taxonID	type	description	language	source
03CFD1313520163BE58FF9A0845F5E10.taxon	description	http: // zoobank. org / urn: lsid: zoobank. org: act: F 7 A 6827 A-F 6 B 4 - 463 C-BDCA- 56 B 461 DD 0189 Figs 1 a, c, e, g, 2 a, c, e, 3 a, c, 4, 5 a, b, c	en	Bernard, Rafał, Daraż, Bogusław (2026): Paragomphus sofiae sp. nov. - a new spotted species of Hooktail from Madagascar (Odonata: Anisoptera: Gomphidae). Zootaxa 5757 (4): 357-368, DOI: 10.11646/zootaxa.5757.4.4, URL: https://doi.org/10.11646/zootaxa.5757.4.4
03CFD1313520163BE58FF9A0845F5E10.taxon	etymology	Etymology. The name, a noun in apposition, refers to the beautiful Sofia River, where the species was discovered. The origin of the river name is unknown; in this or similar spelling it occurs on maps in the 19 th century, earliest found as Soffea (Vandermaelen 1827).	en	Bernard, Rafał, Daraż, Bogusław (2026): Paragomphus sofiae sp. nov. - a new spotted species of Hooktail from Madagascar (Odonata: Anisoptera: Gomphidae). Zootaxa 5757 (4): 357-368, DOI: 10.11646/zootaxa.5757.4.4, URL: https://doi.org/10.11646/zootaxa.5757.4.4
03CFD1313520163BE58FF9A0845F5E10.taxon	materials_examined	Type material. Holotype male (NHC-T-INS- 000018) and two paratype males (NHC-T-INS- 000019, NHC-T-INS- 000020) from eastern banks of the Sofia River, 2 km N of Antsatramalaza, 15.5983 ° S, 48.5613 ° E, 250 m asl, Sofia Region, Madagascar, 28.11.2023, leg. R. Bernard and B. Daraż. Third paratype male (NHC-T-INS- 000021) obtained at a water body at the Bemarivo River (in the Sofia River basin), 4 km ESE of Tsarahasina, 15.7785 ° S, 47.6134 ° E, 30 m asl, Sofia Region, Madagascar, 16.12.2023, leg. R. Bernard and B. Daraż.	en	Bernard, Rafał, Daraż, Bogusław (2026): Paragomphus sofiae sp. nov. - a new spotted species of Hooktail from Madagascar (Odonata: Anisoptera: Gomphidae). Zootaxa 5757 (4): 357-368, DOI: 10.11646/zootaxa.5757.4.4, URL: https://doi.org/10.11646/zootaxa.5757.4.4
03CFD1313520163BE58FF9A0845F5E10.taxon	description	Description. Holotype male. Total length 47.0, abdomen (excluding appendages) 32.3, Fw 26.3, Hw 24.2, Fw Pt 3.1, cerci 3.1. Fw Ax 12, Fw Px 9, Hw Ax 9, Hw Px 8 – 9. Head. Face black, marked extensively with yellow: the four black bands / areas are interspersed with four yellow ones (Fig. 5 b). Vertex black. Postfrons black centrally with two large yellow spots laterally (Figs 1 a, 5 c). Antefrons mostly yellow with a black dorsal frontlet (Fig. 5 b). Postclypeus mostly yellow with a black trapezoidal band along its lower margin. Anteclypeus yellow (Fig. 5 b). Mandibles (mandible bases, mandibular cheeks) yellow with narrow darkened margins. Genae yellow. Labrum variegated, with brown to black margins, black centre and two large lateral yellow spots (Fig. 5 b). Lateral lobes of labium yellow with brown inner margins and the median lobe yellow to brown distally. Occiput yellow with black lateral margins. Postgenae black to dark brown with yellow markings. Eyes when alive are grassy green: light with a yellowish admixture in the dorsal half and dark with a whitish streak in the ventral half (Figs 5 b, c). Thorax. Prothorax black dorsally to dark brown ventrally, with a lateral yellow spot. Sides of the synthorax dark brown to black, yellow-spotted (Figs 1 a, 5 a). Crest of the middorsal carina black with a brownish-yellowish spot (Fig. 1 a). Collar yellow, its two spots narrowly separated from each other (Fig. 5 c). On the mesepisternum, the postdorsal stripe represented by a large oblong yellow spot well-separated from the collar. The yellow antehumeral stripe divided into a large elongated spot situated more ventrally and a small semioval dorsal spot (Figs 1 a, 5 a, c). Mesokatepisternum brownish dorsally to yellowish ventrally. The mesepimeral stripe consists of two barely touching, uneven yellow spots, the smaller and more squarish upper spot, and the more elongated, drop-like lower spot (Fig. 1 a). The metepisternal stripe divided into three well-separated spots, one on the metakatepisternum below the