anteriorly with row of about five strong puncture. Proepis- Description ternum with few weak punctures in anterior part. The description by Régimbart (1900) and the re- Metaventral process on each side with a distinct im- description by van Vondel (2010) were based on a pression, moderately punctured (Fig. 7). Metacoxal female. The discovery of a male specimen makes it lobes moderately punctured. Punctures on fifth and possible to figure the male aedeagus (Figs 19–21). sixth sternites strong and dense, last sternite strongly The male has the first three tarsomeres of fore and punctured. Hind tibia with setiferous striole on two mid-legs widened and ventrally provided with sucker third of length of tibia, longer tibial spur about half hairs. of length of first tarsal segment (Fig. 16). Male: unknown. Distribution (Fig. 22) Endemic to Madagascar and known from the Etymology provinces of Antsiranana and Toliara in the western This species is named after the shape of the most part. anterior elytral mark, resembling the Greek character Additional material examined. Paralectotype: 1 , gamma (γ ). “Data in NHRS, JLKB-000030298; MUSEUM PARIS MADAGASCAR, NOSSI-BÉ, H. PIER- Biology RON, 1885; ; M. Régimbart dét.; Coll. Guig- No details known. not; Paralectotype Haliplus incrassatus Régimbart 1899”(MNHN). 1 , Madagascar, leg. Perrier de Distribution (Fig. 22) la Bathie. 1 , Madagascar, Ankavandra, vii.1949 Endemic to Madagascar and only known from the (MNHN). type locality Soalala in the province of Mahajanga. Haliplus (Liaphlus) insularis Guignot Haliplus (Liaphlus) incrassatus Régimbart Figs 3, 9, 10, 14, 17 Figs 2, 19–21 Haliplus insularis Guignot, 1960: 95. Holotype : Haliplus incrassatus Régimbart, 1900: 371. Lecto- “Data in NHRS, JLKB-000030005; Holotype; type : “MUSEUM PARIS, MADAGASCAR, INSTITUT SCIENTIFIQUE MADAGASCAR;

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Andobo, 190 m, forêt Antsingy, dct Antsalova, Biology -II-1957, P. Griv.; Guignot det. 1958, Haliplus No details known. (Liaphlus) insularis n. sp. Holotype; coll. IRSM, Guignot types” (MNHN) [examined]. Distribution (Fig. 22) Endemic to Madagascar and only known from Remarks the type locality Antsingy forest, Andobo, in the Van Vondel (2010) could not find type material province of Mahajanga. of Haliplus insularis in the collections of MNHN. Additional material examined. We have not seen any Recently, the junior author discovered the female other specimens of this species. holotype. This species is redescribed below. Haliplus (Liaphlus) madagascariensis sp. n. Figs 6, 11, 12, 15, 18 Diagnosis Type material. Holotype : “MADAGASCAR BEA- The marks on the even intervals of the posterior HITSE, Dam of Beahitse, 13.ii.2010, S24.17183 half of the elytra should distinguish it from related E44.46172, 318 m, M. Ferreux & P. Obrdlik species. [WWF]; coll. Jiri HAJEK, National Museum Prague, Czech Republic; Holotype Haliplus madagascarien- Description sis Vondel & Bergsten 2012” (NMPC). Length 3.5 mm. Width 2.0 mm. Body oval to subparallel in the middle. Remarks Head: Red-brown, densely punctured. Antennae Although only one female specimen is available, its yellow, first five or six segments red-brown (Figs 3, very distinct characters clearly define this as repre- 14). Palpi red-brown. senting a separate species. Pronotum: Red-brown, weak dark line in the middle from anterior to posterior edge. Lateral borders Diagnosis about straight and weakly serrate, finely margined. The discal elytral dark mark and the absence of other Densely, but not very strongly punctured, stronger markings distinguish it from related species. punctures along base, punctures not or hardly darkened. Weakly depressed along base (Fig. 3). Description Elytra: Yellow-brown with dark mark along suture Length 2.7 mm, width 1.5 mm. Body oval to sub- to secondary puncture row, long marks on even in- parallel. tervals in posterior 2/3 (Fig. 9). Primary punctures Head: Yellow-brown to brown, moderately, on moderately strong and dense, about 35 punctures in vertex more strongly punctured. Distance between first row (Fig. 3). Secondary punctures a little weaker eyes 1.3× width of one eye. Antennae yellow, dark- than primary punctures. Puncture rows about regu- ened towards base (Figs 6, 15). Palpi yellow-brown. lar. Completely margined, shoulders weakly serrate, Pronotum: Yellow-brown without any dark mark. apical part distinctly serrate. Lateral borders slightly convex, lateral margin with Ventral side: Body red-brown to brown. Epi- a row of strong punctures. Anterior edge hooked in pleura yellow-brown, moderately punctured in ante- the middle. Strongly punctured (Fig. 6). rior half. Prosternal process flat with a very weak im- Elytra: Yellow-brown, suture darkened to first sec- pression on each side, almost parallel, a little wider ondary puncture row, dark mark on disk in the mid- posteriorly than anteriorly, densely punctured in an- dle to third secondary puncture row tapering to apex terior half and along sides in posterior half (Figs 9, (Fig. 6). Primary punctures moderately strong and 10). Proepisternum weakly punctured. Mesepimeron dense (Fig. 6). Ninth primary puncture row im- with few weak punctures. Metasterrnal process with pressed. Secondary punctures only somewhat weaker two almost round impressions, weakly punctured than primary punctures. Secondary puncture row (Fig. 9). Metacoxal lobes strongly and densely punc- 4, 6 and 8 absent or with only few punctures. All tured. Punctures on fifth and sixth sternites in a punctures outside maculation not or hardly dark- dense row, last sternite sparsely punctured except in ened. Completely margined, shoulder and apex very the middle. Hind tibia with setiferous striole on one weakly serrate. third of length of tibia, longer tibial spur about two Ventral side: Yellow-brown to brown. Elytral epi- third of length of first tarsal segment (Fig. 17). pleura yellow-brown, reaching sixth sternite, strongly Males: unknown. punctured in anterior part. Proepisternum weakly

Downloaded from Brill.com10/02/2021 02:55:32AM via free access van Vondel & Bergsten: Review of the Haliplidae of Madagascar (Coleoptera) 65 punctured in anterior part. Mesepimeron with about Horák (NMW). 1 , Madagascar: Toamasina: Antsi- seven strong punctures. Prosternal process about par- nanana: RN2, Ambodivoatra village, S18.23794 allel in posterior half, slightly narrowed before coxae, E049.27577, 10 m, 14.xi.2011, leg. J. Bergsten, strongly channeled in the middle, strongly punctured R. Bukontaite, T. Ranarilalatiana & J.H. Randri- (Figs 11, 12). Metaventral process impressed in the amihaja (NHRS). 1 , Madagascar: Toamasina: middle, strongly and densely punctured (Fig. 11). Antsinanana: RN2, Ambodivoanio 41 km N Toa- Metacoxal lobes moderately punctured. Punctures masina: S17.87002 E049.46249, 20 m, 16.xi.2011, on fifth and sixth sternites strong and dense, last ster- leg. J. Bergsten, R. Bukontaite, T. Ranarilalatiana nite strongly punctured in apical part. Hind tibia & J.H. Randriamihaja (NHRS). 1 , Madagas- with setiferous striole on two-third of length of tibia, car: Toamasina: Alaotra Mangoro: Analamazoa- longer tibial spur about two-third of length of first tra SR. Close to park entrance: stagnant pools tarsal segment (Fig. 18). in tributary to Analamazoatra river, S18.93573 Males: Unknown. E048.41741, 930 m, 08.xi.2011, leg. J. Bergsten, R. Bukontaite, T. Ranarilalatiana & J.H. Randri- Etymology amihaja (NHRS). 1 , Madagascar: Toamasina: The name of this species refers to the country where Antsinanana: RN2, Onibe river 60 km N Toa- it is found. masina by bridge: ricefield, S17.65545 E049.4737, 20 m, 15.xi.2011, leg. J. Bergsten, R. Bukontaite, Biology T. Ranarilalatiana & J.H. Randriamihaja (NHRS). No details known. 