AbstractCopelatus Erichson, 1832  of Madagascar is revised in two parts. This review is restricted to the Copelatus species that have fewer than ten elytral + one submarginal stria, including all species except those of the erichsonii species group. Both morphological and molecular  data are used in an integrative taxonomic approach. Thirteen species are recognised, of which five are described as new: Copelatusankaratrasp. nov., Copelatuskelysp. nov., Copelatuspseudostriatussp. nov., Copelatussafiotrasp. nov. and Copelatusvokokasp. nov.Copelatusunguicularis Régimbart, 1903 and Copelatusapicalis Fairmaire, 1898 are both transferred to the genus Madaglymbus Shaverdo & Balke, 2008 (comb. nov.). CopelatusmimetesCopelatuspulchellus  (syn. nov.). Copelatusmarginipennis  is reinstated (stat. nov.) as a valid species with Copelatusaldabricus Balfour-Browne, 1950 and Copelatusaldabricusvar.simplex Guignot, 1952 as junior synonyms (syn. nov.). We designate lectotypes for Colymbetesmarginipennis Laporte, 1835 and Copelatusobtusus Boheman, 1848. Copelatusperidinus Guignot, 1955 is recorded for Madagascar for the first time and Copelatusnodieri Régimbart, 1895 is rejected as a species present in Madagascar.The genus  Copelatinae of diving beetles is a diverse group of aquatic beetles represented by eight genera: Agaporomorphus Zimmermann, 1921, Aglymbus Sharp, 1880, Capelatus Turner & Bilton, 2015, Copelatus Erichson, 1832, Exocelina Broun, 1886, Lacconectus Motschulsky, 1855, Liopterus Dejean, 1833, and Madaglymbus Shaverdo & Balke, 2008. This subfamily is relatively homogeneous morphologically, and only the single tribe Copelatini is recognised  and one originally mislabelled species removed from the fauna by The subfamily cognised , 2016. Tcognised . After taglymbus , this su species . Rocchi Copelatus species have traditionally been organised in a number of species groups based on the number of elytral and submarginal striae following Copelatinae with mitochondrial CO1 and 16S, found the number of elytral striae to be a highly variable and homplastic character. High variability in number of striae, both intraspecifically and between closely related species, was recently documented in Antillean species , longicornis , irinus , consors  and erichsonii  groups. About half of the described species are members of the erichsonii species group, and the other half of the remaining species groups with fewer striae. In this paper, we will treat the Copelatus species of Madagascar with fewer than 10 elytral and one submarginal stria, i.e., all the non-erichsonii group species. Again, this is a practical rather than a phylogenetic division and in fact Copelatussafiotra sp. nov., described below, we believe is most closely related to the species of the erichsonii group due to the shape of male genitalia.The species that occur in Madagascar fall into five of these species groups: the irinus species group, together with type material and some additional museum accessions. We use both morphological and molecular data with a species delimitation analysis, in an integrative taxonomic approach to the revision. We provide an identification key and for each species a diagnosis, description, known distribution and habitat and ecology notes. Each species is also illustrated with a dorsal habitus photograph, and ventral and lateral views of male penis and parameres.This study is motivated by recent collecting efforts of aquatic beetles in Madagascar 2009–2018 in the Water Beetles of Madagascar Project. The project is a collaboration between the Swedish Museum of Natural History and the University of Antananarivo. The current study is based on this rich new material containing five new species of the Copelatus can be found in diverse aquatic habitats, both running (lotic) and standing (lentic) waters. Along forest streams they are best sought for in stagnant parts such as side-pools and rock pools with dead leaves or vegetation or in residual pools in a partly dried out streambed. Leaf-choked forest pools, vegetation-rich edges of ponds and lakes and marshes are also very good habitats for Copelatus. Some species are very good fliers and readily come to light, but the proportion of light catches (using a 22W black-ringlight) in the studied material is relatively small. A surprising number can be found even in minute seeps especially if visited with head lamp at night when they are more active and swim around. Copelatus was sampled mainly using white pans and GB water nets and sieves with various sizes depending on waterbody type. GB water net is often best for larger habitats while sieves were used for sampling smaller habitats like rockpools. Specimens were collected into plastic tubes with 95% ethanol for conservation.New collecting efforts of aquatic beetles were conducted mainly in National Parks, reserves and natural forests but also in degraded forests, open areas and along main roads from all parts of Madagascar, except scant from the very south Fig. . The fieEach locality was given a collecting event code and associated metadata included geographic name(s), forest type, waterbody type, habitat description, eventual disturbance, collecting date and collectors. Altitude, latitude and longitude were recorded with a handheld GPS (Garmin). Each locality was also documented with photographs using compact Canon and Olympus digital cameras.Madagascar is since the 2007 revision of the constitution divided into 22 regions as a first level administrative division, followed by districts as a second level, a recent change from the former six provinces as first level Fig. . StudiedNHRS), Parc Botanique et Zoologique de Tsimbazaza/ Madagascar Biodiversity Center (PBZT/MBC) and Department of Entomology, Antananarivo University (DEUA). Some material originated from efforts organised by the Natural History Museum in London (NHMUK) 2004–2007.Most of the studied material came from the new fieldwork and is shared between the Swedish Museum of Natural History , Museum of Natural History, London (NHMUK), California Academy of Sciences (CAS), Naturhistorisches Museum, Vienna (NMW), and National Museum , Prague (NMPC). Studied older types are housed in Paris (MNHN), London (NHMUK), Stockholm (NHRS) and Prague (NMPC), while holotypes of newly decribed species are housed in Stockholm (NHRS).Additional studied material came from earlier expeditions housed at museum collections in NHRS, DEUA & PBZT/MBC” given as depository will also be shared with other central institution collections.Depositories of studied specimens are referred to by above abbreviations . Genitalia were extracted with a fine forceps or pin from the tip of the abdomen and glued onto cards on the same pin as the specimen. Dry-preserved specimens were first relaxed in warm water for 5–20 minutes before genitalia were carefully extracted. Photos of habitus were taken with a Canon EOS 5D Mark II DSLR camera equipped with a Canon MP-E 65mm 1–5X super macrolens and mounted on a motorised rail (Stackshot) from Cognisys. The system was operated using Canon EOS Utility and Zerene Stacker (Zerene Systems) softwares, the latter also used for stacking the Z-stack of captured images with the PMax or DMap algorithms. Photographs of dry-mounted genitalia were taken with a Canon EOS 7D DSLR camera mounted on a BALPRO 1 Universal bellow from Novoflex with a long working distance 10X Plano apochromatic microscope objective from Mitutoyo. The bellow was mounted on a motorised rail (Stackshot) from Cognisys and operated with the same software given above.In describing the male penis we use the terminology suggested by NHRS”, “BMNH” (=NHMUK) or “CAS”, followed by a number. A series of specimens with consecutive catalogue numbers are given as a range. The following additional abbreviations are used:Label data are given as written and separated by “//” if on separate labels and “|” if on different rows on the same label. Text within square brackets “” are our comments, explications or interpretations. Most examined specimens  have been given unique catalogue numbers and these are listed first, starting with “Alc. in alcohol tube,Ex. exemplars ,GP  male genitalia have been examined,HT Holotype,LT Lectotype,PLT Paralectotype,PT Paratype,ST Syntype,TL Type locality,DNA was extracted from one mesoleg or from soft abdominal tissue retrieved in association with dissection of male genitalia. The leg or soft tissue was incubated in lysis buffer at 56 °C overnight. Post-incubation protocol followed the cell and tissue DNA kit on a KingFisher Duo Prime. This system provides automated nucleic acid purification at a running time of approx. 25 minutes.cox1)  (CASENT-8135000) and Copelatus sp. female (Andasibe) (NHRS-JLKB000065698), we used the primers Jerry  – Hal 1450rw ; and Hal 1450fw  – PatDyt  to amplify the same segment but in two shorter sections  and PatDyt  to amplify an 825 bp fragment of mitochondrial cytochrome c-oxidase subunit 1 gene (COI or cox1) . For twosections . The PCRSuccessful PCR products were purified using EXOSAP Clean-up mix of two enzymes  and run in a PCR machine with the programme 37 °C for 30 min followed by 80 °C for 15 min and finally 12 °C (∞). PCR products were sent to Macrogen for sequencing.Copelatus from MK878825-MK878877 (Table Gene regions were sequenced in both directions and sequence chromatograms were edited with SEQUENCHER version 4.10.1 (Gene Codes Corporation). The contigs were assembled from forward and reverse reads, and primer regions trimmed. New sequence data were then exported in fasta format and aligned together with GenBank sequences of Copelatusbefasicus. We used PARTITIONFINDER Ver. 2.1.1 , the genetree was artificially rooted using r. 2.1.1  to inferr. 2.1.1  to inferth generation, and treeannotator was used to calculate a maximum clade credibility tree with median node heights, removing 10% as burn-in from each run. The GMYC analysis was carried out in R using the SPLITS package  and 447 bp (NHRS-JLKB000065698) respectively and enabled inclusion in the phylogenetic analyses.Our amplification of CO1 was successful for 53 samples which, together with sequences downloaded from GenBank, gave 77 terminals  closest to the morphological delimitations, the Copelatusinsuetus complex was divided into two species, but also the northernmost Copelatussafiotra sp. nov. specimen was separated from geographically more southern populations. The dry-preserved female from Bemaraha (CASENT-8135000) undoubtedly represents a fourth Malagasy species from the Copelatuspulchellus complex. However, the female from Andasibe (NHRS-JLKB000065698) was merged with Copelatusperidinus, despite a genetic distance of approximately 3%, and this is discussed below.The GMYC species delimitation analysis of the strict clock tree resulted in 11 separate evolutionary units that were largely but not entirely consistent with our morphological delimitation Fig. . EspeciaCopelatus species of Madagascar with fewer than ten discal and one submarginal elytral striae. Note that for some species males are necessary.Key to Copelatus species from Madagascar and were listed as such by The following three taxa were described as Taxon classificationAnimaliaColeopteraDytiscidae0A694D306A4456EB8ADB140766C419B7Copelatussubjectus Sharp, 1882: 568.CopelatusbilunatusTL: Madagascar .Mahajanga. Betsiboka: Maevatanana: -ST ♂ (GP) : // Data in NHRS | JLKB | 000030208 // Madag. Perrier | // Type [red writing] // Museum Paris | Coll. Régimbart | apicalis // Copelatus | apicalis n. sp. //Copelatusapicalis was described by Copelatus and currently placed in the Copelatushydroporoides species group  comb. nov.; in fact Aglymbus “Peut-être c’est une Aglymbus?” [Eng. translation “Maybe it is an Aglymbus”], which was correct since this was before Madaglymbus was erected for the Madagascar species of Aglymbus comb. nov.45AB55AB28805B229F01D47FDED687E7Copelatusunguicularis Régimbart, 1903:19.Madagascar, Suberbieville [= Maevatanana].Mahajanga. Betsiboka: Maevatanana: -HT ♂ (GP) : // Data in NHRS | JLKB | 000030225 // Suberbieville | Madag. Perier // Type [red label] // MUSEUM PARIS | Coll. Maurice Régimbart | 1908 // C. unguicularis | Type // unguicularis Rég. //Copelatusunguicularis was described by consors species group sensu Copelatusapicalis, it has a stout spine-like setae on a protruding anterodistal