metastigma, and two above the metastigma, the middle one being the largest (Figs 1 a, 5 a). The metepimeral stripe is a large, broadly T-shaped spot covering most of the metepimeron (Figs 1 a, 5 a). Poststernum brown with a lighter brownish to yellowish anterior corner (Figs 1 a, 5 a). Legs. Coxae and trochanters yellow to brownish. Femora combining brown and yellow in various proportions and blackening distally: they are mostly brown in hindlegs and with yellow ventrolateral surfaces in mid- and forelegs (Figs 1 a, 5 a, c). Tibiae black, with thin longitudinal yellow stripes on the dorsolateral surface in fore- and midlegs (Figs 5 b, c). Wings. Membrane hyaline, costa yellow. Pt relatively wide compared to the width of adjacent rows of cells and black (Fig. 1 a). Abdomen. Colour pattern divided into two sections: S 1 – 6 black with a bright yellow pattern of paired spots, while S 7 – 10, forming a club, mostly reddish to rusty brown with less regular, partly diffused yellow markings (Figs 1 a, 4, 5 a). The lateroventral spot on each side of S 1 large. The similarly located spot on each side of S 2 even larger, two branched, with the anterior branch also covering the auricle (Figs 1 a, 5 a). Still on S 2, a single spot on the dorsal carina, extending along the almost whole of the segment (Fig. 4). The dorsal pattern on S 3 is more complex and consists of two spots on each side along the dorsal carina; the anterior spot does not cross the supplementary transverse carina and the posterior one crosses half of the segment’s length (Figs 4, 5 a). On S 3, the stripe along the ventral carina broadly yellow; at base the colour extended laterally but not reaching the dorsal spots. On S 4 – 6, the yellow pattern gradually reduces (Figs 1 a, 4, 5 a). It consists of two spots: (a) an anterior transverse spot, extending along the whole segment’s height and being narrow centrally and broadened dorsally and ventrally; and (b) a posterior dorsal spot, on S 4 reaching half of the segment’s length, on S 5 already small, and on S 6 totally lacking. S 7 with an anterior dorsolateral yellow marking, crossing the supplementary transverse carina and reaching 40 % of the segment’s length dorsally. Along the ventral carina, this marking is prolonged as a stripe far posteriorly (Figs 4, 5 a). The rest of S 7 dark reddish brown. S 8 and S 9 reddish to rusty brown. On S 7 – 9, a small black spot situated laterally on rings of posterior field of tergites (Figs 1 a, 5 a). S 10 reddish to rusty brown with diffuse yellowish markings. Foliations much larger on S 8 than on S 9, reddish brown with a broad black margin (Figs 1 a, 4, 5 a). Secondary genitalia. In lateral view, the anterior hamule evenly sickle-shaped, tapering markedly towards the apex, and directed posteriorly (Figs 2 e, 3 a). The posterior hamule distinctly more compact in structure, in lateral view with two adjacent branches on a common stout stem: the anterior branch is much longer than the posterior one and both are rounded apically (Figs 2 e, 3 a). In ventral view, a stout centripetal protrusion is visible at the base of the posterior branch of the posterior hamule (Fig. 3 c). Hood of penial peduncle (vesicle) oval, brown (Figs 2 e, 3 a). Caudal appendages. Almost uniformly reddish orange, darker basally (Figs 1 c, e, g, 5 a). In lateral view, cerci stout in their basal half and gradually narrowing distally, tapered to a finely pointed apex (Figs 1 c, 2 c). They are rather gently and gradually curved downwards in the distal third of their length, with an obtuse angle (about 140 °) between the straight anterior part and the curving posterior part. Eleven tiny black denticles along the ventral ridge of cerci tips (Figs 1 c, g, 2 c). In dorsal view, tips of cerci divergent (Figs 1 e, 2 a). In ventral view, cerci appear twisted with diverging and narrowing tips (Fig. 1 g). Epiproct, in lateral view, half as long as the cerci, arched down, with its distal half ascending and broadening gradually, being broadest at the subapical level, and