Dactylonotus Parent, 1934 - a disjunct generic distribution in southern Africa, New Zealand and New Caledonia (Diptera, Dolichopodidae)

Directeur De la publication / Publication director : Gilles Bloch Président du Muséum national d’Histoire naturelle réDactrice en chef / editor-in-chief : Laure Desutter-Grandcolas assistante De réDaction / assistant editor : Anne Mabille ([email protected]) Mise en page / Page layout : Anne Mabille coMité scientifique / scientific board : Nesrine Akkari (Naturhistorisches Museum, Vienne, Autriche) Maria Marta Cigliano (Museo de La Plata, La Plata, Argentine) Serge Gofas (Universidad de Málaga, Málaga, Espagne) Sylvain Hugel (CNRS, Université de Strasbourg, France) Marco Isaia (Università degli Studi di Torino, Turin, Italie) Rafael Marquez (CSIC, Madrid, Espagne) Jose Christopher E. Mendoza (Lee Kong Chian Natural History Museum, Singapour) Annemarie Ohler (MNHN, Paris, France) Jean-Yves Rasplus (INRA, Montferrier-sur-Lez, France) Wanda M. Weiner (Polish Academy of Sciences, Cracovie, Pologne) couverture / cover : Dactylonotus univittatus (Loew, 1858) South Africa. 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Diffusion – Publications scientifiques Muséum national d’Histoire naturelle CP 41 – 57 rue Cuvier F-75231 Paris cedex 05 (France) Tél. : 33 (0)1 40 79 48 05 / Fax : 33 (0)1 40 79 38 40 [email protected] / https://sciencepress.mnhn.fr © Publications scientifiques du Muséum national d’Histoire naturelle, Paris, 2025 ISSN (imprimé / print) : 1280-9551/ ISSN (électronique / electronic) : 1638-9387 799 ZOOSYSTEMA • 2025 • 47 (34) © Publications scientifiques du Muséum national d’Histoire naturelle, Paris. www.zoosystema.com urn:lsid:zoobank.org:pub:16EC7EF8-367C-46EF-B6D6-301076E147C2 Bickel D. J. 2025. — Dactylonotus Parent, 1934 – a disjunct generic distribution in southern Africa, New Zealand and New Caledonia (Diptera, Dolichopodidae). Zoosystema 47 (34): 799-807. https://doi.org/10.5252/zoosystema2025v47a34. http://zoosystema.com/47/34 ABSTRACT The genus Dactylonotus Parent, 1934 (Diptera, Dolichopodidae) is characterized by a finger-like projection of the antennal pedicel along the dorsal margin of the postpedicel. It comprises six species from southern Africa, Dactylonotus formosus (Parent, 1933) from New Zealand and D. eurypterus n.sp. from New Caledonia. Of morphological interest, Dactylonotus eurypterus n.sp. totally lacks anterior preapical setae on femora II and III, present in all congeners. The disjunct distribution of Dactylonotus is unusual, suggesting a relictual distribution comprising Africa, New Zealand and New Caledonia. The genus is unknown from Australia. RÉSUMÉ Dactylonotus Parent, 1934 – distribution générique discontinue en Afrique australe, en Nouvelle-Zélande et en Nouvelle-Calédonie (Diptera, Dolichopodidae). Le genre Dactylonotus Parent, 1934 (Diptera, Dolichopodidae) se caractérise par une projection en forme de doigt du pédicelle antennaire le long de la marge dorsale du postpédicelle. Il est constitué de six espèces d’Afrique australe, Dactylonotus formosus (Parent, 1933) de Nouvelle-Zélande et D.eurypterus n.sp. de Nouvelle-Calédonie. À noter sur le plan morphologique, Dactylonotus eurypterus n.sp. est totalement dépourvu de soies préapicales antérieures sur les fémurs II et III, présentes chez tous ses congénères. La répartition disjointe de Dactylonotus est inhabituelle, suggérant une répartition relictuelle entre l’Afrique, la Nouvelle-Zélande et la Nouvelle-Calédonie. Le genre est inconnu en Australie. Daniel J. BICKEL Australian Museum, 1 William Street, Sydney, NSW 2010 (Australia) [email protected].au Submitted on 20 March 2025 | Accepted on 21 May 2025 | Published on 19 December 2025 Dactylonotus Parent, 1934 – a disjunct generic distribution in southern Africa, New Zealand and New Caledonia (Diptera, Dolichopodidae) MOTS CLÉS Dolichopodidae, biogéographie, répartition gondwanienne, Nouvelle-Calédonie, Nouvelle-Zélande, Afrique australe, espèce nouvelle. KEY WORDS Dolichopodidae, biogeography, gondwanan distribution, New Caledonia, New Zealand, southern Africa, new species. 