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Research Article
Odorous and monophyletic – systematic revision of the endemic Madagascan genus Tritonaclia with the first record of wing scent scales in Syntomini Polka Dot Moths (Noctuoidea: Erebidae: Arctiinae)
expand article infoMarcin Wiorek‡, David C. Lees§, Niklas Wahlberg|, Łukasz Przybyłowicz‡
‡ Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland
§ Department of Sciences, Natural History Museum, London, London, United Kingdom
| Department of Biology, Lund University, Lund, Sweden

Corresponding author: Łukasz Przybyłowicz ( lukasz@isez.pan.krakow.pl )

Academic editor: Andreas Zwick
© 2026 Marcin Wiorek, David C. Lees, Niklas Wahlberg, Łukasz Przybyłowicz.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
ZooBank: urn:lsid:zoobank.org:pub:49DCFB91-C573-4307-A0CE-2E478ADE5E26
Open Access

Abstract

In the highly diverse, endemic Madagascan lineage of Syntomini, the “black-and-white” general body pattern appears to be present in a few closely unrelated clades (and in six existing genera), causing uncertainties in their systematics. Based on materials from five natural history collections, we confirm the monophyletic character of the morphologically variable and largely montane genus Tritonaclia Hampson, 1898, with six existing species, inferred from both molecular and morphological evidence. The taxonomy of the genus is revisited, with the description of two new species: T. vonifotsy Wiorek, sp. nov. and T. ombilahy Wiorek & Przybyłowicz, sp. nov.. Naclia melania Oberthür, 1923 syn. nov. is treated as a junior subjective synonym of Glaucopis tollini Keferstein, 1870, and lectotypes of both taxa, now placed in Tritonaclia, are designated. We provide determination keys for the genus, based on adults, and their male and female genitalia. All available data on the distribution and ecology of Tritonaclia species are presented, including new geolocalisation of some important insect collecting places in Madagascar. Additionally, we provide descriptions and illustrations of male scent scales (androconial patches) on fore- and hindwings, present in two Tritonaclia species: T. tollini and T. vonifotsy Wiorek, sp. nov. Such structures are recorded in the tribe Syntomini for the first time.

Keywords

Madagascar, tiger moths, androconia, SEM, new species, Tsaratanana, Marojejy, Anosyenne Mountains

1. Introduction

The endemic Madagascan group of Syntomini is one of the largest evolutionary radiations of Macroheterocera in the island (Przybyłowicz et al. 2019; Lees and Minet 2022). Its members are highly variable morphologically, with both aposematic and cryptic species present (Griveaud 1964; Wiorek et al. 2026). Within that diversity, a few general body pattern ground plans can be indicated, with “black-and-yellow” and “black-and-white” being arguably the most common ones. They are characterised by the predominantly uniform brown to almost black body colouration, with yellow and white wing patterns, respectively. The “black-and-white” pattern is present in a few closely unrelated taxa: four of seven species of Melanonaclia Griveaud, 1964, Juliennaclia moerens (Oberthür, 1911), one of two species in the genus, all species in Tenuinaclia Griveaud, 1964 and Tritonaclia Hampson, 1898, Mortinaclia perplexa (Griveaud, 1964) and Vadonaclia marginepuncta Griveaud, 1964 – the latter two belonging to monotypic genera. In the past, their superficial similarity led to the establishment of systematics based on polyphyletic or paraphyletic genera, as recently figured out and partly revised by Wiorek et al. (2026).

Mountain areas of Madagascar are important habitats from the perspective of the diversity of Syntomini. Many species are local endemics, occurring only in certain mountain massifs and highland areas of the island – especially in Marojejy, but also e.g. Tsaratanana, Andringitra, Montagne d’Ambre or Tampoketsa d’Ankazobe (Griveaud 1964, 1971, 1974; Wiorek et al. 2021; see also Wiorek et al. 2026). Mountains are the least studied environments in Madagascar (Clark et al. 2025). Their Lepidoptera fauna is no exception and many species, including some Syntomini, so far have been collected only once, during single expeditions (‘missions’) to remote mountainous localities (Viette 1991).