corner of protarsomere IV and is not a Copelatus. In addition, the elytral striae are very irregular, more like very elongate and deep strioles and possibly not homologous to the regular impressed striae in Copelatus. We here transfer it to Madaglymbus: Madaglymbusunguicularis  comb. nov.Copelatus species lacking elytral striae, but it is very probably not monophyletic and likely still mixed with some copelatine species that should be transferred to other genera.This group is defined by the lack of elytral striae . It is cTaxon classificationAnimaliaColeopteraDytiscidaeGuignot, 19550DA77899943B5226BBA57EB36328B122Copelatusbaculiformis Guignot, 1955b: 193.Madagascar, Massif Ankaratra, Manjakatompo, alt. 1700–1800 m.based on a single female specimen (holotype), collected December 1951 by R. Benoist.Antananarivo. Vakinankaratra: Ambatolampy: -HT ♀ (MNHN “coll. Guignot”): // Data in NHRS | JLKB | 000030226 // Madagascar: Massif An- | karatra 1700/1800 Man- | jakatompo XII-51 Benoist // Type [red label] // Guignot det., 1954 | Copelatus | baculiformis n. sp. | Type ♀ //Madaglymbus. The absence of deep impressed elytral striae  separates this species from all other Copelatus of Madagascar except Copelatusperidinus, a larger species (5.7–6.6 mm).Small size (4 mm). Elytra uniformly dark brown ferrugineous, without a basal testaceous area Fig.  which se(based on holotype ♀):Body length 4 mm. Body shape elongate oval and dark brown to blackish ferrugineous. Head uniformly dark brown ferrugineous to slightly darker posteriorly inside eyes, with thin sparse punctation. Pronotum dark brown ferrugineous, same colour medially and laterally but darker along anterior and posterior third. Disc of pronotum less densely punctuated, posterolateral corners with dense superficial strioles. The entire dorsal surface covered with a microsculpture. Elytra uniformly coloured in same dark brown to blackish ferrugineous colour as anterior and posterior parts of pronotum Fig. . ImpressVentral side testaceous to weakly infuscated. Prosternal process carinate also onto apical process. Lateral parts of metaventrite  rather broad. Metacoxal lines anteriorly diverging and abbreviated before metaventral margin. Metacoxa with fine and long longitudinal strioles, continuing onto abdominal ventrites, and with 6–7 transverse “wrinkles” anterolaterally.Male: unknown.Madagascar, central highlands, only known from type locality Manjakatompo, Ankaratra Massif Fig. .Unknown, but according to original description collected at at an altitude of 1700–1800 m. See hydroporoides species group of Copelatus, but two other Malagasy species placed in this group have turned out to be misidentified Madaglymbus or Exocelina (see above). Copelatusbaculiformis was described by Guignot the same year (1955) that he described C.bilunatus, considered mislabeled , Manjakatompo , 1700–1800 m , XII-51 (collecting month and year), and R. Benoist (collector), speaking against such a mistake. Based on general external morphology, body shape and lack of striae the species resembles C.peridinus. It is much smaller and therefore likely not conspecific, but they could be closely related, as well as with the female sequenced from Andasibe (NHRS-JLKB000065698).No other specimen than the female holotype is known of this species and it is a bit of a “mystery species”. We have conducted fieldwork at the type locality Manjakatompo multiple times  but never found any specimens resembling this species. The species belongs in the slabeled  from theTaxon classificationAnimaliaColeopteraDytiscidaeGuignot, 195525AF5D96C7285A06B37C8F9CC93796E0Copelatusperidinus Guignot, 1955c: 188.CopelatusseydeliTL: Elisabethville, Zaire . Guignot, 1958: 107; Elisabethville, Zaire .PT ♀ : // Data in NHRS | JLKB | 000030317 // Allotype [red label] // [female symbol] // Elisabethville | XI. 1951 // Copelatus | peridinus | Allotype ♀ //-GP) : // Data in NHRS | JLKB | 000030318 // Congo Belge Lac | Edouard: Vitshumbi | U.V. 27.XI.1953 | 3091 // Antananarivo. Analamanga: Antananarivo: -3♂(GP), 3♀ : // Data in NHRS | JLKB | 000030319–24 // MADAGASCAR | TANANARIVE | BETONGOLO | 2 XII 1946 // Piège | lumineux // Museum Paris | 1983 | Coll. Cl. Legros // Toamasina. Alaotra Mangoro: Moramanga: -1♂(GP), 2♀ : // Data in NHRS | JLKB | 000030316, 30325–6 // Madagascar Est | P.K.57-Rte d’Anosibe | Moramanga | II.58 R. Vieu // -1♂(GP), 1♀ : // Data in NHRS | JLKB | 000030327, 30328 // Madagascar | Lokato, near | Andasibe Mantadia NP | M. Tryzna leg., 9–10.i.2007 // coll. Jiri HÁJEK | National Museum | Prague, Czech Republic // [“Lokato near Andasibe Mantadia NP” interpreted as the bifurcation of the road to Lakato which is near Andasibe Mantadia NP. Lakato itself is not near Andasibe Mantadia NP.] -2♂(GP), 1♀, 1♀, 1♂  (NHRS): // NHRS-JLKB | 000010887, 65737, 65703, 10888(Alc.) // MAD: TOAM: Alaotra Mangoro: | Andasibe Mantadia NP, Analamazaotra: | 150m E of park entrance: Mad14-14 | shallow partly dried out forest pond: | 18.9355S | 48.4166E: 930m: 27.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja & T. Ranarilalatiana //-1♂ at which it curves slightly more abruptly creating two gentle angles  indicate. In Madagascar, only know from the eastern central parts: Betongolo (Antananarivo), the Analamazaotra NP, and P.K.57 Route d’Anosibe [RN23] Fig. .A series of six specimens collected with light trap (“piège lumineux”) in the capital Antananarivo 1946, indicates flight capacity and anthropogenically disturbed habitats. All records from DR Congo are also from light trap catches . We collThis species was described from Lubumbashi, DR Congo, and has not been recorded from Madagascar before. Earliest record found is from November 1946. It seems to be a dispersive good flier and often collected at light so its presence in Madagascar is therefore not surprising. However, it is not widespread in Madagascar as far as we know. In fact, the known distribution is restricted to the surroundings of the capital and east of the capital along the main national route towards Toamasina, which could suggest a recent incidental human-mediated introduction from mainland Africa.Copelatus sp. 2 below).Note that it may be that this is a species with intraspecific variation with regards to elytral striation, ranging from five puncture lines out of which two are more distinct, to five weakly impressed striae : // Data in NHRS | JLKB | 000030021 // Morondava | fôret sud | de Befasy | I-56 R.P // Type [red label] // INSTITUT | SCIENTIFIQUE | MADAGASCAR // Guignot det., 1956 | Copelatus | befasicus n. sp. | Type // -2PT ♀ : // Data in NHRS | JLKB | 000030300-1// Morondava | fôret sud | de Befasy // I-56 R.P // Paratype [red label] // [female symbol] //Mahajanga. Melaky: Morafenobe: -1♀ (NHRS): // NHRS-JLKB | 000010860 (JB204) // Madagascar: Mahajanga: Melaky | Btw. Morafenobe–Ambohijanahary | S18.19091; E045.19986, 290 m.a.o | 19.XII.2009 Water Net, Field# MAD09-74 | Leg: J. Bergsten, N. Jönsson, T. | Ranarilalatiana, H.J. Randriamihaja //C.insuetus and related species in habitus by being small, elongate, and subparalell, but C.befasicus is distinguished from C.insuetus and from all other Malagasy Copelatus species by the presence of only five elytral striae, and without submarginal striae. In addition, the first stria is shortened, present only in the posterior third .The l striae . There al striae : CopelatTaxon classificationAnimaliaColeopteraDytiscidaeRégimbart, 190348693E4733015386A1417218CDEAFB7ACopelatusdistinguendusCopelatusduodecimstriatus Aubé in  Régimbart, 1903: 19  and Fianarantsoa, Madagascar.Based on an unknown number of specimens (syntypes) collected by Sikora (Antananarivo) and Perrot (Fianarantsoa).MNHN not studied, as it was out on loan.Type material in Fianarantsoa. Matsiatra Ambony: Ambalavao, Ambohimahasoa, Lalangina: -2♀, 1♂(GP) (NHMUK): // BMNH-792954–6 // MAD: FIAN: Andringitra | Zomandao R.: River edge: Bottle trap | P39EM08: N: -22.1043: E:46.92: 1420 m | 09/V/2006: Leg. Bergsten et al // BMNH (DNA Voucher) // -1♀ (NMW): // Data in NHRS | JLKB | 000010718 // Madagascar: Ambohimahasoa (Fianarantsoa) | RN7 (Km 378) | 16.04.2011: Leg. R. Gerecke (MD211) // spring area with meadow swamps Exp. E: 21°15'41.5"S, 47°14'10.9"E, 1500 m // -2♀ (NMW): // Data in NHRS | JLKB | 000065754–5 // Madagascar Est, 1100– | 1200m, P.N. Ranomafana // Vohiparara, 21.–24.1.1993 | J. Janák lgt // -1♂(GP), 1♀, 5 ex. (Alc.) (NHRS): // NHRS-JLKB | 000010618–9, 10787(Alc.) // Madagascar: Fianarantsoa: Matsiatra | Ambony: Ranomafana NP: | Sahamalaotra 2Km from Vohiparara: | S21.23807, E047.39489, 1140 m.| 01:XI:2011: stamping with sieves: forest | bog in rainforest: Field# MAD11-12 // Leg. J. Bergsten, R. | Bukontaite, T. | Ranarilalatiana & | H.J. Randriamihaja // -1♀(Alc.) (NHRS): // NHRS-JLKB | 000010830 // MAD: FIAN: Matsiatra Ambony | Ranomafana NP: 450m along | Sahamalaotra trail, left at the first | junction: Mad14-07: forestmarsh: | 21.2382S 49.3947E: 1130 m: 02.XI.2014 // Leg. J. Bergsten, | T. Ranarilalatiana | & S. Holmgren // -1♀ (NHMUK): // BMNH-670601_MSL007 | 06.xii.2004, Ranomafana, | Madagascar: lat - 21.2359 | Lon 47.3963 Coll Balke_M; | Monaghan_M // DNA Voucher | BMNH <670601> | MSL007:E07 // Fianarantsoa. Amoron’i Mania: Ambositra: -3♀ (NHMUK): // BMNH-792962–4 // Col de Tapias: Rte Tana–Fianarantsoa: Pond | P36C: N: -20.772: E:47.179: 1717 m | 06/V/2006: Leg. Bergsten et al // BMNH (DNA Voucher) // -1♀ (NHMUK): // BMNH-792912 // Ambositra: Ankazomivady forest | 01.xii.2005 // BMNH (DNA Voucher) // -9♀ (NHMUK), 8 ex. (Alc.) (NHRS): // BMNH-729890, 729893, 729896–7, 792976–80, 10793(Alc.) // 08.xii.2004, Col de Tapias, | P30MD33: lat -20.238 | Lon 47.1 Coll Balke_M | Monaghan_M // BMNH (DNA Voucher) // Antananarivo. Vakinankaratra: Ambatolampy: -3♂(GP), 3♀, 10 ex. (Alc.) : // NHRS-JLKB | 000010620–5, 10786(Alc.) // Madagascar: Antananarivo: | Vakinankaratra: Manjakatompo Stn. | forestière: 500m E Lac Froid by the | road: S19.34485 E047.33381, 1620 m. | 04.XI.2011: GB Nets and sieves: pond | and inlet stream: MAD11-16 // Leg. J. Bergsten, R. | Bukontaite, T. | Ranarilalatiana & | H.J. Randriamihaja // -1♂(GP) (NHRS): // NHRS-JLKB | 000010626 // Madagascar: Antananarivo: | Vakinankaratra: Manjakatompo Stn. | forestière: Analafandriana 500 m S | fish farm by the road: S19.36191 E | 47.31495, 1730 m, 03.XI.2011: GB | Nets: grassy pond: Field# MAD11-14 // Leg. J. Bergsten, R. | Bukontaite, T. | Ranarilalatiana & | H.J. Randriamihaja // -1♂(GP) (NHRS): // NHRS-JLKB | 000010660 // Madagascar: Ambatolampy: Manjaka- | tompo Ankaratra Reserve: MAD16-03: | “Lac froid”: S-19.34292; E047.33893; | 1651 m: lake with grass at margins: | 03/02/2016 Leg. T. Ranarilalatiana // -4♂(GP), 4♀, 5 ex. (Alc.) : // NHRS-JLKB | 000010662–9, 10791(Alc.) // Madagascar: Ambatolampy: Manjaka- | tompo Ankaratra Reserve: MAD16-46: | “Lac froid”: S-19.34292; E047.33893; | 1651 m: lake with grass at margins: | 17/09/2016 Leg. T. Ranarilalatiana // -1♂(GP) (NHRS): // NHRS-JLKB | 000010661 // Madagascar: Ambatolampy: Manjaka- | tompo Ankaratra Reserve: MAD16-10: | Ankafotra mountain: S-19.33753; | E047.24530; 2466 m: streampools: | 07/02/2016; Leg. T. Ranarilalatiana // -4♂(GP), 2♀, 18 ex. (Alc.) : // NHRS-JLKB | 000010670–5, 10792(Alc.) // Madagascar: Ambatolampy: Manjaka- | tompo Ankaratra Reserve: MAD16-47: | Ankafotra mountain: S-19.33753; | E047.24530; 2466 m: streampools: | 18/09/2016; Leg. T. Ranarilalatiana // -1♀ (Alc.) (NHRS): // NHRS-JLKB | 000010832 // Madagascar: Antananarivo: | Vakinankaratra: Manjakatompo Stn | forestière: Analamitana: S19.363972 E | 047.299083, 1757 m. 22:I:2012: swamp | near stream: Field# MJK12-02: Leg. T. | Ranarilalatiana & J.H. Randriamihaja // Antananarivo. Analamanga: Anjozorobe, Ankazobe: -1♂(GP), 1♀ (NHRS): // NHRS-JLKB | 000010684–5 // Madagascar: Anjozorobe: MAD16-36: | Amboasarianala: S-18.45792; E047. | 93438; 1367 m: Ambatovikinina stream: | 04/04/2016; Leg. T. Ranarilalatiana // -1♂(GP), 1♀ (NHRS): // NHRS-JLKB | 000010686–7 // Madagascar: Anjozorobe: MAD16-43:| Amboasarianala, Antanambe stream: | S-18.4671; E047.93807; 1271 m: Stream | with sidepools: 07/04/2016 | Leg. T. Ranarilalatiana // -2♂(GP), 2♀, 1♀(Alc.) (NHRS): // NHRS-JLKB | 000010688–91, 10790(Alc.) // Madagascar: Anjozorobe: MAD16-44 | Amboasarianala, Mangarivotra stream: | S-18.4676; E047.92535; 1271 m: stream | with bedrock and grass at edge | 07/04/2016; Leg. T. Ranarilalatiana // -2 ex. (Alc.) (NHRS): // NHRS-JLKB | 000010831 // MAD: ANTA: Analamanga: Anjoz | orobe forest reserve: Marsh next | to the stream by Saha forest, 10Km E of Anjozorobe: MAD14- | 78: forestmarsh: 18.4128S | 47.9439E; 1320 m; 23.