800 ZOOSYSTEMA • 2025 • 47 (34) Bickel D. J. INTRODUCTION Dactylonotus Parent, 1934 (Diptera, Dolichopodidae Latreille, 1809) is a distinctive genus that was previously known only from South Africa and Zimbabwe (Grichanov 2016). However, two papers published almost simultaneously (Grichanov 1998; Bickel 1999) referred the New Zealand species Syntormon formosus Parent, 1933 to Dactylonotus. This was based on the species description and illustrations of the male antenna, hypopygium and wing in Parent (1933), and was interpreted as a possible Gondwanan relictual distribution involving southern Africa and New Zealand. Described below is an additional species of Dactylonotus from New Caledonia, adding to the unusual distribution of the genus. It was discovered while sorting large undetermined institutional holdings of New Caledonian Dolichopodidae. MATERIAL AND METHODS Photographs were made with a Leica M205A photomontage system. In describing the hypopygium, or male genital capsule, ‘dorsal’ and ‘ventral’ refer to morphological position prior to genitalic rotation and flexion. Thus, in figures showing a lateral view of the hypopygium, the top of the page is morphologically ventral, while the bottom is dorsal. Morphological terminology follows Cumming& Wood (2017). Measurements were made on representative dry specimens. Body length of males is measured from the base of the antennae to the tip of the seventh abdominal segment. The CuAx ratio is the length of the dm-m crossvein/distal section M4. The position of features on elongate structures such as leg segments is given as a fraction of the total length, starting from the base. The relative lengths of the podomeres are representative ratios and not measurements, and are given for each leg in the following formula and punctuation: trochanter + femur; tibia; tarsomere 1/ 2/ 3/ 4/ 5. The wing venation system follows that in Cumming & Wood (2017). AbbreviAtions Morphology I, II, III pro-, meso-, metathoracic legs; ad anterodorsal; av anteroventral; C coxa; dv dorsoventral; F femur; MSSC Male secondary sexual character(s), the non-genitalic character(s) found only on male body; pd posterodorsal; pv posteroventral; t tarsus; T tibia; t1-5 tarsomeres 1 to 5. Wing venation M1 first medial vein; R1 first longitudinal vein; R2+3 second longitudinal vein; R4+5 third longitudinal vein. Institutions BPBM Bishop Museum, Honolulu, Hawaii; CNC Biosystematics Research Institute, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada; INHS Illinois Natural History Survey, University of Illinois, Champaign; MNHN Musém national d’Histoire naturelle, Paris; NZAC New Zealand Arthropod Collection, Landcare Research, Auckland. TAXONOMY Family DolichopoDiDAe Latreille, 1809 Genus Dactylonotus Parent, 1934 Dactylonotus Parent, 1934: 136. type species.— Dactylonotus grandicornis Parent, 1934, monotypy. DiAgnosis.— Body length 3.8-5.5mm; male face wide, only slightly narrowed beneath antennae; antennal pedicel in both sexes prolonged into dorsal finger-like projection resting in dorsal groove on basal half of postpedicel; postpedicel usually enlarged, and bladelike, larger in male than female, arista basidorsal, median or subapical on postpedicel; acrostichal setae short, biseriate on anterior slope of mesonotum, and posteriorly either short and irregular, or as 4-5 uniseriate setae; median scutellar setae strong, lateral scutellar setae short; lower proepisternum with one or two projecting setae, dorsad of CI base; FII and FIII each with strong anterior preapical seta (totally absent on New Caledonian D.eurypterus n.sp.); male in Afrotropical species sometimes with enlarged pulvilli or other modifications of tarsus I (MSSC); vein M 1 with flexion (bosse alaire) between dm–m crossvein and wing apex; male sternite 8 with two strong black projecting setae on distal margin, and with additional shorter erect setae; epandrium subcylindrical; epandrial lobe short with strong projecting distal