Biology and ecology of the Madagascan lineage of Syntomini remain largely unknown, with a few exceptions (Przybyłowicz et al. 2021; Wiorek et al. 2026), a general case with the Afrotropical members of the tribe (e.g. Przybyłowicz 2009; Staude et al. 2023, but see de Freina et al. 2020). This includes their chemical ecology, even though the presence of both dorsal and ventral female pheromone glands is the main and undoubted synapomorphy of the tribe (Kitching and Rawlins 1999; Wiorek et al. 2026). Also, the male scent (androconial) organs are present across different lineages of Lepidoptera, including Arctiinae, but to the best of our knowledge, they have never been studied or even reported from Syntomini so far (Birch et al. 1990; Simmons et al. 2012).

In this paper, we continue the study of the systematics of “black-and-white” Madagascan Syntomini by revising the genus Tritonaclia. We also examine and illustrate the fore- and hindwing patches of transformed scales in T. tollini (Keferstein, 1870) and T. vonifotsy Wiorek, sp. nov.

2. Materials and methods

2.1. Institutional abbreviations

ISEA PAS Institute of Systematics and Evolution of Animals Polish Academy of Sciences, Kraków, Poland – MLUH-ZNS Martin-Luther-Universität Halle-Wittenberg, Zentralmagazin Naturwissenschaftlicher Sammlungen (ZNS), Halle (Saale), Germany – MNHN Muséum national d’Histoire naturelle, Paris, France – NHMUK Natural History Museum, London, UK, former BMNH – British Museum, Natural History – PZBT Parc Botanique et Zoologique de Tsimbazaza, Antananarivo, Madagascar.

2.2. Other abbreviations

DIANA the Region of Diego, Antsohihy, Nosy Be, and Ambanja – SAVA the Region of Sambava, Antalaha, Vohemar, and Andapa – SEM scanning electron microscopy – PN “Parc National”, National Park – RS “Réserve Spéciale”, Special Reserve – MW Marcin Wiorek – DL David C. Lees – ŁP Łukasz Przybyłowicz

2.3. Materials and sampling methods

The study is based on museum specimens from five collections (ISEA PAS, MLUH-ZNS, MNHN, NHMUK, and PZBT), obtained from the second half of the 19th century to 2019 (Table S1). The specimens collected by DL, ŁP and MW were caught by netting during the day or attracted to light at night. For further details of the methodology of collecting Madagascan Syntomini, see Przybyłowicz et al. (2021).

2.4. Morphological study

The genitalia were dissected with the standard method of maceration in a 10% NaOH solution in a water bath, stained with chlorazol black, and mounted in glycerine without a cover slip for photography. Subsequently, the genitalia were mounted in Euparal to make fixed slides. Wing preparation protocol followed Banasiak (2015), with some modifications. All illustrated morphological structures were photographed with a Leica S9i binocular, connected with LasX software. Wing scales for SEM visualisation were sampled from dry specimens with a needle, placed on a carbon disc, coated with gold and photographed using a HITACHI SU8600 ultrahigh-resolution scanning field emission electron microscope (FE-SEM), in the Laboratory of Scanning Electron Microscopy and Microanalysis of the Institute of Geological Sciences, Jagiellonian University, Kraków. The terminology for wing venation and pattern follows Wiorek et al. (2026), for scale morphology follows Downey and Allyn (1975), and for genitalia we adopted the general terminology of Koda (1987). The illustrated specimens were photographed with a Canon EOS 70D camera, with a macro lens EF 50 mm, except for the MLUH-ZNS specimens, photographed with an Olympus TG-6. The photographs of type specimens from the NHMUK collection are courtesy of the Trustees of the Natural History Museum, and those from MNHN were taken by M. Depraetere (2018), available on https://science.mnhn.fr/institution/mnhn/collection/el/item/search. For the details and comments on the morphology and the diagnostic characters of the remaining black-and-white Madagascan Syntomini, see Wiorek et al. (2026).