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | & T. Ranarilalatiana // -2♂(GP), 2♀, 6 ex. (Alc.) : // NHRS-JLKB | 000010676–9, 10797(Alc.) // Madagascar: Ankazobe: MAD16-24: | Firarazana: S-18.13132; E047.23976; | 1551 m; Lake with grass at margins: | 12/03/2016; Leg. T. Ranarilalatiana // -2♀, 11 ex. (Alc.) : // NHRS-JLKB | 000010680–1, 10789(Alc.) // Madagascar: Ankazobe: MAD16-26: | Maharidaza, Large stream by the road to | military camp: S-18.22102; E047.27087; | 1547 m: stream and bog with grass: | 14/03/2016; Leg. T. Ranarilalatiana // -1♂(GP), 1♀, 7 ex. (Alc.) (NHRS): // NHRS-JLKB | 000010682–3, 10788(Alc.) // Madagascar: Ankazobe: MAD16-29: | Firarazana, SW of Ambohitantely | reserve: S-18.16717; E047.26090; | 1532 m: Bog with grass: 17/03/2016; | Leg. T. Ranarilalatiana // -3 ex. (Alc.) (NHRS): // NHRS-JLKB | 000010829 // MAD: ANTA: Analamanga: Mana- | nkazo river by the bridge of | RN4: Mad14-75: medium size river over bedrock: 18.158S | 47.2104E: 1450 m: 21.XI.2014 // Leg. J. Bergsten, | J.H. Randriamihaja | & T. Ranarilalatiana // -14 ex. (Alc.) : // NHRS-JLKB | 000010834 // MAD: ANTA: Analamanga: Andra- | nofeno river by the bridge of | RN4, next to Andranofeno Sud | village: Mad14-74: medium size, | slow flowing river: 18.0844S | 47.1776E: 1430 m: 21.XI.2014 // Leg. J. Bergsten, | J.H. Randriamihaja | & T. Ranarilalatiana // - 1♀ (Alc.) (NHRS): // NHRS-JLKB | 000010835 // Madagascar: Ankazobe: TR18L10: | Stream by the bridge S of Ambohitantely | reserve: S-18.2023; E047.2780; 1556 m: | Hygropetric rock and stagnant pools: | 11/04/2018 | Leg. T. Ranarilalatiana -1♀ (NMW): // Data in NHRS | JLKB | 000065756 // RM: Betsiboka Bas (PO533) | Andranofeno Sud Riv. | 47°10'46"E, 18°05'00"S | 06.11.1995 | Leg. Elouard, J.-M., Oliarinony. R. // Toamasina. Alaotra Mangoro: Ambatondrazaka, Andilamena, Moramanga: -1♀ (NMW): // Data in NHRS | JLKB | 000065759 // E-Madagascar (09) Ambaton- | drazaka Region, 5Km N Didy | 1100–1200 m. asl. 14–16.01.1995 | G. Dunay & J. Janák coll. // -1♀ (NMW): // Data in NHRS | JLKB | 000065760 // E-Madagascar (10) | Ambatombe, near Andilamena | 900 m asl. 17.01.1995 | G. Dunay & J. Janák coll. // -1♂ (NMW): // Data in NHRS | JLKB | 000065761 // E-Madagascar (11) Ampamoho | near Andilamena, 1200–1300 m. | asl. 18–20.01.1995 | G. Dunay & J. Janák coll. // -1♂ (GP) (NHRS): // NHRS-JLKB | 000010627 // MAD: TOAM: Alaotra Mangoro | Betsabora river by RN2 near | Antsapanana village, 6 Km W | of Moramanga: MAD14-81: river | with side pools: 18.9247S | 48.1828E; 900 m; 24.XI.2014 // Leg. J. Bergsten, | J.H. Randriamihaja | & T. Ranarilalatiana // -1♂(Alc.) (NHRS): // NHRS-JLKB | 000010833 // Madagascar: Toamasina: Alaotra | Mangoro: Analamazaotra SR: bog | at S border of reserve: S18.95456 E | 048.44048, 910 m: 09.XI.2011: GB | Nets and sieves: bog with red mud: | Field# MAD11-27 // Leg. J. Bergsten, R. | Bukontaite, T. | Ranarilalatiana & | H.J. Randriamihaja // -1♀ (NHMUK): // BMNH-677103 // 11.xii.2004, Andasibe, | Madagascar: P27MD36 | Coll Balke_M; | Monaghan_M // (DNA Voucher) // -1♀ (NMW): // Data in NHRS | JLKB | 000065757 // MADAGASCAR (Md-4) | Andasibe, NP Perinet | 1150 m, Pfütze auf waldwiese | und in Kleinen Bach | 7.12.2000, Leg. W. Dolin // -1♀ (NMW): // Data in NHRS | JLKB | 000065758 // E-Madagascar (07) | Andranokobaka, N Moramanga | 800 m. Asl, 13.01.1995 | G. Dunay & J. Janák coll. // Antsiranana. Sava: Andapa: -1♂ (GP), 1♂ (Alc.) (NHRS): // NHRS-JLKB | 000010827, 10828 (Alc.) // MAD: ANTS: Sava: Anjanaharibe | Sud NP: river Marolakana at the | crossing place: Mad14-64: larger | river with rocks; 14.7623S | 49.4834E: 920 m: 15.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | & T. Ranarilalatiana //Copelatus species. Striae 1, 3, and 5 present only in posterior one third (1) or two thirds ; striae 2, 4, and 6 fragmented anteriorly and, except for stria 4, never reach the base as clearly impressed striae. Penis profile in lateral view is characteristic, resembling a “pumpjack”  that what he had described from Madagascar was not conspecific with Aubé’s C.duodecimstriatus from the Mascarene islands and he gave it the new name C.distinguendus, referring to his description from 1895. Copelatusduodecimstriatus lacks submarginal striae, has a distinctly different penis shape, and is endemic to Mauritius 0149666EE3E15D2FA4E180FD888742E5AgabuspulchellusCopelatusafricanusTL: Botswana, Lake Ngami; Sharp, 1882: 583; Copelatusbasalis Boheman, 1848: 244; TL: South Africa (Caffraria interiore);  ? CopelatusdiscoideusTL: Mesopotamia; Sharp, 1882: 582; CopelatusobtususTL: South Africa ; Boheman, 1848: 242; CopelatusstrigulosusTL: Mesopotamia; Sharp, 1882: 582; CopelatusmimetesTL: Madagascar, Sakaraha, Lambomakandro; syn. nov. Guignot, 1957: 73; Egypt, Sinai.HT♂ (GP) (Copelatus mimetes) : // Data in NHRS | JLKB | 000030032 // Sakaraha | Lambomakandro | III-56 A.R. // Type [red label] // F. Guignot Det., 1956 | Copelatus | mimetes n. sp.| Type ♂ // INSTITUT SCIENTIFIQUE MADAGASCAR // -LT♀ (lectotype here designated) (Copelatus obtusus) (NHRS): // Caffra | ria. // J. Wahlb // Type. // HoloTypus  // obtusus Boh. // Cop. pulchellus var: | obtusus Boh. | J. Balfour-Browne det. // 5657 | E91 // NHRS-JLKB | 000065335 // Lectotype | Copelatus obtusus | Boheman, 1842 | Des. Ranarilalatiana | & Bergsten, 2019// -? ST♂  (NHRS): // Caffra | ria. // J Wahlb // Type. // Typus  // 158 | 61 // Copelatus | basalis Bhn. // Copelatus | pulchellus | Klug. | Det. 19.iv.1961 | J. Omer-Cooper. // 5597 | E91 // NHRS-JLKB | 000065337 // -? ST♀  (NHRS): // Caffra | ria. // J. Wahlb // Paratypus  // 160 | 61 // 5596 | E91 // NHRS-JLKB | 000065338 // -? ST♀  (NHRS): // Caffra | ria. // J. Wahlb // Paratypus  // 159 | 61 // 5595 | E91 // NHRS-JLKB | 000065339 //-Antsiranana. Diana: Antsiranana: -1♂(GP) (MNHN): // Data in NHRS | JLKB | 000030259 // Madagascar | Diego-Suarez | Ch. Alluaud 1893 // Mahajanga. Boeny: Mitsinjo: -1♀ (NHRS): // NHRS-JLKB | 000065733 // Madagascar: Mahajanga: Boeny: | Mahavavy Kinkony RS: S 16.13337 | E 045.95778, 19 m.a.o. 04.XII.2009 | Water Net, Field# MAD09-25 | leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // Mahajanga. Melaky: Antsalova: -1♂(GP) (NHRS): // NHRS-JLKB | 000010695 // Madagascar: Mahajanga: Melaky: | Tsingy de Bemaraha NP: S19.03572 | E044.77507, 66 m.a.o. 15.XII.2009 | Water Net, Field# MAD09-58 | Leg. J. Bergsten, N. Jönsson, T. | Ranarilalatiana, H.J. Randriamihaja //C.marginipennis  and C.mahajanga in overall habitus, but body shape more like the elongated shape of C.mahajanga. Copelatuspulchellus on Madagascar is ferrugineous black in overall colouration with or without a rather narrow or vague testaceous band basally on elytra :Body length 5.5–6.1 mm. Body shape oblong oval, rather convex and attenuate posteriorly, dark brown to blackish ferrugineous. Head infuscated brown ferrugineous, somewhat lighter posteriorly, covered with dense microreticulation and sparser punctation.Pronotum dark brown to ferrugineous black with testaceous anterolateral corners. Disc covered with fine microsculpture forming regular cells and regularily spread small punctures of about same size as cells. Punctuation becomes coarser in posterolateral corners with a weak tendency to corrugate.th the length of elytra); submarginal stria starting at approx. middle.Elytra predominantely dark brown to ferrugineous black on disc and along striae with or without a rather narrow and vague basal testaceous band Fig. . Outer iC.marginipennis and C.mahajanga. Lateral parts of metaventrite medium broad. Metacoxal lines short and rather strongly diverging anteriorly. Antennae, palps, pro- and mesolegs testaceous, metalegs somewhat darker testaceous.Ventral side ferrugineous dark brown, with testaceous spots laterally on abdominal ventrites. Metacoxa and ventrites with strioles. Prosternal process more elongate lanceolate and with blunter apex compared with Female: elytral striolation limited to the medial parts of the outer three elytral intervals in the single female studied from Madagascar. From other parts of the distribution a female form is known that has the entire elytra striolated .C.marginipennis and C.mahajanga.Male: protibia bisinuate and angled at base, distally expanded. Penis thin, strongly angled at middle in lateral view, and apex somewhat twisted to the left in ventral view Fig. . ParamerC.pulchellus is currently interpreted, this is a very widely distributed Afrotropical and Middle Eastern species. Copelatuspulchellus was not previously recorded from Madagascar following the revision by C.mimetes with C.pulchellus, Madagascar now forms part of the distribution. From Madagascar we have seen specimens from Antsiranana, Mahavavy Kinkony Reserve, Tsingy de Bemaraha National Park (Bekopaka), and from Lambomakandro, Sakaraha  and in a muddy stagnant pool in a dried-out river bed. Both localities are in dry deciduous forests of lowland western Madagascar.Copelatuspulchellus forms part of a diverse species group with many externally very similar species. C.pulchellus.Copelatuspulchellus is now interpreted as a widespread Afrotropical and Middle Eastern species with the male penis similar to that illustrated in Figure C.obtusus Boheman and later as C.africanus Sharp . We have studied the same type material as Omer-Cooper and agree with this conclusion. However, we are not convinced that the material housed at NHRS as these types are the correctly identified types of the name C.basalis. Copelatuspulchellus Klug; 260 – Copelatusobtusus; 261 – Copelatusstriatellus; and 262 – Copelatusbasalis. For C.pulchellus the disc of elytra is described as having six striae. For the longer description of C.obtusus this is further elaborated to detail that there are six discal stria but entire elytra has seven striae “septem-striata” . This pattern and number of striae is consistent with the Copelatuspulchellus species group. The following species is correctly identified as C.striatellus with nine discal striae, and is clearly stated as such: “disco striis 9 tenuibus”, of which the innermost is much abbreviated. This description matches very well with the types preserved at NHRS. Finally, C.basalis is described as the last Copelatus species in Boheman’s work and elytra is described as “12-striata”. The supposed types at NHRS for both C.obtusus and C.basalis have six discal and one submarginal striae. The type for C.obtusus matches the original description of “septem-striata” but the three types of C.basalis with the same number of striae does not match the original description of “12-striata”. It would be very inconsistent of Boheman to describe the total number of striae on one elytron in one case “elytra…septem-striata”, and in the other only the discal striae but summing up the number from both elytral halves “elytra…12-striata”. It would also be very illogical to place C.basalis after the 9-striated C.striatellus if it has the same number of striae as C.pulchellus and C.obtusus, which come first. Similarily, C.basalis and listed the name both under the 12-striated C.mocquerysi Régimbart with a question mark, and under C.pulchellus. We consider the status of C.basalis Boheman as uncertain but we have not found any alternative potential type material at NHRS. We designate the single undoubted syntype of C.obtusus Boheman, 1848 in the NHRS collection as lectotype to preserve the stability of the name.Taxon classificationAnimaliaColeopteraDytiscidaeC9FE8EA486D557DFA5B1A38A9F232743Colymbetesmarginipennis Laporte, 1835: 102.CopelatusaldabricusTL: Seychelles, Aldabra Islands. J. Balfour-Browne, 1950: 368 syn. nov.; Copelatusaldabricusvar.simplexTL: Madagascar.Senegal [possibly mislabelled].aldabricus: based on male (holotype), J.C. Fryer collection, collected 1908-9 from Aldabra; of simplex: based on male and female syntypes from Madagascar without further locality data.housed in Buquet collection and originating from Senegal; of LT♂ (lectotype here designated) (Colymbetes marginipennis) (MNHN): // Data in NHRS | JLKB | 000065416 // Copelatus marginipennis