with minute black apical denticles (Figs 1 c, 2 c). In ventral view, the epiproct deeply divided into two cudgel-like branches, each with a slightly distended ‘ head’ and a black apical edge located laterally (Fig. 1 g). Variation in paratype males. Total length 47.1 – 48.5, abdomen (excluding appendages) 32.3 – 33.8, Fw 26.3 – 27.4, Hw 24.6 – 25.6, Fw Pt 3.5 – 3.7, cerci 3.0. Fw Ax 12 – 13, Fw Px 7 – 9, Hw Ax 9 – 10, Hw Px 8 – 10. Variation in colouration is limited to small differences in the proportions of dark and yellow colours. A good example is the crest of the synthoracic middorsal carina, which, unlike the mostly black crest in the holotype (Fig. 1 a), is black with a central yellow spot in one paratype and yellow with a black distal tooth in two other paratypes (Figs 5 a, c). The following features also exist in slightly more yellow version (s) in one to three paratypes compared to the holotype: (a) vertex not uniformly black, but with two or three minute yellow spots / stripes; (b) antefrons slightly more yellow with a black frontlet being discontinuous and declining laterally (Fig. 5 b); (c) labrum predominantly yellow, with less marked dark margins; (d) labium more uniformly yellow, without darker inner margins of lobes; (e) occiput more broadly yellow (Figs 5 b, c); (f) a broader yellow stripe along the ventral carina of S 3, extending up to three quarters of the segment’s length (Fig. 5 a); (g) a dorsal posterior yellow spot (minute) on each side of S 6; and (i) S 8 diffusely yellowish basally (Fig. 5 a). On the contrary, two yellow spots of the mesepimeral stripe are more broadly separated by black colour in two paratypes than in the holotype. Female. Unknown.	en	Bernard, Rafał, Daraż, Bogusław (2026): Paragomphus sofiae sp. nov. - a new spotted species of Hooktail from Madagascar (Odonata: Anisoptera: Gomphidae). Zootaxa 5757 (4): 357-368, DOI: 10.11646/zootaxa.5757.4.4, URL: https://doi.org/10.11646/zootaxa.5757.4.4
03CFD1313520163BE58FF9A0845F5E10.taxon	diagnosis	Differential diagnosis. Paragomphus sofiae is remarkably similar to P. fritillarius in the yellow pattern of its head, synthorax and abdomen, and in the reddish brown tip of the lattermost (Figs 1 a, b), but it is slightly smaller, with male Hw 25.0 (24.2 – 25.6, N = 4) versus 27.3 (26.6 – 27.9, N = 3), and distinct by its unique hamules and caudal appendages. The anterior hamule of P. sofiae is much shorter with similar proportions of its parts, the basal stouter section and the decurved distal section, while in P. fritillarius, the distal section is much longer resulting in the much larger overall hamule being more scythe-shaped than the regular sickle of the former species (Figs 2 e, f, 3). In lateral view, the posterior hamule of P. sofiae looks a bit like a mirror image of that in P. fritillarius: in the former species the finger-like anterior branch of the hamule is much longer than its posterior branch, while in the latter species only the claw-like posterior branch is recognisable (Figs 2 e, f, 3 a, b). Cerci in P. sofiae versus P. fritillarius: (a) are uniformly reddish orange versus black-tipped (Figs 1 c, d); (b) are downcurved at an obtuse angle versus abruptly bent down at a right angle (Figs 1 c, d, 2 c, d); (c) in lateral view, are narrowed gradually versus consisting of two contrasting sections, a strong proximal part and a much thinner distal part (Figs 1 c, d, 2 c, d); and (d) are armed with a regular row of 10 – 11 small black denticles along the ventral ridge of the tip versus have only minute and hardly recognisable and less regularly situated denticles there (Figs 1 g, h). Epiproct of both species is deeply divided into two cudgel-like branches whose stems are proportionately much longer in P. sofiae than in P. fritillarius (Figs 1 g, h).	en	Bernard, Rafał, Daraż, Bogusław (2026): Paragomphus sofiae sp. nov. - a new spotted species of Hooktail from Madagascar (Odonata: Anisoptera: Gomphidae). Zootaxa 5757 (4): 357-368, DOI: 10.11646/zootaxa.5757.4.4, URL: https://doi.org/10.11646/zootaxa.5757.4.4