2.5. Molecular study

For the molecular study, two legs of each specimen were sampled with forceps. Extraction of genomic DNA was done with NucleoSpin Tissue kit (Macherey-Nagel, Germany), according to the manufacturer’s protocol. For each specimen, up to eight molecular markers (one mitochondrial: COI, and seven nuclear: CAD, EF1-alpha, GAPDH, IDH, MDH, RpS5, Wingless) were amplified, using primers by Wahlberg and Wheat (2008), and the protocol described in Zenker et al. (2017). Successful PCR products were sequenced by Macrogen Europe (Amsterdam, the Netherlands). Obtained sequences were processed as described in Przybyłowicz et al. (2021). Newly generated sequences were submitted to GenBank (Table S2). A Tritonaclia dataset related to this study, “DS-TRITO26” was also submitted to BOLD (https://doi.org/10.5883/DS-TRITO26; Process IDs SYNMD001-26-SYNMD012-26; Table S2).

2.6. Phylogenetic study

Except for the newly generated sequences, we also used data from Przybyłowicz et al. (2019) and Przybyłowicz and Wiorek (2023) (Table S2). Phylogenetic analyses were conducted within the Maximum Likelihood (ML) framework using IQ-TREE 3 (Wong et al. 2025), with five runs. The data were initially partitioned by gene and codon position, with ModelFinder (Kalyaanamoorthy et al. 2017) used to select the best partitioning scheme and substitution model, and the partitions allowed to be merged (-m TEST and --merge greedy). The third codon positions of all genes except COI, the second positions of CAD, EF1-alpha, GAPDH, MDH and Wingless, and the first positions of IDH and RpS5 were merged, resulting in 11 partitions. The best model selected for the largest partition was TN+F+I, and GTR+F+I+G4 for most of the remaining ones. Node support values were calculated using the Ultrafast Bootstrap (Hoang et al. 2018) and SH-like approximate likelihood test (Guindon et al. 2010), both with 1000 replicates.

2.7. Distributional study

Localities lacking geographical coordinates in the labels were geolocalised based on Viette (1991), Moat and Smith (2007), and Albignac et al. (1970), using Google Maps and Mapy.com. Missing elevational data were secondary inferred with Google Earth. Some of the localities were previously provided with coordinates by Wiorek et al. (2026). Additionally, a single available iNaturalist observation of Tritonaclia, by P. Łukasik (‘sympiotr’), was included in the study, verified by MW. All collecting localities of the specimens included in this study are presented in Table S3.

The general elevational ranges of vegetation follow Du Puy and Moat (1996) and Clark et al. (2025).

3. Results

3.1. Phylogenetic relationships and systematics

Tritonaclia Hampson, 1898

Type species.

Hydrusa kefersteinii Butler, 1882: 2, by original designation

Diagnosis.

Antennae in both sexes piliform, each antennomere with pair of prominent, arched setae and additional pair of shorter, straight ones (Fig. 1). In the male genitalia, uncus basolaterally provided with differently expressed, setaceous protrusion; valva biramous, with costa transformed into variable, claw-like or spatula-like protrusion, and sacculus developed into weakly sclerotised lobe.

Figure 1. 

Male antenna of Tritonaclia kefersteinii, ms – main seta, as – additional seta (see text).

The most similar to Tritonaclia are other “black-and-white” Madagascan Syntomini, especially from the genera Melanonaclia (in the tree represented by M. nigra Griveaud, 1964 and M. toulgoeti Griveaud, 1964), Vadonaclia Griveaud, 1964 (absent in the tree),