Buquet | pulchellus var. Aubé | h. in Senegal D. Buquet // D. Sharp | Monogr. // Ex-Musaeo Déjean // pulchellus // Lectotype | Colymbetes marginipennis | Laporte, 1835 | Des. Ranarilalatiana | & Bergsten, 2019// -HT♂  (NHMUK): // Aldabra, 08-9. J.C.F. Fryer // Perey Sladen Trust | expedition. | 1913-170. // Type [red round label] // Copelatus | aldabricus Type | J. Balfour-Browne det. //-Toamasina. Alaotra Mangoro: Andilamena, Moramanga: -1♀ (NMW): // Data in NHRS | JLKB | 000010726 // Madagascar 17.01.1995 | Ambatombe | nr. Andilamena 900 m | leg. Dunay & Janák (10) // -1♀ (NHMUK): // BMNH-797894 // MAD: AMPA: Moramanga: Andasibe | Andasibe NP: Big Pond | P61BI15: N: -18.937: E:48.416: 940 m | 06/I/2007: Leg. Isambert et al. // DNA Voucher | BMNH <797894> | MSL294:B10 // -2♂ (GP), 15 ex. (Alc.) : // NHRS-JLKB | 000010587–8, 10799(Alc.) // Madagascar: Tamatave: Alaotra Mangoro: | Analamazaotra RS; Bas fond, non-permanent | pond near trail to “lac rouge”MAD15-1| 943 m, 18°56'26.7"S, 048°25'03.9"E, 16.III.2015 | Among vegetation and dead leaves in the pond, | Leg. T. Ranarilalatiana & H.J. Randriamihaja // -2♂ (GP), 2♀ (Alc.) (NHRS): // NHRS-JLKB | 000010604–5, 10800(Alc.) // Madagascar: Tamatave: Alaotra Mangoro: Mantadia NP. | Non-permanent pond at PK18,50 m; E of Park road | 973 m, 18°46'09.9"S, 048°26'10.4"E, 17.III.2015 | Under dead leaves & vegetation at the edge of the pond, | Leg.T. Ranarilalatiana, H.J. Randriamihaja; MAD15-5 // -1♂ (GP) (NHRS): // NHRS-JLKB | 000010815 // Madagascar: Toamasina: Alaotra | Mangoro: RN2, Mangoro river | 10Km W of Moramanga: S18.92438 | E048.18273, 940 m. 06.XI.2011 | GB Nets and sieves: river and | pools: Field# MAD11-21 // Leg. J. Bergsten, R. | Bukontaite, T. | Ranarilalatiana & | J.H. Randriamihaja // -1♀ (Alc.) (NHRS): // NHRS-JLKB | 000010822 // Madagascar: Toamasina: Alaotra | Mangoro: Analamazaotra SR: | close to park entrance: S18.9355 E | 048.41656, 970 m: 08.XI.2011: GB | Nets and sieves: dried up forest | pond: Field# MAD11-25 // Leg. J. Bergsten, R. | Bukontaite, T. | Ranarilalatiana & | H.J. Randriamihaja // -6 ex. (Alc.) (NHRS): // NHRS-JLKB | 000010823 // Madagascar: Toamasina: Alaotra | Mangoro: by RN2 S border of | Analamazaotra reserve 1Km E | Antsampanana: S18.94987 E048.42331| 980m: 09.XI.2011: GB Nets & sieves | ditch next to road: Field# MAD11-29 // Leg. J. Bergsten, R. | Bukontaite, T. | Ranarilalatiana & | J.H. Randriamihaja // Toamasina. Analanjirofo: Maroantsetra: -3♂ (GP), 2♀ (NHMUK): // BMNH-797906–10 // MAD: TOAM: Maroantsetra: Masoala | Masoala NP: Pool | P58BI14: N: -15.758: E: 49.993: 10 m| 17/XI/2006: Leg. Isambert et al. // BMNH (DNA Voucher) // -1♂ (GP) (NHRS): // NHRS-JLKB | 000010794 // MAD: TOAM: Maroantsetra: Masoala | Masoala NP: Pool | P58BI14: N: -15.758: E: 49.993: 10 m| 17/XI/2006: Leg. Isambert et al. // -1♂ (NMW): // Data in NHRS | JLKB | 000010728 // E-Madagascar: Fenerive | Foret de Tampolo | 28.12.1998 | leg. J. Moravec // -1♀(Alc.) (NHRS): // NHRS-JLKB | 000011122 // Madagascar: Toamasina: Analajinrofo: | Masoala NP: degraded lowalt. forest: | MAD18-47: small waterpools on path | ~0.5km NW of Marofototra village, | 15.7606S, 49.9926E, 15 m, 17.II.2018 | Leg. T. Ranarilalatiana // Toliara. Menabe: Morondava: -3♂ (GP) 2♀ (Alc.) (NHRS): // NHRS-JLKB | 000010580, 10730 (JB197), 10751, 10802(Alc.) //Madagascar: Toliara: Menabe: | Kirindy RS: S20.07430 | E044.66307, 52 m.a.o. 12.XII.2009 | Water Net, Field# MAD09-46 | Leg: J. Bergsten, N. Jönsson, T. | Ranarilalatiana, H.J. Randriamihaja // Mahajanga. Boeny: Ambato-Boeny, Mitsinjo: -1♂ (NHRS): // NHRS-JLKB | 000010734 (JB201) // Madagascar: Mahajanga: Boeny: | Ankarafantsika NP. S16.30350 | E046.81068, 87 m.a.o. 29.XI.2009 | Water Net, Field# MAD09-03 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // -6♂ (GP), 3♀, 11 ex. (Alc.) : // NHRS-JLKB | 000010567–71, 10729 (JB196), 10732 (JB199), 10742–3, 10798(Alc.) // Madagascar: Mahajanga: Boeny: | Ankarafantsika NP. S16.30341 | E046.81073, 74 m.a.o. 29.XI.2009 | 22W Black Light, Field# MAD09-07 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // -1♂ (NHRS): // NHRS-JLKB | 000010731 (JB198) // Madagascar: Mahajanga: Boeny | Ankarafantsika NP, S16.30270 | E046.80996; 75 m.a.o. 30.XI.2009 | 22W Black light, Field# MAD09-13 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // -1♀ (NHRS): // NHRS-JLKB | 000010737 (JB191) // Madagascar: Mahajanga: Boeny: | Ankarafantsika NP. S16.31418 | E046.81731, 30.XI.2009 | Hand picking, Field# MAD09-14 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // -3♀, 1♂(GP), 11 ex. (Alc.) : // NHRS-JLKB | 000065749 (JB809), 10738 (JB192), 10744–5, 10806(Alc.) // Madagascar: Mahajanga: Boeny: | Mahavavy Kinkony RS. S16.14653 | E045.94926, 9 m.a.o. 04.XII.2009 | Water net, Field# MAD09-24 | Leg. J. Bergsten, N. Jönsson, T. | Ranarilalatiana, H.J. Randriamihaja // -2♂ (GP), 2♀, 18 ex. (Alc.) : // NHRS-JLKB | 000010578–9, 10739 (JB193), 10746, 10807(Alc.) // Madagascar: Mahajanga: Boeny: | Mahavavy Kinkony RS. S16.13337 | E045.95778, 19 m.a.o. 04.XII.2009 | Water net, Field# MAD09-25 | Leg. J. Bergsten, N. Jönsson | T. Ranarilalatiana, H.J. Randriamihaja // -5♂ (GP), 4♀, 79 ex. (Alc.) : // NHRS-JLKB | 000010572–7, 10740 (JB194), 10747–8, 10808 (Alc.) // Madagascar: Mahajanga: Boeny: | Mahavavy Kinkony RS. S16.05776 | E045.80585, 22 m.a.o. 05.XII.2009 | Water net, Field# MAD09-28 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // -1♂, 1♀, 4 ex. (Alc.) (NHRS): // NHRS-JLKB | 000010735 (JB202), 10749, 10809(Alc.) // Madagascar: Mahajanga: Boeny: | Mahavavy Kinkony RS. S16.06651 | E045.77672, 24 m.a.o. 05.XII.2009 | Water net, Field# MAD09-29 | Leg. J. Bergsten, N. Jönsson, T. | Ranarilalatiana, H.J. Randriamihaja // -1♀ (NHRS): // NHRS-JLKB | 000010741(JB189) // Madagascar: Mahajanga: Boeny | Mahavavy Kinkony RS, 16.05648S | 045.76371E; 55 m.a.o. 05.XII.2009 | Water net, Field# MAD09-30 | Leg: J. Bergsten, N. Jonsson, T. | Ranarilalatiana, H.J. Randriamihaja // -1♂, 6ex. (Alc.) (NHRS): // NHRS-JLKB | 000010750, 10810(Alc.) // Madagascar: Mahajanga: Boeny: | Mahavavy Kinkony RS. S16.01334 | E046.00376, 24 m.a.o. 06.XII.2009 | Water net, Field# MAD09-33 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // Mahajanga. Melaky: Antsalova: -2♂ (GP) (CAS): // CASENT-8135015–6 // Madagascar: Mahajanga | Prov. Parc National Tsingy | de Bemaraha, 2.5 Km 62° ENE | Bekopaka, Ankidrodroa river | elev 100 m: 11–15 Nov 2001 // 19°7'56"S, 44°48'53"E | Coll: Fisher, Griswold et al. | California Acad. of Sciences | sifted litter - tropical dry forest | on Tsingy, code: BLF4340 // -1♂ (GP) (CAS): // CASENT-8131891 // Madagascar: Mahajanga | Prov. Parc National Tsingy | de Bemaraha, 2.5 Km 62° ENE | Bekopaka, Ankidrodroa river | elev 100 m: 11–15 Nov 2001 // 19°7'56"S, 44°48'53"E | Coll: Fisher, Griswold et al. | California Acad. of Sciences | at light- tropical dry forest | on Tsingy, code: BLF4343 // -1♂ (GP) (CAS): // CASENT-8135006 // Madagascar Mahajanga | Prov. Foret de Tsimembo | 11.0Km 346° NNW Soatana | elev 50 m: 21–25 Nov 2001 | 18°59'43"S, 44°26'37"E // Coll: Fisher, Griswold et al. | California Acad. of Sciences | sifted litter  | in tropical dry forest | coll. Code: BLF4508 // -1♂ (GP), 2♀, 3 ex. (Alc.), 2♂(Alc.) (NHRS): // NHRS-JLKB | 000010581–2, 10752, 10803(Alc.), 10821(Alc.)// Madagascar: Mahajanga: Melaky: | Tsingy de Bemaraha NP: S19.03572 | E044.77507, 66 m.a.o. 15.XII.2009 | Water Net, Field# MAD09-58 | Leg: J. Bergsten, N. Jönsson, T. | Ranarilalatiana, H.J. Randriamihaja // -3♂ GP, 1♀, 5 ex. (Alc.) (NHRS): // NHRS-JLKB | 000010583–4, 10736 (JB203), 10754, 10805(Alc.) // Madagascar: Mahajanga: Melaky: | Tsingy de Bemaraha NP: S18.75643 | E044.71398, 119 m.a.o. 17.XII.2009 | Water Net, Field# MAD09-65 | Leg: J. Bergsten, N. Jönsson, T. | Ranarilalatiana, H.J. Randriamihaja // -1♀, 1♂ (GP), 2♀ (Alc.) (NHRS): // NHRS-JLKB | 000010733 (JB200), 10753, 10804(Alc.) // Madagascar: Mahajanga: Melaky: | Tsingy de Bemaraha NP: 19.03419S | 044.77499E, 41 m.a.o. 15.XII.2009 | Water Net, Field# MAD09-59 | Leg: J. Bergsten, N. Jönsson, T. | Ranarilalatiana, H.J. Randriamihaja // Antsiranana. Diana: Ambanja, Ambilobe, Antsiranana II: -1♂ (GP) (NHRS): // NHRS-JLKB | 000010585 // Madagascar: Antsiranana: Diana: | Ambilomagodra: Stream under the | bridge of the road RN6 in | Ambilomagodra village, S13.00780 | E49.13313, 139 m, 30.XI.2012, GB nets: | dried up stream with some pools in | the village: Field# MAD12-33 // Leg. J. Bergsten, R. | Bukontaite, T | Ranarilalatiana & | J.H. Randriamihaja // -1♂ (GP), 2♀ (Alc.) (NHRS): // NHRS-JLKB | 000010586, 10801(Alc.) // Madagascar: Antsiranana: Diana: | Andrafiabe: Antsoha stream | 200m from Andrafiabe, 12.93022S | 49.03466E, 32 m, 01.XII.2012, GB | nets: stream with some pools: | Field# MAD12-39 // Leg. J. Bergsten, R. | Bukontaite, T | Ranarilalatiana & | J.H. Randriamihaja // -1♂ (Alc.) (NHRS): // NHRS-JLKB | 000011123 // Madagascar: Antsiranana: Diana: | Antsaba: 1km W of Antsaba, | 13.63474S; 48.72918E, 67 m, | 28.XI.2012, GB nets: forest | stream: Field# MAD12-32 // Leg. J. Bergsten, R. | Bukontaite, T | Ranarilalatiana & | J.H. Randriamihaja // -1♀ (NMW): // Data in NHRS | JLKB | 000010725 // MADAGASCAR: Sakaramy |  | lake Farihy Makery | 29.03.2011 | leg. R. Gerecke (MD189) // riparian area near outflow | 12°26'20.5"S, 49°14'20.0"E | 377 m. 29.5 °C // -1♂ (NMW): // Data in NHRS | JLKB | 000010727 // N. Madagascar | Antseranana distr. | Sambirano riv. | Marovato vill. | 5–12.12.01, leg. J. Horák // Antsiranana. Sava: Sambava, Vohemar: -1♀ (NHMUK): // BMNH-797876 // MAD: DIEG: Sambava: Marojejy | Marojejy NP: Pool | P57BI31: N: -14.457: E:49.79: 162m | 10/XI/2006; Leg. Isambert et al // DNA Voucher | BMNH <797876> | MSL294:A4 // -1♀ (CAS): // CASENT-8135007 // Madagascar Antsiranana | Foret d’Ampondrabe. | 26.3Km 10° NNE Daraina | elev 175m: 10 December 2003 // 12°58'12"S, 049°42'00"E | California Acad. of Sciences | Coll: B.L.Fisher, sifted | litter  | tropical dry forest, BLF9974 //Copelatusmarginipennis is distinguished from all other Malagasy Copelatus except C.pulchellus and C.mahajanga by the presence of six discal elytral striae and a broadly oval body shape. Copelatusmarginipennis is most easily separated by the distinct shape of the male penis in lateral view  lacks a transverse testaceous band basally on the elytra and females of C.marginipennis can usually be distinguished from C.mahajanga by the normally narrower testaceous band, a longer first elytral stria, and a broader body shape.iew Fig. . Copelatth posterior of elytral base, second to sixth full length, and submarginal stria starting at approx. the middle; all striae approaching the apex of elytron except the sixth, which is abbreviated posteriorly and a little shorter. Sometimes, the first and second elytral striae, like fifth and sixth, unite posteriorly. Elytral surface covered with dense punctures.Body length 5.2–6.6 mm. Body shape oval, rather convex and attenuate posteriorly, dark brown to brown ferrugineous. Head, pronotum and elytra in the same dark brown ferrugineous, covered with fine dense punctation. Lateral sides of pronotum more brownish, with short strioles and the widest striolate area in the posterior corners. Elytra dark brown to brown ferrugineous, with a testaceous transverse band at base Fig. . Six cleVentral side brownish to ferrugineous, metacoxa with microsculpture, densely and finely punctate. Metacoxa and abdominal ventrites striolate. Prosternal process rather short and spear-shaped, medially only weakly raised and rounded. Lateral parts of metaventrites rather broad. Metacoxal lines short and rather strongly diverging anteriorly. Antennae and palps both in the same brown colour. Pro- and mesothoracic legs brown ferrugineous. Metathoracic legs dark brown ferrugineous.Male: Protibia strongly angled basally and expanded in apical two thirds. Penis in ventral view with a small preapical tooth on right side; in lateral view very characteristic with a subbasal dorsal knob, and post-middle with a deep ventral invagination Fig. .Females from Madagascar usually with elytral striolation rather weak and restricted to outer intervals, but rarely the elytra are entirely and distincly striolate. Females on average smaller than males.Endemic to the western Indian Ocean islands as far as is modernly known, but the 1835 type locality