setae; surstylus divided into separate dorsal and ventral arms, sometimes clavate; male cercus often digitiform; female oviscapt with tergite 10 bearing five pairs of blade-like acanthophorites. remArks Dactylonotus Parent, 1934 is readily recognised by the diagnostic finger-like projection of the antennal pedicel that dorsally overlaps the postpedicel in both sexes (Figs1C-D; 3B, D). The genus has a peculiar disjunct distribution, with six species in South Africa and Zimbabwe, a single species in New Zealand, and a new species from New Caledonia. Ihave never seen the genus among large institutional holdings of Australian Dolichopodidae. Dactylonotus has been placed in the subfamily Diaphorinae Schiner, 1864, but a number of characters, such as the strong anterior preapical setae on femora II and III, are discordant with the traditional concept of the subfamily (however, these anterior preapical setae are totally absent on D.eurypterus n.sp.). Based on a preliminary cladistic analysis, Capellari (2013) suggested that a generic group comprising Anepsiomyia Bezzi, 1902, Argyra Macquart, 1834, Dactylonotus Parent, 1934, Somillus Brèthes, 1924, and Symbolia Becker, 1922 is monophyletic and possibly deserving of separate subfamily rank. This requires further investigation. 801 Dactylonotus (Dolichopodidae) from New Caledonia ZOOSYSTEMA • 2025 • 47 (34) The genus Dactylonotus Parent, 1934 now comprises eight species (see Grichanov [2016] for key and treatment of the six Afrotropical species): – D.eurypterus n.sp., New Caledonia; – D.formosus (Parent, 1933), (Syntormon), New Zealand; – D.frater Parent, 1939, Zimbabwe; – D.grandicornis Parent, 1934, South Africa; – D.nigricorpus Grichanov, 2016, South Africa; – D.rudebecki Vanschuytbroeck, 1960, South Africa; – D.meuffelsi Grichanov, 1998, South Africa; – D.tsitsikamma Grichanov, 2016, South Africa; – D.univittatus (Loew, 1858), (Saucropus), South Africa. compArAtive mAteriAl D.univittatus (Loew, 1858): South Africa • 2 ♂, 2 ♀; Natal; 75km WSW of Eastcourt, Cathedral Peaks For. Sta.; 1500m, 7-31. XII.1979; Peck S.& J. leg.; CNC (Fig.1A-C)]. pAcific species Two species of Dactylonotus from the southwest Pacific Ocean, D.eurypterus n.sp. from New Caledonia and D.formosus from New Zealand are treated below. Both species have strong uniseriate acrostichal setae while their southern African congeners have short biseriate or irregular acrostichal setae. In other respects, the two species are morphologically distinct and probably not close sister taxa with the genus. Dactylonotus eurypterus n.sp. (Figs1D; 2A-C) urn:lsid:zoobank.org:act:760FB4E0-CFCB-426E-B4DA-2F68F32B9347 type mAteriAl.— Holotype. New Caledonia • ♂; Province Nord: Mt. Panié track; 20°33’S, 164°46’E; 25.VIII-9.XI.2000; 900m; Malaise trap; J. Skevington& C. Burwell leg.; CNC1731377; MNHN. Paratype. New Caledonia • ♂; same data as holotype; CNC1731365. other mAteriAl exAmineD.— New Caledonia • 1 ♂; Province Nord Yambé (as Yiambi); 500-700m; 14.X.1967; J.& M. Sedlacek leg.; BPBM • 1 ♂; Province Sud; Mt. Koghi; 500-600m; 23-27.X.1967; J.&M.Sedlacek leg.; BPBM • 1 ♂; Mt. Koghi; 500m; 17km NE of Noumea; 1-3.XI.1992; Malaise trap; M.E. Irwin& D. W. Webb leg.; INHS. Fig. 1. — Dactylonotus univittatus (Loew, 1858) (South Africa): A, male habitus, left lateral; B, female, head and thorax, right laterodorsal; C, male, head and thorax, left dorsal. Dactylonotus eurypterus n. sp. (New Caledonia); D, male, head and thorax, right laterodorsal. Scale bars: A, B, 1.0 mm; C, D, 0.5 mm. AB CD 802 ZOOSYSTEMA • 2025 • 47 (34) Bickel D. J. e tymology .