Juliennaclia moerens (congeneric with J. pauliani (Griveaud, 1964), in the tree), as well as Mortinaclia Wiorek, 2026 (monotypic, in the tree) and Tenuinaclia (in the tree represented by T. cf. andapa Griveaud, 1964) (Fig. 2). The genus Melanonaclia (as reviewed by Wiorek et al. 2026) differs in having only intensely yellow, and never white marks on the head, thorax and abdomen, piliform antennae without additional prominent setae, simple male valva without the costal process, and the presence of a pouch-like appendix or diverticulum in the basal portion of the female corpus bursae. Some Melanonaclia species do have a row of cornuti in the male vesica, although differently expressed than in Tritonaclia. The monotypic genus Vadonaclia (V. marginepuncta) differs in intensely orange to reddish head, thorax and abdomen blotches, distinctly bipectinate antennae, and an additional row of small blotches along the fore wing outer margin. The monotypic genus Mortinaclia (M. perplexa), although provided with a row of cornuti in the male vesica, differs from Tritonaclia in much smaller body size, piliform antennae without distinct setae, a differently expressed abdomen pattern extending on its dorsal side, and the presence of distinct lateral arms on the tegumen. The genus Tenuinaclia differs in much smaller body size, a differently expressed pattern of the abdomen: either completely absent or consisting of lateral and ventral rows of at most three small blotches, and only a single distinct cornutus in the male vesica (Griveaud 1964).

Figure 2. 

Phylogenetic relationship of Tritonaclia (blue clade) and selected other “black-and-white” Madagascan Syntomini species, inferred based on Maximum Likelihood analysis of eight markers, in IQ-TREE 3. Nodes support values represent SH-like/Ultrafast Bootstrap results, respectively.

The morphological findings are congruent with the phylogenetic tree (Fig. 2). The genus Tritonaclia, represented in our dataset by four out of six species, for which fresh materials were available for molecular study, forms a separate monophyletic clade. It is not closely related to other groups comprising “black-and-white” species, i.e. Melanonaclia, Tenuinaclia, Juliennaclia Wiorek, 2026 and Mortinaclia. The genus forms three main lineages grouped in two clades, well-supported by both molecular and morphological results. In the tree, the first clade comprises T. kefersteinii and T. quinquepunctata Griveaud, 1967 (represented by the former) and it is sister to all remaining species with a high support. The lineage is characterised by: i) the prominent body coverage with elongate, hairy scales, giving the species “fluffy” appearance; ii) pale yellow to creamy white marks on the abdomen with no blotches on sternites, iii) six or seven pale yellow to creamy white, never semi-transparent forewing wing blotches and one basal yellow blotch at the forewing hind margin (Fig. 3), iv) male genitalia vesica armed with a bunch (not a row) of elongate cornuti (Fig. 4), and v) female genitalia corpus bursae signum developed into a longitudinal plaque (Fig. 5), although the female of T. quinquepunctata remains unknown. The second clade comprises two remaining lineages: T. tollini (sensu Keferstein 1870 nec Griveaud 1964) + T. vonifotsy Wiorek, sp. nov. (i.e. T. tollini, sensu Griveaud 1964), and T. stephania (Oberthür, 1923) + T. ombilahy Wiorek & Przybyłowicz, sp. nov., in the tree represented by all species but the latter, and sister to each other with good support (Fig. 2). They are characterised by: i) white marks on the abdomen, both on tergites and sternites, ii) four or five always semi-transparent pure white forewing wing blotches, with basal blotch always absent, iii) forewing blotch m5 distinctly smaller than m6 (Figs 6, 7, 8); iv) male genitalia vesica armed with a differently expressed row of prominent, tooth-like cornuti (Figs 9, 10, 11, 12). Additionally, the lineage T. tollini + T. vonifotsy Wiorek, sp. nov. is characterised by patches of scent scales on the male fore- and hindwing, and spirally coiled ductus bursae and double signum in the female genitalia (Figs 13, 14). The lineage T. stephania + T. ombilahy Wiorek & Przybyłowicz, sp. nov. has no wing scent scales, wide and short ductus bursae, and a single signum (Fig. 15) – but the female genitalia of T. stephania remain undescribed.

Figure 3. 

