Copelatus distinguendus Regimbart , 1903

Ranarilalatiana, Tolotra, Raveloson Ravaomanarivo, Lala Harivelo & Bergsten, Johannes, 2019, Taxonomic revision of the genus Copelatus of Madagascar (Coleoptera, Dytiscidae, Copelatinae): the non- erichsonii group species, ZooKeys 869, pp. 19-90 : 19

Copelatus distinguendus Regimbart, 1903 Figs 4B , 7D

Copelatus distinguendus Régimbart, 1903: 19 [nom. nov., referring to his description of Malagasy material under the name Copelatus duodecimstriatus Aubé in Régimbart 1895: 163]

Type locality.

Environs de Tananarive [surroundings of Antananarivo] and Fianarantsoa, Madagascar.

Type information from original description.

Based on an unknown number of specimens (syntypes) collected by Sikora (Antananarivo) and Perrot (Fianarantsoa).

Type material studied.

Type material in MNHN not studied, as it was out on loan.

Additional material studied.

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, DEUA & PBZT/MBC): // NHRS-JLKB | 000010620-5, 10786(Alc.) // Madagascar: Antananarivo: | Vakinankaratra: Manjakatompo Stn. | forestière: 500m E Lac Froid by the | road: S 19.34485 E 047.33381, 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, DEUA & PBZT/MBC): // 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, DEUA & PBZT/MBC): // 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, DEUA & PBZT/MBC): // 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, DEUA & PBZT/MBC): // 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, DEUA & PBZT/MBC): // 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 //

Diagnosis.

Body shape elongate oval, convex, and attenuate posteriorly, with uniform black colouration ( Fig. 7D ). The pattern of abbreviation of the elytral striae separates this species from all other Malagasy Copelatus species. Striae 1, 3, and 5 present only in posterior one third (1) or two thirds (3, 5); 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” ( Fig. 4B ).

Description.

Body length: 5.3-6.3 mm. Body shape elongate oval, convex, and attenuate posteriorly. Head, pronotum, and elytra all the same colour, ferrugineous black and finely punctate.

Lateral margin of pronotum rusty ferrugineous, with short sparse strioles. Pronotum with puncture rows and microsculpture. Elytra narrowly testaceous to ferrugineous posterolaterally. Six elytral striae present and one submarginal stria ( Fig. 7D ). Striae 2, 4, and 6 longer than striae 1, 3, and 5, at least as continued fragments; first stria shortest, present only in posterior fourth to posterior third. Second and third striae present as clearly impressed striae in posterior half or a little longer, but second stria continues, albeit fragmentary and less impressed, more distinctly all the way to base. Fourth stria the most complete, more or less present to base but less deeply impressed or fragmentary in anterior third. Fifth stria abbreviated a little less than third stria, present in approximately posterior two thirds, and like stria 3 has small isolated point remnants of stria more anteriorly. Sixth stria longer than fifth and with more distinct remnants of striae present anteriorly. Elytra covered with punctures in apical part.

Ventral side ferrugineous dark brown. Metacoxa and abdominal sternite II, III, IV, striolate. Prosternal process raised medially but rounded, not carinate. Lateral parts of metaventrite broad. Metacoxal lines short and strongly diverging anteriorly. Antenna, palps, pro- and mesothoracic legs brown to yellowish, but metathoracic legs dark brown.

Male: protibia slightly widened at apex, somewhat curved and angulate basally. Penis in lateral view curved with two points where curvature is more abrupt, constricted before apex at a narrow “neck” and expanding to apical part ( Fig. 4B ). As a whole in lateral view, the penis bears resemblance to the silhouette of a “pumpjack”. Penis apex in ventral view left-turned. Parameres broad at base, curved and tapering towards apex, apical part with numerous setae on ventral margin ( Fig. 4B ).

Female: dorsal sculpture similar to male.

Distribution.

Occurs on Madagascar and Mauritius ( Gomy 2016). Guignot (1948) tentatively refer a single female specimen from Lulua in DR Congo to C. distinguendus but this was very likely a different species. In Madagascar, it has been recorded from the provinces of Antananarivo, Fianarantsoa Toamasina, and Antsiranana; more specifically from Anjozorobe Angavo reserve, around Ankazobe, Manankazo, Andranofeno Sud, around Ambohitantely reserve, Manjakatompo Ankaratra reserve; also from the eastern central area around Moramanga, Analamazaotra NP; in the southeast from Ambositra in col de Tapia, Ankazomivady forest, Ranomafana NP, Andringitra NP; and from the northeast in Anjanaharibe Sud reserve ( Fig. 11D ).

Habitat and ecology.

This species has been collected in various localities, mostly from open, partly deforested areas at mid- to high altitudes or open forest marshes. It occurs at altitudes above 900 m and is often associated with grass vegetation along lake shores and in marshes, found by stamping, and at vegetation-rich margins of rivers.

Comments.

Régimbart (1895) described this species under the name Copelatus duodecimstriatus Aubé but realised later (1903) 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. Copelatus duodecimstriatus lacks submarginal striae, has a distinctly different penis shape, and is endemic to Mauritius ( Vinson 1967; Guignot 1961; Gomy 2016). The closest relatives of Copelatus distinguendus are likely a group of species on continental Africa with similar genitalia (e.g., C. ateles Guignot, 1955, C. sylvaticus Guignot, 1952, C. andreinii Régimbart, 1905, and additional species described by Bilardo and Rocchi 2013). These have a variable number of elytral striae from five discal to ten discal plus one submarginal, but similar “pumpjack” -shaped genitalia ( Bilardo and Rocchi 2013). Copelatus distinguendus differs from these especially by the lack of a lateral tooth on the male penis in ventral view.

Figures

Figure 4. Male genitalia. From left to right, aedeagus in right lateral, ventral, left lateral views and left paramere. A Copelatus peridinus B Copelatus distinguendus C Copelatus pseudostriatus sp. nov.

Figure 7. Habitus, dorsal view. A Male: Copelatus peridinus B Female: Copelatus baculiformis C Female: Copelatus sp. 2 (Andasibe) D Female: Copelatus distinguendus.

Figure 11. Distribution maps of Copelatus species. A C. marginipennis (circle), C. baculiformis (square) B C. mahajanga (circle), C. sp. 2 (square) C C. pulchellus (circle), C. peridinus (square) D C. distinguendus (circle), C. sp. 1 (square).