— The specific epithet eurypterus is from Greek meaning “broad wing”, in reference to the broad male wing of this species. Description Male Length: 5.0mm; wing: 4.7× 1.8mm (Fig.2). Head. Subcircular in anterior view, wider than high; dorsal postcranium dark metallic blue-green and flattened; frons metallic blue-green with grey pruinosity, one pair of strong postvertical setae displaced out line with dorsal postocular setae; pair of strong vertical setae and pair of strong diverging ocellar setae present; face wide, only slightly narrower than width of face at antennal base and covered with dense grey pruinosity; eye facets enlarged anteroventrally; palp rather small, pale yellow with short black apical seta; proboscis yellow; antenna mostly yellowish, but with postpedicel infuscated dorsally and distally; scape short, bare of setae; pedicel tg7 d.sur v.sur e.l. pha hyp cer epan st8 tg6 AB C Fig. 2. — Dactylonotus eurypterus n. sp.: A, male habitus, left lateral; B, male habitus, left lateral, more detailed; C, male postabdomen, left lateral. Abbreviations: cer, cercus; d.sur, dorsal surstylus; e.l., epandrial lobe; epan, epandrium; hyp, hypandrium; pha, phallus; st8, sternite 8; tg6, tg7, tergites 6, 7; v.sur, ventral surstylus. Scale bars: A, B, 1.0 mm; C, 0.1 mm. 803 Dactylonotus (Dolichopodidae) from New Caledonia ZOOSYSTEMA • 2025 • 47 (34) prolonged dorsally with twoshort apical setae and resting in dorsal groove on basal half of postpedicel; postpedicel large, and halberdor blade-shaped, dorsally excavated and attached to apex of dorsal pedicel extension, so that ventral postpedicel extends basad from attachment; arista arising dorsally at 2/3, about as long as head height, and relatively thick; ventral postcranium with field of white hairs. Thorax. Mesonotum mostly yellow, glabrous with metallic green reflections, scutellum mostly shining metallic blue with yellow at lateral margins; postscutellum and pleura mostly pale yellow; setae black; fourstrong uniseriate acrostichal setae with tiny biseriate setae anteriormost; six strong dorsocentral setae, slightly decreasing in size anteriorly, with posteriormost seta slightly offset laterally; onepostalar seta; twopostsutural supra-alar setae, onepresutural intra-alar seta; twonotopleural setae; onepostpronotal seta; onepresutural supra-alar seta, onepresutural supra-alar seta; pair of median scutellar setae strong, lateral scutellar setae short; lower proepisternum with single pale yellow seta dorsad of CI base. Legs. Coxae and remainder of legs almost entirely yellow, but apex of TIII and tarsus III light brown; CI with abundant long white anterior hairs and with longer white distolateral setae; CII with black anterior hairs and with strong anterolateral seta at 3/5and apically; CIII with strong black lateral seta at 1/5; legs with short black vestiture; I: 5.8;5.6;5.0/2.5/2.0/1.0/ 0.7; FI with row of 4projecting posterior setae, from ¼ to ½, each seta about width of FI in length (MSSC?), and with preapical pv seta; TI with short dorsal seta at 1/5and 3/5; II: 5.7; 7.3; 6.0; 2.8/1.8/1.2/0.8/0.6; FII lacking anterior preapical seta, but with short preapical av and pv seta; TII with ad-pd setal pair at 1/6, offset setal pair near 2/5, and pd seta at ¾, and with subapical circlet of pd, dorsal, ad, av and pv setae; III: 7.2; 8.5; 2.7/3.6/2.0/1.2/0.6; FIII lacking anterior preapical seta, but with short preapical av and pv setae; TIII with row of fiveshort spaced dorsal setae along length and with subapical circlet of dorsal, ad, av and pv setae. Wing. Membrane hyaline; wing relatively broad and with long curve distal margin (MSSC?); vein R2+3ending in costa near 5/6, with vein R4+5joining margin just anteriad of apex; vein M1with flexion (bosse alaire) one half distance between dm-m and apex, and displaced slightly anteriad, becoming subparallel with R 4+5 , and joining margin at wing apex; CuAx ratio: 0.5; anal angle developed; lower calypter yellow with dark brown rim and fan of brown setae; halter pale yellow. Abdomen. Tergite 1basally yellow with brown distal margin; tergites2-4brown but each with large rounded mediolateral translucent yellow area; tergite5brown; tergites6-7yellow; tergites1-5 covered with short black vestiture, with some longer setae near posterior tergal margins; tergite6 bare of setae; sternite8 yellow, ovate with two strong setae along its distal margin and other scattered setae, and forming cap over left lateral hypopygial foramen; hypopygium (Fig.2C) mostly yellow, and