Tritonaclia adults. A T. kefersteinii male (MNHN, EL83627–CLV_1); B T. kefersteinii female (MNHN, EL83626–CLV_2); C T. kefersteinii female with reduced wing pattern (NHMUK, NHMUK015109820), see text; D T. quinquepunctata, male (MNHN, EL74630). Scale bars: 1 cm.

Figure 4. 

Male genitalia of Tritonaclia. A, B Ventral view and phallus of T. kefersteinii (MNHN, preparation ♂ nr 124); C, D ventral view and phallus of T. quinquepunctata (MNHN, slide P. Griveaud, No. 501).

Figure 5. 

Female genitalia of Tritonaclia kefersteinii. A MNHN, EL83626-CLV_2, preparation ♀ No. 123; B–D NHMUK, NHMUK015109820, slide NHMUK, NHMUK014331971.

Figure 6. 

Tritonaclia tollini adults. A–C Original specimens of G.A. Keferstein, males; A, B, F, G paralectotypes (MLUH-ZNS), upper- and underside; C, H lectotype (MLUH-ZNS), upper- and underside; D lectotype of Tritonaclia melania syn. nov. (NHMUK, NHMUK010620989); E original illustration of T. tollini in Keferstein (1870); I, J male, upper- and underside (ISEA PAS, DL_2959); K male, lateral view of abdomen (ISEA PAS, DL_01-216). Scale bars: 1 cm.

Figure 7. 

Tritonaclia adults. A–E T. vonifotsy Wiorek, sp. nov., males; A, B holotype, upper- and underside (ISEA PAS, DL_1864); C Lectotype, upperside (MNHN, EL74622); D, E lectotype, upper- and underside (MNHN, EL74624); F T. vonifotsy Wiorek, sp. nov., female, lectotype (NHMUK, NHMUK010918662 – DL1863); G T. tollini, female (ISEA PAS, DL_2981); H, I unidentified female of T. tollini / T. vonifotsy Wiorek, sp. nov., upper- and underside (NHMUK, NHMUK015109812). Scale bars: 1 cm.

Figure 8. 

Tritonaclia adults. A–C T. stephania; A female holotype (NHMUK, NHMUK010620988); B, C male, upper- and underside (ISEA PAS, MAD_166); D–F T. ombilahy Wiorek & Przybyłowicz, sp. nov., male holotype, upper-, underside and lateroventral view of the abdomen (MNHN, EL74649); G T. ombilahy Wiorek & Przybyłowicz, sp. nov. female, originally “neallotype” of T. stephania (MNHN, EL65139, see text). Scale bars: 1 cm.

Figure 9. 

Male genitalia of Tritonaclia tollini. A, B Specimen ISEA PAS, DL_01-216, slide S554; C–E specimen ISEA PAS, DL_2959, slide S555; A–C ventral, dorsal and lateral view, note the shape of tegumen (arrowhead in B), and uncus; D, E phallus; F, G protruding genitalia of the lectotype (MLUH-ZNS), note shape of uncus bifurcation (arrowhead).

Figure 10. 

Male genitalia of Tritonaclia vonifotsy Wiorek, sp. nov. A Ventral view, arrowhead indicates protrusions of juxta; B dorsal view, arrowhead indicates lateral protrusion of uncus; C phallus; D–F lateral view, note variation in uncus length, arrowhead in D indicates protrusion of sacculus; A, B, D paratype, MNHN, EL74622, préparation ♂ no. 155; C, E paratype, MNHN, EL74624, préparation ♂ no. 156; F holotype, ISEA PAS, DL_1864, slide S553.

Figure 11. 

Male genitalia of Tritonaclia stephania, specimen ISEA PAS, MAD_166, slide S556. A Ventral view; B dorsal view; C phallus.

Figure 12. 

Male genitalia of Tritonaclia ombilahy Wiorek & Przybyłowicz, sp. nov., holotype, MNHN, EL74649, préparation ♂ no. 157. A Ventral view, arrowhead indicates elongate arms of juxta; B dorsal view; C lateral view, arrowhead indicates lateral protrusion of tegumen; D, E phallus.

Figure 13. 

