Taxonomic revision of the genus Copelatus of Madagascar (Coleoptera, Dytiscidae, Copelatinae): the non- erichsonii group species Author Ranarilalatiana, Tolotra Author Raveloson Ravaomanarivo, Lala Harivelo Author Bergsten, Johannes text ZooKeys 2019 869 19 90 http://dx.doi.org/10.3897/zookeys.869.33997 journal article http://dx.doi.org/10.3897/zookeys.869.33997 1313-2970-869-19 B7C88A64C06E4B67A352F2F9C8FB0D1C C44C602AB0235FB2922DC2A4915C3288 Copelatus safiotra Ranarilalatiana & Bergsten sp. nov. Figs 6E , 10C Type locality. Anjanaharibe Sud reserve, [14.7414S, 049.4975E] [Madagascar, Sava region, Andapa district] Type material. 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, NHMUK, DEUA & PBZT/MBC): // 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. (3♂, 3♀) (Alc.) (NHRS, NHMUK, DEUA & PBZT/MBC): // 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♂ (teneral) (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. (1♂, 5♀) (Alc.) (NHRS, DEUA & PBZT/MBC): // 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: S 22.48665 E 45.37966, 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 // Diagnosis. Somewhat similar to 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 Copelatus owas ; penis has a large medial expansion in lateral view followed by an apical blade ( Fig. 6E ). Description. Body length 4.3-5.2 mm. Body shape elongate and subparallel along a very long part of the body. Pronotum and head broad and eyes small creating a very wide interocular distance. Maximum width of pronotum clearly in front of hind corners. Head rufotestaceous with weak or absent infuscation in between and posterior of eyes. Pronotum infuscated medially with broadly testaceous lateral sides. Elytra brown, with a broad testaceous transverse band basally ( Fig. 10C ). Testaceous band broader and transitioning posteriorly more diffusely into the brown colour compared with C. insuetus . Antennae, palps and legs testaceous. Elytra with six discal and one submarginal stria. Fifth stria abbreviated anteriorly, variably also first and third striae. In some individuals especially fifth but also first and sixth striae are rudimentary or with very shallow impressions. Interval between fifth and sixth striae narrow, approx. half interval between first and second striae. Submarginal stria short, starting around middle. Head, pronotum, and elytra microreticulate and finely micropunctate. Pronotum not striolate. Ventral side entirely testaceous except last three abdominal ventrites may be vaguely infuscated and have lighter lateral spots. Metacoxa and abdominal sternite II-IV striolate, but strioles shallower and finer than in C. insuetus . Compared with C. insuetus , the metacoxal lines are shorter, and the anterior traces suggest an inward curve towards the posterior metaventral margin. The lateral part of metaventrite is narrower than in C. insuetus at level of mesocoxa, equal to the width of mesofemur at middle. Posterior metaventral margin not straight but slightly angular at level of apex of mesotrochanter. Prosternal process is slightly more elongate. Male: first three pro- and mesotarsomeres widened, ventrally equipped with suction cups. Pattern of suction cups same as for C. insuetus but tarsomeres not as wide and less developed as an integrated protarsal palett. Protibia modified, bisinuate, angled basally, and broadened distally. Pro- and mesotarsal claws unmodified. Penis very characteristic, rather broad and short and in lateral view with a medial expansion followed by a sharp constriction before the narrow blade-like apex ( Fig. 6E ) which is characteristic of several species related to C. owas Regimbart , 1895; apical blade curved leftwards in ventral view and serrated by transverse ridges on the convex right side. Parameres as in Figure 6E , rather long and thin with a medial emargination on the concave side. Female: on average smaller than males (Table 3). At least some females with elytral microreticulation slightly more strongly impressed than in males and therefore appearing more matt. Table 3. Measurements of body length summarised as Min, Max, Mean, and Standard Deviation (SD) for each species, separated by sex. N = number of measured individuals, F = females, M = males.

Species	Sex	N	Min	Max	Mean	SD
C. befasicus	F	4	4.13	4.19	4.15	0.03
C. befasicus	M	-	-	-	-	-
C. marginipennis	F	26	5.16	6.32	5.75	0.30
C. marginipennis	M	45	5.48	6.58	6.02	0.26
C. mahajanga	F	12	5.16	6.13	5.63	0.24
C. mahajanga	M	16	5.42	6.19	5.88	0.20
C. pulchellus	F	1	5.74	5.74	5.74	-
C. pulchellus	M	3	5.48	6.06	5.83	0.30
C. distinguendus	F	44	5.29	6.32	5.75	0.21
C. distinguendus	M	26	5.48	6.26	5.92	0.18
C. peridinus	F	7	5.74	6.58	6.05	0.27
C. peridinus	M	7	5.81	6.26	6.10	0.16
C. baculiformis	F	1	4.00	4.00	4.00	-
C. baculiformis	M	-	-	-	-	-
C. kely sp. nov.	F	7	3.81	4.26	4.01	0.16
C. kely sp. nov.	M	5	3.87	4.32	4.03	0.17
C. insuetus	F	27	4.26	5.03	4.60	0.20
C. insuetus	M	24	4.26	4.90	4.61	0.17
C. safiotra sp. nov.	F	3	4.32	4.71	4.58	0.22
C. safiotra sp. nov.	M	13	4.52	5.16	4.86	0.19
C. vokoka sp. nov.	F	6	3.94	4.32	4.18	0.13
C. vokoka sp. nov.	M	8	4.06	4.45	4.29	0.12
C. ankaratra sp. nov.	F	19	4.39	5.03	4.70	0.17
C. ankaratra sp. nov.	M	30	4.65	5.16	4.87	0.16
C. pseudostriatus sp. nov.	F	2	5.42	5.61	5.52	0.14
C. pseudostriatus sp. nov.	M	1	5.29	5.29	5.29	-

Etymology. The species name safiotra is a Malagasy noun for hybrid, here referring to the unusual combination of a male genitalia type, typical of the Copelatus owas species complex, in a body with a 6+1 striated elytra very much resembling the C. insuetus complex of species. It is a non-latinised noun in apposition. Distribution. Endemic to Madagascar. This species has a rather large distribution in the eastern humid forest from Anjanaharibe Sud reserve in the NE, all along the eastern escarpment including Zahamena NP, Mantadia NP, and Ranomafana NP, and even extending to the rather isolated western patch of subhumid forest at Isalo NP ( Fig. 12D ). Habitat and ecology. This species seems to be strongly associated with clean streams having sandy substrate in humid forests. At these localities, individuals can be found in sidepools, at margins or sites protected from waterflow (e.g., by fallen logs) 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. Comments. Copelatus safiotra sp. nov. falls in the irinus group, based on the number of elytral striae (six discal and one submarginal 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 ( Balke et al. 2004 ).