of the original description and labels read “Senegal”, which indicates either mislabeling or that the species in fact also occurs on continental Africa. Known from Madagascar, Reunion, Comores, Seychelles, and Aldabra Island . WidesprC.mahajanga in Betsabora river near Moramanga and at a 22W black light trap by a forest pool in Ankarafantsika NP.This is a lowland species that seems to be most common in the deciduous western parts of Madagascar. It was common in the newly designated protected areas of Mahavavy Kinkony when we visited it in 2009. In the deciduous forest biome it has been recorded from Kirindy in the south to Ankarana NP in the north. The species seems to be a generalist and can as well show up on lowland humid east coast and at midaltitudes up to at least 1000 m. It is an apt flier collected with light traps and often in very temporary and small shallow pools including water-filled wheel tracks. Found in all kinds of temporary pools, as well as in streams and in residual pools in dried-up riverbeds. It occurred sympatrically with Copelatusaldabricus was described by J. simplex, and again in 1961 as var. aequabilis (non-available infrasubspecific name) from Andranofotsy (E of Maroantsetra NE Madagascar), and stated (1961) that females of C.aldabricus are normally striolated, but that on Madagascar the non-striolated var. aequabilis is most common or even ubiquitous. We have found at least one female on Madagascar  with the elytra entirely striolated  has been treated as a junior synonym of Copelatuspulchellus Klug, 1834 since C.marginipennis in MNHN, Paris. This could be identified based on a folded blue label bearing the information “Copelatusmarginipennis Buquet, pulchellus var Aubé, h. in Senegal, D. Buquet”, which fits perfectly the description by C.aldabricus and not of C.pulchellus. The name marginipennis has to our knowledge not been used as a valid name after 1899, fulfilling the first of the two criteria for protecting a younger currently used synonym from an older available name (ICZN 23.9.1.1). However, Copelatusaldabricus is a relatively recent name from a region with a low-intensity taxonomic research and we do not believe it fulfils the second criteria of at least 25 papers published by at least ten authors in the immediately preceding 50 years and encompassing a span of not less than ten years (ICZN 23.9.1.2). Therefore it is not justifiable to protect the younger synonym C.aldabricus and the older C.marginipennis is therefore brought back as the valid name of this species. We designate the discovered syntype of Colymbetesmarginipennis Laporte, 1835 in the MNHN collection as lectotype to preserve stability of the name. At the same time as this type specimen was discovered, also a likely “type” specimen of the nomen nudum Copelatusfimbriolatus Dejean, 1837 from “Ile de France” [= Mauritius] was found with the same type of blue folded label (see supplementary file 1: NHRS-JLKB000065420). This nomen nudum is currently considered a synonym of C.pulchellus Klug (Nilsson and Hajek 2018). Unfortunately, it is a female specimen, which is why its identity cannot be established with certainty, but it is more likely identical with C.marginipennis since C.pulchellus is not yet known from any of the Mascarene Islands , 3 male and 4 female (paratypes). I. Janis 1–10 December 1996. Mahajanga. Boeny: Mahajanga I/Mahajanga II [district cannot be verified based on original description or labels]: -HT♂(GP) (NMPC): // MADAGASCAR 1996, Mahajanga Distr., Mahajanga env., Dec. 1–10., I. Janiš leg. // HOLOTYPE, COPELATUS, mahajanga sp. nov., F. Pederzani & J. Hájek det.2005 //Mahajanga. Boeny: Ambato-Boeny, Mitsinjo, Soalala: -3♂ (GP), 1♀ (NMW): // Data in NHRS | JLKB | 000010719–22 // RM: Betsiboka Bas (PO124) | Loc. Ambohimanatrika | Kamoro Riv. // 47°10'06", 16°28'55" | alt. 40 m 01.04.1993 // Leg. ORSTOM ANTANANARIVO // -1♂ (GP) (NHRS): // NHRS-JLKB | 000010717 // Madagascar: Mahajanga: Boeny: | Ankarafantsika NP. S16.30341 | E046.81073, 74 m.a.o. 29.XI.2009 | 22W Black Light, Field# MAD09-07 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // -2♀ (NHRS): // NHRS-JLKB | 000065747 (JB802), 65732 // Madagascar: Mahajanga: Boeny: | Mahavavy Kinkony RS. S16.13337 | E045.95778, 19 m.a.o. 04.XII.2009 | Water net, Field# MAD09-25 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // -2♀ (NHRS): // NHRS-JLKB | 000010724 (JB195), 65748 // Madagascar: Mahajanga: Boeny: | Mahavavy Kinkony RS. S16.01334 | E046.00376, 24 m.a.o. 06.XII.2009 | Water net, Field# MAD09-33 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // -2♂ (GP), 3♀ (CAS): // CASENT-8131811–5 // Madagascar: Mahajanga | Province, Parc National de | Namoroka 16.9 km 317° NW | Vilanandro 12–16 Nov 2002 // 16°24'24"S, 045°18'36"E // Colls. Fisher, Griswold et al.| California Acad. of Sciences | sifted litter,  in tropical dry forest | elev 100 m code: BLF6582 // -1♀ (CAS): // CASENT-8135013 // Madagascar: Mahajanga | Prov. Parc National de | Namoroka 16.9 Km 317° NW | Vilanandro elev 100 m | 12–16 November 2002 // 16°24'24"S, 045°18'36"E | Coll. Fisher, Griswold et al. | California Acad. of Sciences | malaise trap- | tropical dry forest | collection code BLF6581 // -1♂ (GP) (CAS): // CASENT-8135001 // Madagascar: Mahajanga | Province Parc National de | Namoroka | 17.8 Km 329° WNW Vilanandro | elev 100 m: 08–12 Nov 2002 // 16°22'36"S, 045°19'36"E | Colls. Fisher, Griswold et al. | California Acad. of Sciences | pitfall trap- in tropical dry forest, | collection code BLF6506 // Mahajanga. Melaky: Antsalova: -1♂ (GP) (NHRS): // NHRS-JLKB | 000010723 (JB190) // Madagascar: Mahajanga: Melaky: | Tsingy de Bemaraha NP: S19.03572 | E044.77507, 66 m.a.o. 15.XII.2009 | Water Net, Field# MAD09-58 | Leg. J. Bergsten, N. Jönsson, T. | Ranarilalatiana, H.J. Randriamihaja // Toamasina. Alaotra Mangoro: Moramanga: -5♂ (GP), 3♀, 11 ex. (Alc) : // NHRS-JLKB | 000010554–5, 10596–601, 10795 (Alc.) // MAD: TOAM: Alaotra Mangoro: | Betsabora river by RN2 near | Antasmpanana village, 6Km W | of Moramanga: MAD14-81: river | with side pools: 18.9247S | 48.1828E: 900 m: 24.XI.2014 // Leg. J. Bergsten, | J.H. Randriamihaja | & T. Ranarilalatiana // -3♂ (GP) 26 ex. (Alc.), 7 ex. (Alc.) : // NHRS-JLKB | 000010602–3, 10825, 10796(Alc.), 10826(Alc.) // Madagascar: Toamasina: Alaotra | Mangoro: RN2, Betsabora river | 10Km W of Moramanga: S18.92438 | E048.18273, 940 m. 06.XI.2011: | GB Nets and sieves: river and | pools: Field# MAD11-21 // Leg. J. Bergsten, R. | Bukontaite, T. | Ranarilalatiana & | H.J. Randriamihaja //C.marginipennis in habitus appearance but body shape more elongate. On average C.mahajanga have a broader transverse testaceous band at base of elytra  and was previously only known from the type series and lowland west type locality “Mahajanga env.” without further details. The discovery of the species in Betsabora River at mid-altitude in the east indicates that the species may have a much wider distribution and ecological niche. As the species is very similar to C.marginipennis habitually, it might be misidentified as such in collections if male genitalia were not examined.Taxon classificationAnimaliaColeopteraDytiscidaeGuignot, 1941573F0C84F70555E7A4F6FC4974616368Copelatusinsuetus Guignot, 1941: 39.Madagascar, Perinet [= Analamazaotra NP].Based on single male specimen (holotype). Madagascar, Perinet.Toamasina. Alaotra Mangoro: Moramanga: -HT♂ (GP) : Data in NHRS | JLKB | 000030227 // Madagascar | Perinet // Type [red label] // [male symbol] // Copelatus | insuetus | guign. Type //Toamasina. Alaotra Mangoro: Moramanga, Ambatondrazaka, Andilamena: -2♀ : Data in NHRS | JLKB | 000030228–9 // Perinet // INSTITUT | SCIENTIFIQUE | MADAGASCAR // [female symbol] // -5♀ (NMW): Data in NHRS | JLKB | 000030302–6 // Madagascar (Md 4/5) | Andasibe NP, Perinet | 1250 m Pfütze im Urwald | 5.12.2000, leg. W. Dolin // -1♂ : Data in NHRS | JLKB | 000030312 // Madagascar 10.1.2007 | Andasibe NP. | Lokato-near Andasibe | Mracek Z. leg. // coll. Jiri HÁJEK | National Museum | Prague, Czech Republic // -1♀, 1♂ (NMW): Data in NHRS | JLKB | 000030307–8 // E-Madagascar (10) | Ambatombe near Andilamena | 900 m asl. 17.01.1995 | G. Dunay & J. Janák coll. -1♀, 1♂ (NMW): Data in NHRS | JLKB | 000030309–10 // E-Madagascar (11) Ampamoho | near Andilamena, 1200–1300 m. | asl. 18–20.01.1995 | G. Dunay & J. Janák coll. // -1♀ (NMW): Data in NHRS | JLKB | 000030311 // Madagascar 18–20.1.1995 | 5km S Ampamoho | nr. Andilamena, 950–1000 m | Dunay & Janák (11) // -1♀ (NHMUK): // BMNH-797884 // MAD: AMPA: Moramanga: Andasibe | Andasibe NP: Chanel| P61BI02: N: -18.935: E:48.418: 944 m | 04/I/2007 Leg. Isambert et al // DNA Voucher | BMNH <797884> | MSL294:A12 // -2♀ (NHRS): // NHRS-JLKB | 000010692–3 // Madagascar: Tamatave: Alaotra Mangoro: | Analamazaotra RS; Bas fond, non-permanent | pond near trail to “lac rouge”MAD15-1 | 943 m, 18°56'26.7"S, 048°25'03.9"E, 16.III.2015 | Among vegetation and dead leaves in the pond | Leg.T. Ranarilalatiana & H.J. Randriamihaja // -1♀, 2♀ (Alc.) (NHRS): // NHRS-JLKB | 000065745, 10783 (Alc.) // Madagascar: Toamasina: Alaotra | Mangoro: Analamazaotra RS: close to | park entrance: 18.93573S; 048.41741E | 930 m. 08.XI.2011 GB Nets and sieves: | stagnant pools in tributary to | Analamazaotra river; Field# MAD11-26 // Leg. J. Bergsten, R. | Bukontaite, T. | Ranarilalatiana & | H.J. Randriamihaja // -1♂ (GP), 1♀, 3 ex.  (Alc.) (NHRS): // NHRS-JLKB | 000010839, 65702, 10840 (Alc.) // MAD: TOAM: Alaotra Mangoro | Andasibe Mantadia NP, Analamazaotra | 250m E of park entrance: Mad14-18: | dried up river bed with stagnant pools: | 18.9357S 48.4174E: 930 m: 27.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | & T. Ranarilalatiana // -1♀ (NHRS): // NHRS-JLKB | 000010841 // MAD: TOAM: Alaotra Mangoro | Andasibe Mantadia NP, Analamazaotra | pond: 100m E from Lac vert: Mad14-15: | shallow forest pool: 18.9385S 48.4219E: | 940 m: 27.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | & T. Ranarilalatiana // -1♂ (GP), 1♀, 1♀ (Alc.) (NHRS): // NHRS-JLKB | 000065705, 65777, 10844(Alc.) // MAD: TOAM: Alaotra Mangoro | Andasibe Mantadia NP, Analamazaotra | 150 m E of park entrance: Mad14-14: | shallow partly dried out forest pond: | 18.9355S 48.4166E: 930 m: 27.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | & T. Ranarilalatiana // -1♂ (GP), 34 ex. (Alc.) : // NHRS-JLKB | 000010842, 10843(Alc.) // MAD: TOAM: Alaotra Mangoro | Andasibe Mantadia NP, | Mantadia: 12Km N of park | entrance: Mad14-83: | forest pond with some | vegetation: 18.7911S | 48.4259E: 960 m: 28.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | & T. Ranarilalatiana // -6♂ (GP), 4♀, 33 ex. (Alc.) : // NHRS-JLKB | 000010534–7, 10606–11, 10780(Alc.) // MAD: TOAM: Alaotra Mangoro: | Betsabora river by RN2 near | Antasmpanana village, 6Km W | of Moramanga: MAD14-81: river | with side pools: 18.9247S | 48.1828E: 900 m: 24.XI.2014 // Leg. J. Bergsten, | J.H. Randriamihaja | & T. Ranarilalatiana // -1♀ (NHMUK): // BMNH-797895 // MAD: TOAM: Ambatondrazaka | Zahamena: Zahamena NP: Stream | P60BI15:N: -17.52: E:48.721: 1075 m| 31:XI:2006: Leg. Isambert et al // DNA Voucher | BMNH <797895> | MSL294:B11 // -4♂ (GP), 1♀ 11 ex. (Alc.) : // NHRS-JLKB | 000010817, 10872–4, 65700, 10880(Alc.) // Madagascar: Toamasina: Alaotra Mangoro: | Zahamena NP: Sect. Antanandava: | path towards Camp Cascade: S17.5166; | E048.7227; 1040 m. 07.III.2018; GB Nets, | white pan & sieves: Waterfilled goldigging | hole in forestswamp: Field# MAD18-80 | Leg. J. Bergsten, & T. Ranarilalatiana // -1♀ (Alc.) (NHRS): // NHRS-JLKB | 000010885 // Madagascar: Toamasina: Ambatondrazaka | Zahamena NP: the way to Camp | Cascade: 19.III.2013, GB Nets & sieves: slow | stream, Leg. J.H. Randriamihaja & | T. Ranarialalatiana: Field# ZAH13-02 // -1♀ (Alc.) (NHRS): // NHRS-JLKB | 000010881 // Madagascar: Toamasina: Alaotra Mangoro: | Zahamena NP: Sect. Antanandava: | Analamaitso forest: S17.5179; E048.7232; | 1050 m. 07.III.2018; GB Nets, white pan & | sieves: small foreststream: Field# MAD18-81 | Leg. J. Bergsten, & T. Ranarilalatiana // -3♀ (Alc.) (NHRS): // NHRS-JLKB | 000010882 // Madagascar: Toamasina: Alaotra Mangoro: | Zahamena NP: Sect. Antanandava: | Ambavahala forest: S17.5300; E048.7161; | 1070 m. 08.III.2018; GB Nets, white pan & | sieves: larger shaded forestswamp: Field# MAD18-90 | Leg. J. Bergsten, & T. Ranarilalatiana // -1♂ (GP), 2♀ (Alc.) (NHRS): // NHRS-JLKB | 000010875, 10883(Alc.) // Madagascar: Toamasina: Alaotra Mangoro: | Zahamena NP: Sect. Antanandava: | Pandanus Swamp: S17.5166; E048.7227; | 1040 m. 08.III.2018; GB Nets, white pan & | sieves: forestswamp: Field# MAD18-91 | Leg. J. Bergsten, & T. Ranarilalatiana -1♀ (Alc.) (NHRS): // NHRS-JLKB | 000010884 // Madagascar: Toamasina: Alaotra Mangoro: | Zahamena NP: Sect. Antanandava: | Analamaitso forest: S17.5225; E048.7250; | 1090 m. 08.III.2018; GB Nets, white pan & | sieves: waterfilled golddigging hole in flat: Field# MAD18-92 | Leg. J. Bergsten, & T. Ranarilalatiana // -2♂ (GP), 3♀, 6♀ (Alc.), 6ex.  (Alc.), : // NHRS-JLKB | 000010816, 10876–9, 10886(Alc.), 65753(Alc.) // Madagascar: Toamasina: Alaotra Mangoro: | Zahamena NP: Sect. Antanandava: 150 m | upstream to Camp Cascade: S17.5458; E048.7244; | 1270 m. 10–11.III.2018; GB Nets, white pan & | sieves: waterpool with dead leaves: Field# MAD18-109, | Leg. J. Bergsten, & T. Ranarilalatiana // Toamasina. Analanjirofo: Maroantsetra: -1♂, 1♀ : Data in NHRS | JLKB | 000030314-5 // Maroantsetra | Vadon IV.50 // 1/6 //? insuetus | guign. //  Fianarantsoa. Vatovavy Fitovinany: Ifanadiana: -1♀ (CAS): // CASENT-8131927 // Madagascar: Province | Fianarantsoa, Parc National | Ranomafana, radio tower | at forest edge, elev 1130 m | 19–26 February 2002 // 21°15.05'S, 47°24.43' | coll: M. Irwin, R. Harin’Hala | California Acad. of Sciences | malaise. Mixed tropical | forest MA-02-09B-17 // -1♂  : Data in NHRS | JLKB | 000030313 // Madagascar 26–31.I.2007 | Ranomafana NP | Ranomafana vill. env. | Z. Mracek leg. // coll. Jiri HÁJEK | National Museum | Prague, Czech Republic // Fianarantsoa. Matsiatra Ambony: Lalangina: -1♀ (CAS): // CASENT-5004000 // Madagascar, Fianarantsoa | Province, Ranomafana National | Park, Vohiparara village, 1160m | mixed tropical forest | 2–22 January 2001 | 21.23906S / 47.38487E | COL-DHK-2001–003 // D.H. & K.M. Kavanagh | R.L. Brett, E. Elsom, F. Vargas, | R. Ranaivosolo, E.F. Randrianirina, | N. Rasoamanana, T.J. | Ravelomanana and H.C. | Raveloson collectors // -4♂ (CAS): // CASENT-5004056, 5004070–1, 5004075 // Madagascar, Fianarantsoa | Province, Ranomafana | National Park, Vohiparara | area, 1170m mixed tropical | forest 2–22 January 2001 | 21.22644S, 47.36979E, | Stop# DHK-01-004 // D.H. & K.M. Kavanagh | R.L. Brett, E. Elsom, F. | Vargas, R. Ranaivosolo, | E.F. Randrianirina, N. | Rasoamanana, T.J. | Ravelomanana and H.C. | Raveloson collectors // Antsiranana. Diana: Nosy-Be: -1♂(GP) (MNHN): // Data in NHRS | JLKB | 000030230 // Nosy-Be | Sokobe [=Lokobe NP] | V. 57 Hoyt Coll // ♂ [male symbol] // Mahajanga. Boeny: Ambato-Boeny: -1♀ (NHRS): // NHRS-JLKB | 000010694 (JB206) // Madagascar: Mahajanga: Boeny: | Ankarafantsika NP. S16.30341 | E046.81073, 74 m.a.o. 29.XI.2009 | 22W Black Light, Field# MAD09-07 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja // -1♀ (NHRS): // NHRS-JLKB | 000010781 (JB205) // Madagascar: Mahajanga: Boeny: | Ankarafantsika NP. S16.30350 | E046.81068, 87 m.a.o. 29.XI.2009 | Water Net, Field# MAD09-03 | Leg. J. Bergsten, N. Jönsson, | T. Ranarilalatiana, H.J. Randriamihaja //Copelatusinsuetus can be distinguished from the following species based on the penis ventral outline in lateral view with a distinct “shoulder”  and also spots laterally on abdominal sternites II–VII and along some sternite margins are lighter. Metacoxa and abdominal sternites II–IV marked with strioles. Prosternal process short, broad, with blunt apex. Lateral parts of metaventrite broad. Metacoxal lines long, abbreviated only slightly before metaventral margin, diverging anteriorly.Male: first three pro- and mesotarsomeres widened and ventrally equipped with suction cups; number of suction cups per segment for I–III: 7:4:4 for both pro- and mesotarsus. Protibia modified, narrow with a bisinuate angulate ventral margin at base, broadened distally. Pro- and mesotarsal claws unmodified. Penis in ventral view thin and simple, apical part slightly left-turned. Penis in lateral view angled after basal third forming a “shoulder”. Apex in left lateral view with a characteristic dorsal ridge crossing posterior dorsal margin Fig. . Penis sFemale: anterior half of elytra finely striolated from second or third elytral interval to external margin. Degree of elytral striolation in females quite variable between specimens. These strioles are finer than the strioles on pronotum found in both sexes.NHRS) from Ankarafantsika NP is likely this species.Endemic to Madagascar. Known with certainty from the eastern escarpment, Ranomafana NP in the south, Analamazaotra NP, and Mantadia NP in the central region, and Zahamena NP and Andilamena further north Fig. . We haveWe have collected this species in Analamazaotra NP in stagnant forest pools with vegetation or with plentiful of dead leaves. Also found in a sidepool next to a river in semi-open degraded area near Moramanga. The eastern escarpment localities range in altitude between 900–1300 m, but if the localities of Nosy Be and Maroantsetra are correct this species can also occur at lowland sea level altitudes. It seems to be most abundant in tropical eastern forests, but if the Ankarafantsika females belong to this species, then it can also occur in western deciduous forests.C.insuetus should be regarded with caution in light of the habitually very similar new species described below.Eventual older records of Taxon classificationAnimaliaColeopteraDytiscidaeRanarilalatiana & Bergstensp. nov.9FD551E5A9DF5CB19C529BDE43F5F8D0http://zoobank.org/3AFB0F44-539F-4621-9002-F2E44EEDC73AAmbohidray reserve, Andriambe  .Toamasina. Alaotra Mangoro: Moramanga: -HT♂(GP) (NHRS): // NHRS-JLKB | 000010890 // Madagascar: Moramanga: Ambohidray | reserve: TR18L14: Andriambe stream: | S-18.61317; E48.32593; 1044m: | stagnant pool in pitfall holes: | 23/05/2018; Leg. T. Ranarialalatiana // Holotype | Copelatus kely sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // Paratypes: -2♂(GP), 3♀, 11 ex.  (Alc.) : // NHRS-JLKB | 000010891, 65741–4, 10889(Alc.) // Madagascar: Moramanga: Ambohidray | reserve: TR18L14: Andriambe stream: | S-18.61317; E48.32593; 1044m: | stagnant pool in pitfall holes: | 23/05/2018; Leg. T. Ranarialalatiana // Paratype | Copelatus kely sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -2♂GP , 2♀, 10 ex. (Alc.)  (NHRS): // NHRS-JLKB | 000010858–9, 65738–9, 10861(Alc.) // Madagascar: Moramanga: Ambohidray | reserve: TR18L04: Andriambe stream: | S-18.6132; E48.3262; 1044m: | stagnant pool in pitfall holes: | 07/04/2018; Leg. T. Ranarialalatiana & al. // Paratype | Copelatus kely sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -2♀ , 5 ex. (Alc.)  (NHRS): // NHRS-JLKB | 000065701, 65740, 10862(Alc.) // Madagascar: Moramanga: Ambohidray | reserve: TR18L07: Andriambe stream: | S-18.6131; E48.3257; 1046m: | stagnant pools in path: | 07/04/2018; Leg. T. Ranarialalatiana & al. // Paratype | Copelatus kely sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -1♂(GP) (NHRS): // NHRS-JLKB | 000065746 // MAD: TOAM: Alaotra Mangoro | Andasibe Mantadia NP, Analamazaotra | 250m E of park entrance: Mad14-18: | dried up river bed with stagnant pools: | 18.9357S; 48.4174E: 930m: 27.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | & T. Ranarilalatiana // Paratype | Copelatus kely sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 //C.insuetus but C.kely is smaller than all other species in the Copelatusinsuetus complex on Madagascar; body length 3.8–4.4 mm . Protibia modified, bisinuate and angled basally and broadened distally. Pro- and mesotarsal claws unmodified. Penis thin and simple, in ventral view with apical part slightly leftturned; in lateral view slightly angled after basal third giving a suggested “shoulder” . Parameres as in Figure Male: first three pro- and mesotarsomeres widened, but less so than in er” Fig.  but whicFemale: with very weak, faint and dispersed strioles on anterior half of elytra from third or fourth interval to the lateral margin.C.insuetus species complex on Madagascar. It is a non-latinised adjective.The new species is named after the Malagasy word for small, “kely”, referring to the small body size. It is the smallest species so far known from the Known from the eastern central part of Madagascar, at Ambohidray Reserve and in Analamazaotra NP Fig. .Mantella frogs, but we’ve also found it in muddy, stagnant, forest pools with dead leaves. The altitude of known localities ranges from 930 to 1050 m. At Analamazaotra NP, we found one specimen occurring sympatrically with Copelatusinsuetus at the same locality.This species occurs in the eastern central rainforest. Most specimens were collected from small waterfilled pitfalltrap holes for Mantella frog species. Copelatuskely adds to the importance of this reserve for conservation of rare and microendemic eastern rainforest species. Ambohidray reserve was established in 2013 and managed through collaboration between the local people association (VOI MMA) and Antananarivo University. During fieldwork at the reserve in April 2018 however, we observed worrying signs of disturbances; “tavy” or slash and burn of the forest for agriculture, the cutting of woods for charcoal, signs of zebu-cattle along forest paths. These factors could cause a serious threat to the aquatic insect fauna of the reserve. The reserve of Ambohidray harbours some species not known from anywere else on Madagascar. If the reserve has any ambition to serve as a refugium for these species, activities destroying or degrading the forests or aquatic habitats should be avoided. Copelatuskely is very close to C.insuetus, and the two species were not reciprocally monophyletic in the CO1 gene tree  (NHRS): // NHRS-JLKB | 000010849 // Madagascar: Fianarantsoa: Ihorombe: R.S. | Pic d’Ivohibe Corridor: at the confluence of | two rivers Inganga and Anefitany: | S22.456683; E046.956283; 874 m; 09.XII.2013 | GB Nets & sieves: forest pools with dead | leaves, Leg. J.H. Randriamihaja & | T. Ranarialalatiana: Field# MAD13-55 // Holotype | Copelatus vokoka sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // Paratypes: -6♂ (GP), 2♀, 16 ex.  (Alc.) : // NHRS-JLKB | 000010850–4, 10857, 65735–6, 10855(Alc.) // Madagascar: Fianarantsoa: Ihorombe: R.S. | Pic d’Ivohibe Corridor: at the confluence of | two rivers Inganga and Anefitany: | S22.456683; E046.956283; 874 m; 09.XII.2013 | GB Nets & sieves: forest pools with dead | leaves, Leg. J.H. Randriamihaja & | T. Ranarialalatiana: Field# MAD13-55 // Paratype | Copelatus vokoka sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -1♂ (GP), 1♀, 3♀ (Alc.) (NHRS): // NHRS-JLKB | 000010512–3, 10782(Alc.) // Madagascar: Fianarantsoa: Ihorombe: | R.S. Pic d’Ivohibe Corridor: The | confluence of rivers Inganga and | Anefitany: S22.457283; E046.95535; 870 m, | 09.XII.2013, GB Nets & sieves: big | muddy pool, Leg. J.H. Randriamihaja & | T. Ranarialalatiana: Field# MAD13-57 // Paratype | Copelatus vokoka sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // Fianarantsoa. Vatovavy Fitovinany: Ifanadiana: -1♂ (GP), 4♂, 11♀ : // Madagascar. Ex-prov. Fiana- | rantsoa. ca. 3.3 km WSW | Ranomafana. 