surstylus brownish; epandrium ovate; hypandrium forming flat elongate cover over phallus, apically curved; epandrial lobe short, with three apical setae; surstylus with two distinct arms, ventral lobate arm with short apical setae, and dorsal clavate arm, distally expanded with setae (see Fig.2C); cercus yellow, tapering and digitiform with strong apical setae. Female None associated. remArks Dactylonotus eurypterus n.sp. is known from three mid-elevational sites (500-900m) on Grande Terre, the main island of New Caledonia. Like many of its southern African congeners, this New Caledonian species has the thorax and legs mostly yellow (cf. D.univittatus Loew, 1858 from South Africa, Fig.1C, and D.eurypterus n.sp. from New Caledonia, Fig.1D). Diagnostic characters for Dactylonotus eurypterus n.sp. include: 1) males with a row of four erect posterior setae on the basal half of femur I, absent on other congeners (MSSC); 2)only a single long curved seta on the lower proepisternum (Fig.1D) while other members of the genus have two setae, one long and one short; and 3) the hypandrium in lateral view being relatively straight (hyp, Fig.2C) whereas in most congeners, the hypandrium appears somewhat contorted in lateral view (e.g., Grichanov2016, figs4, 10). However, most importantly from a wider taxonomic viewpoint, Dactylonotus eurypterus n.sp. totally lacks anterior preapical setae on femora II and III, present in all other congeners. The presence of strong anterior preapical femoral setae has been used as an important character in defining dolichopodid subfamilies, so such intra-generic loss or nonexpression appears rather anomalous. Dactylonotus formosus (Parent, 1933) (Fig.3) Syntormon formosus Parent, 1933: 367. t ype mAteriAl .— Parent described Syntormon formosus from specimens collected at various locales in New Zealand, but two males were designated as types, and later this was wrongly interpreted as holotype and paratype. Two males, both from Wellington, New Zealand, have been designated holotype and paratype, possibly during curation and data capture (https://scd.landcareresearch.co.nz/ Specimen/NZAC04021516), NZAC, examined. As noted in the Introduction, this species was referred almost simultaneously to Dactylonotus by both Grichanov (1998) and Bickel (1999), based on the original species description and illustrations of the male antenna and hypopygium. ADDitionAl mAteriAl.— New Zealand • 1 ♂; Auckland: Kauri Park, North Shore; 15.XI.1973; J. Dugdale leg.; NZAC • 2 ♀; Henderson Park, Opanuku Walkway, at Border Rd.; 7.XII.2005, 16.XII.2005; S. E. Thorpe leg.; NZAC. reDescription Male Length. 5.7mm; wing: 5.2× 2.0mm (Fig.3). 804 ZOOSYSTEMA • 2025 • 47 (34) Bickel D. J. Head. Frons blue-green with grey pruinosity; row of short postocular setae, white on ventral half and black on dorsal half; with dorsalmost setal pair (=postvertical setae) stronger and displaced onto surface of dorsal postcranium; palp enlarged, ovate, yellow with one short black apical seta (MSSC); antenna mostly yellow basally, with pedicel and postpedicel infuscated; pedicel prolonged dorsally and resting in dorsal groove on basal half of postpedicel; postpedicel large and blade-shaped, so that ventral postpedicel extends basad from attachment at apex of pedicel (MSSC). Thorax. Mesonotum mostly dark brown with iridescence and complex pruinosity: mostly grey, with narrow brown stripes between acrostichal and dorsocentral rows, and laterally over humeral region; extending posteriorly almost to edge of mesonotum acrostichal band becoming silvery white on posterior mesonotal slope, and narrow brown stripes laterad of acrostichal band becoming wider and dark brown posteriorly; scutellum metallic blue; pleura mostly brown with grey pruinosity, but with areas of yellow cuticle (Fig.3B); metepimeron yellow; 6-7 strong uniseriate acrostichal setae; six strong dorsocentral setae, slightly decreasing in size anteriorly, lateral scutellar seta about one third length of median seta; lower proepisternum with long and short pale