1 , Madagascar: Toamasina: Antsinanana: RN5, btw. Toamasina and Fenerive: swamp, stream and pool, 15.xi.2011, leg. J. Bergsten, R. Bukontaite, Distribution (Fig. 22) T. Ranarilalatiana & J.H. Randriamihaja (NHRS). Endemic to Madagascar and only known from the 2 ,6, Madagascar: Toamasina: Antsinanana: type locality Beahitse in the province of Toliara. RN2, Fanandrana by the bridge, river Ivondro: S18.25612 E049.26886, 10 m, 14.xi.2011, leg. J. Bergsten, R. Bukontaite, T. Ranarilalatiana & J.H. Peltodytes quadratus Régimbart Fig. 4 Randriamihaja (NHRS). 1 , BMNH(E)<794141> DNA Voucher, Peltodytes quadratus Régimbart 1895 Peltodytes quadratus Régimbart, 1895: 10. Lec- Det. J. Bergsten, 2006, MAD:FIAN: Andringitra, totype : Madagascar, “Madagasc., Lac Am- Zornandao R. Bedrock, P39M12, N: −22.136: bod° [Lac Ambodinandohalo], R.P. Camboué; E46.889: 1601 m, 19.v.2006, leg. Andrianizehy et al. Muséum Paris, COLL. MAURICE RÉGIM- (BMNH). 1 , BMNH(E)<794143> DNA Voucher, BART 1908; TYPE; quadratus Rég.; Muséum Peltodytes quadratus Régimbart 1895 Det. J. Berg- Paris coll. gen. Peltodytes quadratus; Lectotype sten, 2006, MAD:TOLI: Zombitse, Manamboay designated by B.J. v. Vondel 1999; Peltodytes R.;muddyriverwaterP42E;N:−22.829 E44.6: quadratus Régimbart” (MNHN) (designation by 485 m, 14.v.2006, leg. Bergsten et al. (BMNH). 1 , van Vondel 2010). NHRSJLKB-000000732 DNA Voucher, Peltodytes quadratus Régimbart 1895 Det. J. Bergsten, 2012, Remarks Madagascar: Mahajanga; Melaky; btw. Antsalova- Régimbart (1895) based his description on more Maintirano S18.30233, E044.18071, 37 m, than one specimen. Van Vondel (2010) only found 18.xii.2009, leg. J. Bergsten, N. Jönsson, T. Ra- one of the syntypes (a female) and designated it narilalatiana, H.J. Randriamihaja (NHRS). 1 , lectotype. Recently the junior author discovered a Madagascar: Mahajanga; Melaky; btw. Antsalova- male syntype, which will be labeled as paralectotype. Maintirano S18.30233 E044.18071, 37 m, 18.xii.2009, leg. J. Bergsten, N. Jönsson, T. Ra- Distribution (Fig. 22) narilalatiana, H.J. Randriamihaja (NHRS). 1 , Endemic to Madagascar and known from all six NHRSJLKB-000000733 DNA Voucher, Peltodytes provinces. quadratus Régimbart 1895 Det. J. Bergsten, 2012, Additional material examined. Paralectotype : “Data Madagascar: Mahajanga; Melaky; btw. Maintirano- in NHRS, JLKB-000030268; Madagascar, Diego Morafenobe S18.03666 E044.49432, 212 m, Suarez 3, Ch, Alluaud 1893; TYPE; Paralectotype 18.xii.2009, leg. J. Bergsten, N. Jönsson, T. Ra- Peltodytes quadratus Régimbart 1895” (MNHN). narilalatiana, H.J. Randriamihaja (NHRS). 2 , 1 , N. Madagascar, Antsiranana distr., Sambi- Madagascar: Mahajanga; Melaky; btw. Maintirano- rana river, Moarovato vill., 5–12.xii.2001, leg. J. Morafenobe S18.03666, E044.49432, 212 m,

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18.xii.2009, leg. J. Bergsten, N. Jönsson, T. Ranari- Ranarilalatiana, Jacquelin Herisahala Randriamihaja, lalatiana, H.J. Randriamihaja (NHRS). Rasa Bukontaite and Niklas Jönsson helped with fieldwork. The Department of Entomology at An- tananarivo University and Madagascar Institute for Implications for conservation the Conservation of Tropical Ecosystems (MICET) The focus on Madagascar’s threatened biodiversity is are thanked for continuous fieldwork support. The on the rainforest belt in the eastern and northern part Uyttenbogaart-Eliasen Foundation is thanked for fi- of the country containing the highest level of diver- nancial support of the senior author’s research on sity. The western regions of Madagascar are consid- Haliplidae in general. ered less diverse but are also less explored. It is clear from this review that Haliplidae is one group that contradicts the main pattern and is most diverse in References the drier western parts of Madagascar (Fig. 22). The Benstead, J.P., P.H. De Rham, J.L. Gattolliat, F.M. Gibon, dry deciduous forest in western Madagascar is also P.V. Loiselle, M. Sartori, J.S. Sparks & M.L.J. Stiassny, the habitat most endangered, much less remaining 2003. Conserving Madagascar’s freshwater biodiversity. compared to the eastern rainforests, and it is con- – BioScience 53: 1101–1111. tinuously diminishing through subsistence agricul- Ganzhorn, J.U., P.P. Lowry, G.E. Schatz & A. Sommer, ture (“slash and burn”) and small-scale tree harvest 2001. The biodiversity of Madagascar: one of the (Ganzhorn et al. 2001). More than 97% of Mada- world’s hottest hotspots on its way out. – Oryx 35: gascar’s dry deciduous forests have been lost and the 346–348. remnants are severely fragmented and affected by Guignot, F., 1957. Un Haliplide et deux Dytiscides nou- human activities (Whitehurst et al. 2009). Aquatic veaux de Madagascar. – Le Naturaliste Malgache 9: 71– habitats change dramatically as a result of deforesta- 74. tion (Benstead et al. 2003). The main factors are Guignot, F., 1959. Revision des Hydrocanthares d’Afrique the increased sediment load caused by erosion, in- (Coleoptera Dytiscoidea), part 1. – Annales du Musée Royal du Congo Belge Tervuren (8) Sciences Zo- creased insolation and water temperature, increased ologiques 70: 1–316. nutrient load and changed food resources (Benstead Guignot, F., 1960. Haliplides et Dytiscides nouveaux ou et al. 2003). Increased nutrient load is also caused by interessants de Madagascar. – Le Naturaliste Malgache Zebu, introduced cattle, and, together with the ero- 11: 95–101. sion, can completely change the invertebrate fauna Hanski, I., H. Koivulehto, A. Cameron & P. Rahagalala, of a forest river from endemic Madagascan species to 2007. Deforestation and apparent extinctions of en- widespread non-endemic, often continental African, demic forest beetles in Madagascar. – Biology Letters taxa. It is not unlikely that some of the Haliplidae 3: 344–347. species only known from single specimens from the Régimbart, M., 1895. Revision des Dytiscidae et Gyrinidae beginning (H. gamma sp. n.) or middle (H. insu- d’Afrique, Madagascar et iles voisines. En contribution laris)ofthe20th century are already extinct. Hanski a la faune entomologique du Congo. – Mémoires de la et al (2007) found that the main explanatory vari- Société Entomologique de Belgique 4: 1–244. able of dung-beetle species not re-found in recent Régimbart, M., 1900. Diagnoses d’espèces nouvelles de times was the loss of forest within the historical range Dytiscidae de la région Malgache (Col.). – Bulletin de of species. Only about 10% of Madagascar’s orig- la Société Entomologique de France 1899(19): 371– inal forest cover remain and ecological theory sug- 374. gests that this is enough to maintain around 50% of Régimbart, M., 1903. Coleopteres aquatiques (Halipli- dae, Dytiscidae, Gyrinidae et Hydrophilidae) recueillis the diversity (Hanski et al. 2007). This means that dans le sud de Madagascar par M. Ch. Alluaud (Juil- 50% of the original biodiversity of Madagascar is ei- let 1900–Mai 1901). – Annales de la Société Ento- ther already extinct or doomed to extinction in the mologique de France 72: 1–51. long term. Conservation efforts in Madagascar must Vondel, B.J. van, 2010. Revision of the Haliplidae of the treat freshwater systems as a high priority, especially Afrotropical region, including North Africa (Coleop- in the dry western parts, and more research is needed tera). – Tijdschrift voor Entomologie 153: 239–314. to map the invertebrate freshwater diversity. Whitehurst, A.S., J.O. Sexton & L. Dollar, 2009. Land cover change in western Madagascar’s dry deciduous forests: a comparison of forest changes in and around Acknowledgements Kirindy Mite National Park. – Oryx 43: 275–283. We wish to express our sincere thanks to the cura- tors of the museums in Paris, Prague, London and Received: November 17, 2011 Vienna for placing material at our disposal. Tolotra Accepted: May 7, 2012

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