28 XII 2017. | Ramahandrison & Manuel leg. // 21°16'05"S, 47°25'28"E Alt. 993 m. | Shallow shaded pond with large | quantity of dead tree leaves and | some Poaceae, in forest. | Ranomafana NP. // Coll. | M. Manuel | Paris // Paratype | Copelatus vokoka sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 //C.insuetus, and eyes smaller. Elytral striae more deeply impressed and intervals therefore slightly more convex . Protibia modified, bisinuate and angled basally and broadened distally. Pro- and mesotarsal claws unmodified. Penis long, thin and simple; apex in ventral view straight and not leftturned; in lateral view rather evenly arched, lacking the distinct “shoulder” characteristic of C.insuetus but with a different type of postmedial and preapical suggested humps in the curvature. Apex in lateral view also broder closer to apex. Apex in left lateral view with a dorsal ridge crossing posterior dorsal margin but at a more acute angle .We collected this species in 2013 from forest pools with dead leaves next to streams at the Ivohibe reserve in pristine humid forest at an altitude of 870 m. The reserve of Pic d’Ivohibe was established in 1964 and is managed through collaboration between Madagascar National Parks, local people associations, and other partners. During our visit in 2013, there were little signs of degradation except at the entrance and at the west edge of the reserve. Local people sometimes take zebu cattle with them on a path through the forest. At a larger scale zebu excrement could influence freshwater quality and species assemblages through eutrophication, but there were no signs of this inside the intact pristine forest. In Ranomafana NP it was collected in a shallow clear-water shaded forest pond with large quantity of dead tree leaves and some Copelatusvokoka sp. nov. falls in the irinus group, based on the number of elytral striae. The new species is most closely related to C.ankaratra according to the mitochondrial gene CO1 and belongs to the C.insuetus species complex on Madagascar.Taxon classificationAnimaliaColeopteraDytiscidaeRanarilalatiana & Bergstensp. nov.2CC38FF3A07F541E8F8A801E6B92EAB4http://zoobank.org/2E3F790E-F146-4865-8BBE-4B521FF74094Manjakatompo Ankaratra Reserve, Tsiafajavona mountain  Antananarivo. Vakinankaratra: Ambatolampy: -HT♂ (GP) (NHRS): // NHRS-JLKB | 000065412 // Madagascar: Ambatolampy: Manjaka- | tompo Ankaratra Reserve: MAD16-11: | Tsiafajavona mountain: S-19.35163; | E47.24278; 2597 m: stream near source: | 07/02/2016; Leg. T. Ranarialalatiana // Holotype | Copelatus ankaratra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // Paratypes: -12♂ GP, 10♀, 77 ex. (Alc) : // NHRS-JLKB | 000010644–59, 10863–7, 65704, 10778(Alc.) // Madagascar: Ambatolampy: Manjaka- | tompo Ankaratra Reserve: MAD16-11: | Tsiafajavona mountain: S-19.35163; | E47.24278; 2597 m: stream near source: | 07/02/2016; Leg. T. Ranarialalatiana // Paratype | Copelatus ankaratra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -6♂ (GP), 3♀, 60 ex. (Alc) : // NHRS-JLKB | 000010612–7, 65750–1, 10870, 10776(Alc.) // Madagascar: Antananarivo: | Vakinankaratra: Manjakatompo Stn | forestière: Anosiarivo: S19.344889 E | 047.304139, 2073 m. 24.I.2012: lake near source | Field# MJK12-13 : Leg. T. | Ranarilalatiana & J.H. Randriamihaja // Paratype | Copelatus ankaratra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -12♂ GP, 6♀, 115 ex. (Alc.) : // NHRS-JLKB | 000010628–43, 10868–9, 10777(Alc.) // Madagascar: Ambatolampy: Manjaka- | tompo Ankaratra Reserve: MAD16-08: | Anosiarivo: S-19.34505; E47.30384; | 2062 m: large pond in stream source: | 06/02/2016; Leg. T. Ranarialalatiana // Paratype | Copelatus ankaratra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 //C.insuetus complex. The penis also lacks any apical dorsal ridge crossing dorsal posterior margin in left lateral view and is devoid of any sulcation ventrolaterally other than a faint microsculpture.The best diagnostic character for the species is the angled base of penis in posteroventral view Fig.  which seC.insuetus. All appendages testaceous.Body length 4.4–5.2 mm. Body shape very elongate and subparallel. Head infuscated around eyes and medially, testaceous on clypeus and as a posterior band. Pronotum entirely infuscated except lateral margins and especially the anterolateral corners more broadly testaceous. Elytra in same dark brown colour as infuscated parts of head and pronotum, except an irregular basal testaceous band Fig. . Testacerd to 1/2 posterior of base and does not reach apex. Head, pronotum, and elytra microreticulate and finely micropunctate. Posterior third to posterior half of pronotum striolate. Strioles on average coarser than in C.insuetus and in some specimens more extensive onto disc and posteromedially.Elytra with six clearly impressed discal and one submarginal stria. First to fourth elytral striae more or less full length, fifth and sixth striae slightly abbreviated anteriorly; submarginal stria starting 1/3C.insuetus, slightly darker on average so that medial light area around metacoxal lines may be more contrasting. Strioles on metacoxa rather short. Prosternal process more rhomboid and apex more strongly raised medially than in C.insuetus. Lateral parts of metaventrite rather broad. Metacoxal lines long and anteriorly diverging, similar to C.insuetus.Ventral side similar to C.insuetus). Protibia modified, bisinuate, angled basally, and broadened distally. Pro- and mesotarsal claws unmodified. Penis in posteroventral view distinctly angled at base .Known only from a few localities in the Ankaratra Massif Reserve on the central highland plateau of Madagascar Fig. .Copelatusankaratra seems to be a high-altitude alpine species associated with spring water.This new species was collected in the mountains of Ankaratra at altitudes above 2000 m. The first locality, Anosiarivo forest, consisted of water from a source flowing over grass vegetation at an altitude of 2070 m. The second locality, Tsiafajavona Mountain, was a small stream with grass vegetation downstream but very near to the source at an altitude of 2600 m near Ankaratra peak. Copelatusankaratra sp. nov. belongs to the irinus group, based on the number of elytral striae. It belongs to the Copelatusinsuetus species complex radiation on Madagascar and based on its CO1 it is most closely related to C.vokoka from Ivohibe. Notably, both these species have a densely striolated female form. There was a surprisingly large genetic distance (2.3–2.5% uncorrected p-distance) between the locality near the peak (2600 m) and the locality in the forested region ca. 5 km away at a lower altitude (2070 m). We find the male genitalia and other characters very similar and treat them here as conspecific.Boophiswilliamsi  and Mantidactyluspauliani Guibé, 1974 only found in montane streams at elevations above 2000 m.The Ankaratra Massif is an area known for several microendemic species not known from anywhere else on Madagascar. This includes two critically endangered micro-endemic frogs, Manjakatompo forestry station was established in 1923 in the forested part of the mountains to preserve an area of 8320 ha, out of which only 650 ha is natural forest and 2300 ha has been replanted with exotic trees . Even thTaxon classificationAnimaliaColeopteraDytiscidaeRanarilalatiana & Bergstensp. nov.5433FAE18DE15C1298549264274DB2EFhttp://zoobank.org/576CE645-9356-483E-806F-3030D772B260Tsaratanana reserve, Antetikalambazaha Camp  .Mahajanga. Sofia: Bealanana: -HT♂ (GP) (NHRS): // BMNH-672729 // HOLOTYPE // Madagascar: Tsaratanana NP | [Antetykalambazaha Camp], 14.1824S, | 48.9448E, 1700 m, 20–24.xii.2004 | P32, Leg. Lees_D, Ranaivosolo_R // DNA Voucher | BMNH <672729> | MSL027:A07 // Holotype | Copelatus | pseudostriatus sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // Paratypes: -1♀ (NHRS): // BMNH-672728 // PARATYPE // Madagascar: Tsaratanana NP | [Antetykalambazaha Camp], 14.1824S, | 48.9448E, 1700 m, 20–24.xii.2004 | P32, Leg. Lees_D, Ranaivosolo_R // DNA Voucher | BMNH <672728> | MSL027:A06 // Paratype | Copelatus | pseudostriatus sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -1♀ (NHMUK): // BMNH-672727 // PARATYPE // Madagascar: Tsaratanana NP | [Antetykalambazaha Camp], 14.1824S, | 48.9448E, 1700 m, 20–24.xii.2004 | P32, Leg. Lees_D, Ranaivosolo_R // DNA Voucher | BMNH <672727> | MSL027:A05 // Paratype | Copelatus | pseudostriatus sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 //Copelatus species from Madagascar; the penis is gently curved from base to apex and parameres are long and thin  and pronotum may have been infuscated medially prior to exposure to lysis buffer. Elytra uniformly testaceous brown with a faint suggestion of a darker transverse field preapically Fig. . Appendath to 1/3rd posterior from the base. Between first and second, and between second and third striae, there are irregular traces of intermediate striae, or “pseudostriae”, extending from just after base until 1/4th from the apex  and striatus (furrowed or striated) and refers to the intermediate non-complete striae in-between the complete continuous striae on the elytra in this species. It is the only species of Copelatus on Madagascar with this characteristic. The word striatus (masculine) is a participle (verb as adjective) in the nominative singular (ICZN 11.9.1.1).The name Endemic to Madagascar, only known from the type series from Tsaratanana Massif Fig. .Not known, but the type series of specimens were collected in 2004 likely from a stream, near Antetikalambazaha Camp at an altitude of 1700 m.Copelatuspseudostriatus sp. nov. is hardly possible: based on the complete striae it would fall in the irinus group, but the incomplete pseudostriae are likely reduced striae of an ancestor with a higher number of complete striae. This is a very distinct species with no recognisable close relatives, either based on genitalia or the CO1 gene. Tsaratanana Massif contains the highest peak in Madagascar at 2876 m and possibly this species is endemic to the Tsaratanana Massif. However, Rhantusmanjakatompo Pederzani & Rocchi, 2009, collected at the same place and time as the type series of C.pseudostriatus, is a species shared between Tsaratanana and Ankaratra Massif in central Madagascar.Species group assignment of Taxon classificationAnimaliaColeopteraDytiscidaeRanarilalatiana & Bergstensp. nov.851D9EEE861E5E29812C70504B087522http://zoobank.org/137C62A2-22DD-4550-8697-9E81CED4ED45Anjanaharibe Sud reserve,  Antsiranana. Sava: Andapa: -HT♂ (GP) (NHRS): // NHRS-JLKB | 000065415 // MAD: ANTS: Sava: Anjanaharibe | Sud NP: stream next to Camp | site: Mad14-62: medium size | sandy forest stream: 14.7414S | 49.4975E; 910 m: 14.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | & T. Ranarilalatiana // Holotype | Copelatus safiotra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // Paratypes: -2♀ (Alc.) (NHRS): // NHRS-JLKB | 000010785 // MAD: ANTS: Sava: Anjanaharibe | Sud NP: stream next to Camp | site: Mad14-62: medium size | sandy forest stream: 14.7414S | 49.4975E; 910 m: 14.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | & T. Ranarilalatiana // Paratype | Copelatus safiotra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -4♂ (GP), 1♀, 12 ex. (Alc.), 3 ex. (Alc.): : // NHRS-JLKB | 000010506–7, 10566, 10595, 65414, 10775(Alc.), 10845(Alc.) // MAD: ANTS: Sava: Anjanaharibe | Sud NP: Camp site: Mad14-70: | forest stream: 14.7414S | 49.4975E; 910 m: 16.XI.2014 // Leg. J. Bergsten, R. | Bukontaite, S. Holmgren, | J.H. Randriamihaja | & T. Ranarilalatiana // Paratype | Copelatus safiotra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // Toamasina. Alaotra Mangoro: Ambatondrazaka, Moramanga: -4♂ (GP), 29 ex. (Alc.), 6 ex.  (Alc.) : // NHRS-JLKB | 000010818, 10836–7, 65413, 10838(Alc.), 65752(Alc.) // Madagascar: Toamasina: Alaotra | Mangoro: Zahamena NP: Sect. | Antanandava: close to Camp Bemoara | S17.5108; E048.7287; 1060 m. 07.III.2018 | GB Nets, white pan and sieves: Waterfilled | goldigging holes: Field# MAD18-87 // Leg. J. Bergsten, & | T. Ranarilalatiana // Paratype | Copelatus safiotra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -1♂  (NHRS): // NHRS-JLKB | 000010871 // Madagascar: Toamasina: Alaotra | Mangoro: Zahamena NP: Sect. | Antanandava: Manambato stream by Camp | Cascade: S17.545; E048.7237; 1290 m. 09.III.2018 | GB Nets, white pan and sieves: large | foreststream: Field# MAD18-100 // Leg. J. Bergsten, & | T. Ranarilalatiana // Paratype | Copelatus safiotra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -4♂ GP, 2♀, 6 ex.  (Alc.) : // NHRS-JLKB | 000010589–94, 10774(Alc.) // Madagascar: Toamasina: Alaotra | Mangoro: Mantadia NP: Waterfall | 6km from park entrance: S18.83396 | E048.43777, 1000 m, 11.XI.2011 GB | Nets and sieves: forest stream in | rainforest: Field# MAD11-37 // Leg. J. Bergsten, R. | Bukontaite, T | Ranarilalatiana & | J.H. Randriamihaja // Paratype | Copelatus safiotra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // -4♀ (Alc.) (NHRS): // NHRS-JLKB | 000010784 // Madagascar: Toamasina: Alaotra | Mangoro: Mantadia NP: River | Sahanody 9 km from park entrance: S | 18.80973 E 048.42861, 930 m. 11.XI.2011 | GB Nets and sieves: forest stream in | rainforest: Field# MAD11-34 // Leg. J. Bergsten, R. | Bukontaite, T | Ranarilalatiana & | J.H. Randriamihaja // Paratype | Copelatus safiotra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // Fianarantsoa. Matsiatra Ambony: Lalangina: -1♂ (GP) (NHRS): // NHRS-JLKB | 000010846 // MAD: FIAN: Matsiatra Ambony | Ranomafana NP: next to Sahamalaotra | trail entrance: Mad14-04: forest stream: | with sandy bottom: 21.2395S 49.3947E: 1130 m: 02.XI.2014 // Leg. J. Bergsten, | T. Ranarilalatiana | & S. Holmgren // Paratype | Copelatus safiotra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 // Fianarantsoa. Ihorombe: Ihosy: -2♂ (GP) (NHRS): // NHRS-JLKB | 000010847–8 // Madagascar: Fianarantsoa: Ihorombe: | Isalo NP: 300m into the canyon de | Makis: S22.48665 E45.37966, 700 m | 13.XI.2012, GB nets and sieves: canyon | river with side pools: Leg. R. Bukontaite | & J.H. Randriamihaja: Field# MAD12-03 // Paratype | Copelatus safiotra sp. nov. | Det. Ranarilalatiana | & Bergsten, 2019 //C.insuetus on habitus appearance, but sturdier, broader pronotum and head with a much greater interocular distance compared to width of eyes, and subparalell along a longer distance of body with more rapidly attenuating anterior and posterior ends; easily distinguished from all other irinus group species of Madagascar by the penis shape, which is of a type otherwise found in species closely related to Copelatusowas; penis has a large medial expansion in lateral view followed by an apical blade  where dead leaves and debris accumulate. The species has been found at altitudes between 700 and 1300 m but most numerous at elevations above 900 m in primary humid forest. The discovery in Isalo NP, in a sandy river running through a Canyon, indicates that subhumid forests may also be part of the species’ niche.Copelatussafiotra sp. nov. falls in the irinus group, based on the number of elytral striae . However, the genitalia is of a very different type and characteristic of the complex of species close to C.owas in the erichsonii group with ten discal and one submarginal striae. We hypothesize that this species, despite the body shape and number of elytral striae, is not related to the other species treated here, but belongs to the radiation of species around C.owas. This would reinforce the idea that the number of elytral striae is a very homoplastic character and not reliable to create phylogenetically sound groups : sp. 1ea805ca0-ac56-5ec7-aeed-92758eebdba0C.nodieri & C.nodierivar.somalicus for comparison with sp1of ]: -LT♀ : // Data in NHRS | JLKB | 000065417 // Haut Sénégal | Khayes | Dr. Nodier | 11–12. 1881 // Type [red label] // ♀ [female symbol] // nodieri // Coll. Guignot // -PLT♀ : // Data in NHRS | JLKB | 000065418 // Khayes | H Senegal // Nodieri Reg // Coll. Régimbart // -PT♀ : // Data in NHRS | JLKB | 000065419 // SOMALIA IT. | Belet Amin. | (Giuba) Apr. 1923 | Patrizi // Paratype [red label] // ♀ [female symbol] // Museo Civico | di Genova // Copelatus | nodieri | var. ♀ somalicus | Guign. Paratype // var. somalicus | Det. J. Bergsten, 2011 //[Mahajanga. Melaky: Antsalova: -1♀ (CAS): // CASENT 8135000 // Madagascar: Mahajanga | Prov Parc National Tsingy | de Bemaraha, 10.6 km ESE | 123° Antsalova, Elev 150 m | 16–20 November 2001 // 19°42'34"S, 44°43'5"E |coll: Fisher, Griswold et al | California Acad. Of Sciences | sifted litter - tropical dry forest | on Tsingy, BLF4432 // Copelatus sp. nov. | C. pulchellus complex | Det. Ranarilalatiana | & Bergsten, 2019 //pulchellus complex. Three species of the pulchellus complex are known from Madagascar, C.pulchellus, C.marginipennis, and C.mahajanga. It is most similar to C.marginipennis based on its very short and broad body shape. However, it differs in being almost entirely black dorsally, only slightly rufous laterally on the elytra, pronotum, and part of head. As such it is most similar in colouration to the dark form of C.pulchellus on Madagascar, but the body is shorter and broader. A second distinguishing characteristic is the very abbreviated first stria, present only in the posterior third, hence abbreviated even more than in C.mahajanga. Finally, this female specimen has fine but dense striolations mediolaterally on the elytra, and is densely punctured in the posterolateral corners of the pronotum  of this female which differed substantially from the three other species in the complex (K2P: 8.2–13.6%). DNA matching of the male, once discovered, would be straightforward.It is plausible that it was one or several female specimens of this species that ale Fig. . We consTaxon classificationAnimaliaColeopteraDytiscidae(Andasibe and Ranomafana): sp. 20aac52e2-8b52-5955-a42c-c87201c92ee7Toamasina. Alaotra Mangoro: Moramanga: -1♀ : // Data in NHRS | JLKB | 000065698 // Madagascar, E, Andasibe | Analamazoatra Res. | 48°25'12.1"E /18°56'14.2"S | 938 m (26) 17.1.2015, | leg. Berger & Dostal // coll. Wewalka // Copelatus | longicornis group? | Det. Wewalka 2017 // Copelatus sp. nov.? | near C. peridinus | Det. Ranarilalatiana | & Bergsten, 2019 // Fianarantsoa. Vatovavy Fitovinany: Ifanadiana: -1♀ (NMW): // Data in NHRS | JLKB | 000065762 // RM: Namorona Bas. (PO221) | Loc. 1km de Vohiparara | Aff. de Namorona Riv. | 47°22'43"E, 21°13'53"S | Alt. 1200 m; 20.04.1994 | Leg. Elouard, J.-M., Sartori, M. // Copelatus sp. nov.? | near C. peridinus | Det. Ranarilalatiana | & Bergsten, 2019 //longicornis species group together with C.befasicus. However, the five striae are likely not homologous as C.befasicus has an abbreviated first stria, whereas the first stria is entirely lacking in this specimen, therby creating an interstriae space double in width compared to the outer intervals. But despite the lack of the first stria, it has five striae on the central and lateral parts of elytra where C.befasicus only has four. In addition, the striae are very faintly impressed, intermediate between real striae and the puncture lines found in C.peridinus. The colouration is uniformly brownish black like in C.peridinus and the body size is also similar. This specimen is possibly a different species compared to all others presented here; however, we cannot rule out that intraspecific variation of C.peridinus ranges from two defined puncture lines to five weakly impressed striae. They are very similar in all other aspects. We managed to sequence a partial fragment of CO1 (447 bp) from the Andasibe specimen (NHRS-JLKB000065698) and this indeed showed that this specimen is closely related to, or possibly conspecific with, C.peridinus. Genetic distance between them was 3.0% (K2P). Such a level of intraspecific variation is not impossible but unlikely given they were from the same locality . Together with the morphological characters we believe this may be a different and distinct species, but we await the discovery of a male before the identity can be established with confidence. The second female from Namorona River near Ranomafana NP (NHRS-JLKB000065762) is substantially larger (6.4 mm) than the female from Andasibe (5.6 mm) but otherwise shares the same characteristics : sp. 370c03b1e-e8d2-5a8f-8f48-8ffe0eb3ceb6Fianarantsoa. Ihorombe: Ivohibe: -3♀ (NHRS): // NHRS-JLKB | 000010856, 65699, 65734 // Madagascar: Fianarantsoa: Ihorombe: R.S. Pic | d’Ivohibe: Andaranovory: close to botanical | transect R.S. Pic d’Ivohibe: S22.47511667 | E046.9559, 1106 m, 10.XII.2013, GB Nets and | sieves: small lake with dead leaves and | vegetation, Leg. J.H. Randriamihaja & | T. Ranarialalatiana: Field# MAD13-61 // Copelatus sp. nov. | C. insuetus complex | Det. Ranarilalatiana | & Bergsten, 2019 // Toamasina. Atsinanana: Toamasina, Toamasina II: -1♀ (NHRS): // NHRS-JLKB | 000010811 // Madagascar: Toamasina II: Analalava | reserve: MAD17-12: S of nursery plants: | S17.71055; E49.45002; 39 m: Forest | stream with side pools: 09/03/2017; | Leg. T. Ranarilalatiana // Copelatus sp. nov. | C. insuetus complex | Det. Ranarilalatiana | & Bergsten, 2019 // -1♀ (NHRS): // NHRS-JLKB | 000010779 // Madagascar: Toamasina: | Antsinanana: RN2, 6Km N | Toamasina by bridge: S18.06493 | E049.37856, 0 m. 15.XI.2011, | GB Nets and sieves: river and | sidepool: Field# MAD11-52 // Leg. J. Bergsten, R. | Bukontaite, T | Ranarilalatiana & | J.H. Randriamihaja // Copelatus sp. nov. | C. insuetus complex | Det. Ranarilalatiana | & Bergsten, 2019 //C.insuetus complex , a distance that does not rule out conspecificity as the geographic and altitudinal distance are substantial between these localities , there are currently three endemic genera. Other Dytiscidae genera like Copelatus are not endemic but may still contain endemic species radiations within. Here we have revised the species of Copelatus on Madagascar excluding the erichsonii species groups. We recognise 13 species with names and three additional non-named species based on females. Of the named species nine are endemic to Madagascar, two (C.marginipennis and C.distinguendus) are regional endemics to Madagascar and nearby west Indian Ocean islands, and two (C.pulchellus and C.peridinus) also occur on the African continent. This gives an endemic proportion at species level of approximately 70%, an intermediate level compared to other insect groups on Madagascar , longicornis (1 species), irinus (10 species), and erichsonii (> 20 species) species groups. Copelatusunguicularis of the consors species group turned out to be a species of the genus Madaglymbus. As has been flagged before, these species groups are commonly not monophyletic  mostly occupy open, often anthropogenically disturbed, habitats of the Central Highlands. The C.pulchellus complex , and C.befasicus seem to be most abundant in the dry deciduous western forests, lowlands with open to semiopen landscapes. Finally, some species are specialists like the high altitude crenophile C.ankaratra sp. nov. and the sandy stream specialist C.safiotra sp. nov. Not until we have this knowledge – what are the species, how can we recognise them, where do they occur and how do they live – can we attempt to protect them and their habitats in the face of constantly increasing human pressure on the environment.


The correct name is: Ariana Paulina-Carabajal. The correct citation is: Paulina-Carabajal A, Sterli J, Müller J, Hilger A (2013) Neuroanatomy of the Marine Jurassic Turtle Plesiochelys etalloni . PLoS ONE 8(7): e69264.


This article has been corrected: The correct affiliation information is given below:1 Department of Surgical Oncology, Nanjing First Hospital, Nanjing Medical University, Nanjing 210008, Jiangsu Province, Chinahttps://doi.org/10.18632/oncotarget.21195Original article: Oncotarget. 2017; 8:86410-86422.


The correct name is: Ghareib W. Ali. The correct citation is: Mahmoud SS, Ibrahim IH, Sallam ASM, Ali GW (2018) Paradox response of cornea to different color intensities of visible light: An experimental study. PLoS ONE 13(5): e0196827.


Scientific Reports 10.1038/s41598-017-11728-6, published online 12 September 2017Correction to: This Article contains errors in Reference 17, which was incorrectly given as:Applied Physics Letters101, 203105 (2016).Rodrigues, M. S., Costa, L., Chevrier, J. & Comin, F. Why do atomic force microscopy force curves still exhibit jump to contact? The correct reference is listed below as reference


The correct name is: Ian H. Spicknall. The correct citation is: Bulla I, Spicknall IH, Gromov D, Romero-Severson EO (2018) Sensitivity of joint contagiousness and susceptibility-based dynamic optimal control strategies for HIV prevention. PLoS ONE 13(10): e0204741.


Author Hendrik-Jan Mijderwijk appears incorrectly. The correct citation is: Mijderwijk H, Stolker RJ, Duivenvoorden HJ, Klimek M, Steyerberg EW (2018) Prognostic model for psychological outcomes in ambulatory surgery patients: A prospective study using a structural equation modeling framework. PLoS ONE 13(4): e0193441.


The correct reference is: Patel, N. et al. A dyad of lymphoblastic lysosomal cysteine proteases degrades the antileukemic drug L-asparaginase. J Clin Invest 119, 1964–1973,


PubMed PMID: 12837737.Reference 25 is incorrect. The correct reference is: Casewell M, Friis C, Marco E, McMullin P, Phillips I. The European ban on growth-promoting antibiotics and emerging consequences for human and animal health. The Journal of antimicrobial chemotherapy. 2003;52(2):159–61. Epub 2003/07/03. doi:


Metagenomics in ophthalmology: current findings and future prospectives. BMJ Open Ophth 2019;4:e000248. doi: 10.1136/bmjophth-2018-000248.Borroni D, Romano V, Kaye SB, The license type for this paper has changed from CC BY-NC to CC BY.