yellow curved setae. Legs. Coxae and remainder of legs almost entirely yellow; anterior face of CI basally with short white setae and distal half with short black hairs; with three strong black lateral setae along distal half and row of 4-5 short apical setae; CII with strong anterolateral seta at 1/3 and 4/5; CIII with strong black lateral seta at 2/5; I: 7.0;7.2; 4.2/2.4/1.8/1.0/0.8; TI with short ad seta at 1/4, 1/3 and 3/5, short pd seta at 1/4 and 1/2, and with short subapical dorsal, pd, and posterior setae; II: 7.7; 8.3;5.7/2.8/1.5/0.8/0.7; FII with strong anterior preapical seta, and with short subapical av and pv seta and av and pv rows of short setae in basal two-thirds; TII with offset ad-pd setal pairs at 1/5, 2/5, and 3/5, and with subapical circlet of pd, dorsal, ad, av and pv setae; III: 9.0;10.2;3.0/4.3/1.9/1.0/0.8; FIII with strong anterior preapical seta, with short preapical av and pv setae; and with short av and pv setae; TIII with ad seta at 1/8, 1/4, and 1/2, and AB CD Fig. 3. — Dactylonotus formosus (Parent, 1933), New Zealand: A, male habitus, left lateral; B, male, head and thorax, left laterodorsal; C, male postabdomen, left laterodorsal; D, female male, head and thorax, right lateroventral. Scale bars: A, 2.0 mm; B, D, 1.0 mm; C, 0.5 mm. 805 Dactylonotus (Dolichopodidae) from New Caledonia ZOOSYSTEMA • 2025 • 47 (34) with five dorsal setae along basal two thirds; row of short ventral setae along entire length, and subapical circlet of dorsal, ad, av and pv setae. Wing. Membrane hyaline; vein R4+5 joining margin anteriad of apex; vein M1 with flexion (bosse alaire) halfway between dm-m and wing apex, and distally displaced, becoming subparallel with distal R4+5, and joining wing apex; CuAx ratio: 0.6; anal angle developed; lower calypter pale yellow with fan of pale yellow setae; halter pale yellow. Abdomen. Tergite 1 yellow with long posterior setae; tergites 2-5mostly yellow, with dark brown posterior margin; tergites 2-5 with short black vestiture, with longer setae near posterior tergal margins; tergites 6-7mostly yellow; sternite 8 yellow, ovate with two strong setae along its distal margin and other scattered setae; hypopygium (Fig.3C) mostly yellow, epandrium ovate; surstylus with two subparallel arms, with ventral arm truncate and dorsal arm tapering with pale curve hairs; cercus brown, digitiform with fine whitish hairs. Female (Fig.3D) Similar to male except as noted: postpedicel also bladelike but much shorter and basally yellowish; FII and FIII without rows of short av and pv setae; female oviscapt with epiproct (tergites 9+10) bearing five pairs of blade-like acanthophorites. remArks Dactylonotus formosus (Parent, 1933) is known from scattered locales on the North Island and the Banks Peninsula on the South Island of New Zealand. The complex thoracic pruinosity pattern is distinctive. Photos of live specimens have been posted on https://www.inaturalist.org/taxa/380089Dactylonotus-formosus. DISCUSSION The historical biogeography of the unusually rich New Caledonian flora and fauna has been the subject of considerable discussion (e.g., Holloway 1979; Grandcolas etal. 2008; Heads 2023; Malem etal. 2023). Previously, the main island (Grande Terre) was regarded as a Gondwanan fragment supporting a unique biota of vicariant origin. However, geological evidence suggests that New Caledonia was submerged during the Palaeocene and without subaerial exposure until the late Eocene-early Oligocene when its ultramafic lithosphere was uplifted. Thus, many workers now consider the New Caledonian biota to be the result of post-Eocene transoceanic dispersal following geological emergence, and this view is supported by various taxonomic studies. Even so, Giribet& Baker (2019) discuss the problem of limited salt-water dispersal capability in many New Caledonian “paleoendemic” ground-dwelling invertebrates. Dactylonotus formosus eurypterus frater grandicornis nigricorpus rudebecki tsitsikamma univittatus Fig. 4. — Distribution map of Dactylonotus Parent, 1934 species.