Female genitalia of Tritonaclia tollini, specimen ISEA PAS, DL_2981, slide S557. A Ventral view; B dorsal view; C magnification of distal part of genitalia, note coiled ductus bursae; D magnification of signum.

Figure 14. 

Female genitalia of Tritonaclia vonifotsy Wiorek, sp. nov., lectotype NHMUK, NHMUK010918662-DL1863, genitalia slide NHMUK010316589. A Ventral view; B magnification of distal part of genitalia, arrowhead indicates lateral protrusion of lamella postvaginalis (see text); C magnification of coiled ductus bursae; D magnification of signum.

Figure 15. 

Female genitalia of Tritonaclia ombilahy Wiorek & Przybyłowicz, sp. nov., specimen MNHN, EL65139, slide P. Griveaud No. 291, originally “neallotype” of T. stephania (see text).

Taxonomic note.

The genus Tritonaclia (monotypic at the moment of establishment, with T. kefersteinii) was characterised by Hampson (1898: 125) based on the shape of palpi, setation of antennae, and wings venation scheme. Griveaud (1964: 74) sustained this set of characters, indicating additional ones in the male genitalia. According to Griveaud (1964), the forewing veins M2 and M3 should be basally stalked in Tritonaclia and completely separate in Melanonaclia (Griveaud 1964: figs 154, 172), which turned out to be a variable character (see Wiorek et al. 2026: figs 25E–H, 34E, F).

The species Tritonaclia inauramacula Griveaud, 1964, was described only from females and placed in the genus based on the alleged forewing venation scheme mentioned above. However, they appeared to be conspecific with the female of Melanonaclia luctuosa (Oberthür, 1911) collected in copula and described in Griveaud (1969) and therefore synonymised in Wiorek et al. (2026). Thus, in the arrangement proposed here, the genus Tritonaclia consists of six species.

Despite the morphological variation and clear division into three lineages, the genus Tritonaclia as a whole is well-defined by the proposed synapomorphies, especially the antennae setation and the shape of the male valva. Thus, to maintain a possibly stable systematics of the group, we refrain from establishing subgenera, especially within a group of six species.