03CFD1313520163BE58FF9A0845F5E10.taxon	distribution	Distribution and ecology. Paragomphus sofiae is known from two localities situated 103 km apart in the Sofia River basin in lowlands of northern Madagascar (Fig. 8), in the ecoregion of Madagascar Dry Deciduous Forest (One Earth 2025) and the hydrographic ecoregion of Northwestern Basins (Sparks & Stiassny 2022). The Sofia River, certainly providing the larval microhabitats for P. sofiae, is large but shallow, with the sandy river bed 200 – 300 m broad, in many places only partly covered by water at the beginning of the rainy season. This is a braided river consisting of a network of multiple shallow channels splitting and merging around many sand bars (Fig. 6). The flow rate is rather low, water is transparent and mostly a few to 40 cm deep, and bottom deposits are sandy and unvegetated. The river flows through an open, partly cultivated landscape, only with a narrow tree and bushy rim along the riverbanks. Males of P. sofiae were only observed in these wooded edges, where they perched on vegetation in sunny places. In the second location, the male obtained at standing water most probably originated from the adjacent Bemarivo River which also is quite large in its lower reaches (about 115 – 200 m broad) and sandy, but more concentrated and with far fewer sand bars. It also flows through an open, cultivated landscape.	en	Bernard, Rafał, Daraż, Bogusław (2026): Paragomphus sofiae sp. nov. - a new spotted species of Hooktail from Madagascar (Odonata: Anisoptera: Gomphidae). Zootaxa 5757 (4): 357-368, DOI: 10.11646/zootaxa.5757.4.4, URL: https://doi.org/10.11646/zootaxa.5757.4.4
03CFD1313529163AE58FFE4C801A5E80.taxon	materials_examined	Paragomphus fritillarius, used for the comparison with P. sofiae, was collected at two localities in the Sofia Region, north-eastern Madagascar: (a) a small sandy and rocky river in Antsiatsiaka, 16.0589 ° S, 49.1088 ° E, 783 m asl, 30.11.2023, 1 male, leg. R. Bernard and B. Daraż; and (b) a large sandy stream in the foothills of the Marotandrano massif, 16.2654 ° S, 48.836 ° E, 644 m asl, 09.12.2023, 2 males and 1 female, leg. R. Bernard and B. Daraż (Fig. 8).	en	Bernard, Rafał, Daraż, Bogusław (2026): Paragomphus sofiae sp. nov. - a new spotted species of Hooktail from Madagascar (Odonata: Anisoptera: Gomphidae). Zootaxa 5757 (4): 357-368, DOI: 10.11646/zootaxa.5757.4.4, URL: https://doi.org/10.11646/zootaxa.5757.4.4
03CFD1313529163AE58FFE4C801A5E80.taxon	description	The key features of these individuals are identical to those of the species lectotype and paralectotype. The lectotype is shown here (Fig. 7) since the original description (Selys 1892), lacking illustrations and some features, turned out to be ambiguous and insufficient in the face of the discovery of the new, similar species. A clarifying addition also is necessary for the species locus typicus. Selys (1892) gave “ Rumena valley, Mayanga ”. Fraser (1956) most probably did not find this site because he asked if it was perhaps Ramena in Majunga Province. In fact, the name “ Ramena ” (not Rumena) is given on the lectotype and paralectotype oldest labels, being slightly unclearly written on the former (Fig. 7 b) but almost calligraphed on the latter. We have not found any Ramena valley in the former Mahajanga province (= Majanga or Majunga on the 19 th century maps). At the beginning of the 1890 s, however, there was no province yet, and simply the region associated with Majanga, the nearest large town, was given. Thus, in our opinion, the locus typicus was most probably the Ramena River, a sandy tributary of the Sambirano River, which was located near the border of the later Mahajanga province, about 25 km from the seacoast.	en	Bernard, Rafał, Daraż, Bogusław (2026): Paragomphus sofiae sp. nov. - a new spotted species of Hooktail from Madagascar (Odonata: Anisoptera: Gomphidae). Zootaxa 5757 (4): 357-368, DOI: 10.11646/zootaxa.5757.4.4, URL: https://doi.org/10.11646/zootaxa.5757.4.4