3.2. Tritonaclia Determination keys

Adults
1 Wing blotches creamy yellow, not semi-transparent (Fig. 3) 2
1’ Wing blotches white, semi-transparent (Figs 6, 7, 8) 3
2 Forewing with six blotches, three distal ones of similar shape and size (Fig. 3A, B) T. kefersteinii
2’ Forewing with five blotches (second medial absent), three distal blotches of unequal size, first one (m4) subrectangular and parallel to costal margin, second one (m5) smaller than third one (m6) (Fig. 3D) T. quinquepunctata
3 Upper surface of hindwing in proximal part with large pale area, provided with suboval blotch of darker colour (Figs 6A–D, 6I, 7A, 7C, 7D) 4
3’ Upper surface of hindwing background uniformly dark brown to black, without additional colour blotch (Figs 7F–H, 8A, 8B, 8D, 8G) 5
4 Hindwing medial suboval blotch ochraceous-creamy to pale yellow (Fig. 7A, C, D) T. vonifotsy Wiorek, sp. nov., male
4’ Hindwing medial suboval blotch ochraceous-brown to grey (Fig. 6A–D, I) T. tollini , male
5 Hindwing subtrapezoidal in general outline (Fig. 7F–I), outer margin slightly sinuous with shallow incision at vein CuA2 T. vonifotsy Wiorek, sp. nov. / T. tollini, female*
5’ Hindwing suboval in general outline, without incision (Fig. 8) 6
6 Small moth, wingspan around 25 mm T. stephania
6’ Big moth, wingspan around 35 mm T. ombilahy Wiorek & Przybyłowicz, sp. nov.
* females of these two species appear indistinguishable morphologically, but see the note for T. vonifotsy Wiorek, sp. nov.
Male genitalia
1 Uncus terminally distinctly divided into two lobes (Figs 9A–C, 10A, 10B, 10F) 2
1’ Uncus terminally undivided (Figs 4A, 4C, 11A, 11B, 12 A–C) 3
2 Terminal lobes of uncus open at around 120 degrees, sacculus without subtriangular protrusion (Fig. 9A, B, F) T. tollini
2’ Terminal lobes of uncus open at around 90 degrees, sacculus basomedially with subtriangular protrusion (Fig. 10A, B) T. vonifotsy Wiorek, sp. nov.
3 Vesica with longitudinal row of around 10–15 heavily sclerotised, tooth-like cornuti (Figs 9D, 9E, 10C, 11C, 12D, 12E) 4
3’ Vesica terminally with bunch of elongate, spine-like cornuti (Fig. 4B, D) 5
4 Uncus with bulbous tip, juxta dorsally without distinct, spine-like protrusions, vesica with row of around 10 massive cornuti (Fig. 11) T. stephania
4’ Uncus terminally widened but not bulbous, juxta dorsolaterally with two prominent, spine-like protrusions, vesica with around 15 massive cornuti (Fig. 12) T. ombilahy Wiorek & Przybyłowicz, sp. nov.
5 Protrusion of costal margin of valva elongate, narrow, distinctly curved downwards (Fig. 4A) T. kefersteinii
5’ Protrusion of costal margin of valva straight, only slightly bent, parallel to main axis of valva (Fig. 4C) T. quinquepunctata
Female genitalia
Remarks. The females of T. quinquepunctata and T. stephania remain undescribed (but for the latter see the taxonomic note)
1 Ductus bursae distinctly spirally coiled (Figs 13A–C, 14A, 14C) T. tollini / T. vonifotsy Wiorek, sp. nov.*
1’ Ductus bursae wide and straight (Fig. 15) 2
2 Signum in form of longitudinal plaque along entire corpus bursae (Fig. 5A) T. kefersteinii
2’ Signum in form of single, rounded plaque located medially on corpus bursae (Fig. 15) T. ombilahy Wiorek & Przybyłowicz, sp. nov.
* female genitalia of these two species appear indistinguishable in the limited available materials; for the potential differences, see the taxonomic note for T. vonifotsy Wiorek, sp. nov.

3.3. Taxonomy

Tritonaclia kefersteinii (Butler, 1882)

Figures 1, 3A–C, 4A, B, 5A, 16A

Hydrusa kefersteinii Butler, 1882: 2.

Tritonaclia kefersteinii Hampson 1898: 136, fig. 154; Griveaud 1964: 74, figs. 181–183, pl. I, fig. 45; Viette 1990: 170; Przybyłowicz and Wiorek 2023: 476, fig. 1A, B, D–F.

Material.

31 specimens, including holotype (MNHN - 9 ♂♂, 1 ♀; NHMUK - 5 ♂♂, 2 ♀♀; PZBT - 12 ♂♂, 2 exx. sex unknown) (Table S1).

Holotype.

♂, NHMUK, NHMUK010621038, by monotypy (in the original description as ♀, in error, see Przybyłowicz and Wiorek 2023).

Diagnosis.

The most similar species is T. quinquepunctata, the diagnostic characters are provided in the determination keys. For more details, see Przybyłowicz and Wiorek (2023).

Morphological remarks.

The female of T. kefersteinii was first described, and the male genitalia were redescribed by Przybyłowicz and Wiorek (2023).

A single female specimen (NHMUK015109820) collected by DL in 2018 has five forewing blotches (the second medial absent), and it lacks the hindwing distal blotches (Fig. 3C). The specimen, although relatively fresh and unspread before this study, appears faded, with head, thorax and basoproximal portion of forewing dark brown, almost black, and the remaining body parts paler, ochraceous brown. It was unavailable for molecular study, but we interpret it as a female of T. kefersteinii with a reduced wing pattern, since the general shape and arrangement of the remaining forewing blotches, especially distal ones, are typical for that species (see the determination key), rather than T. quinquepunctata (see Fig. 3A, B vs. Fig. 3D). Comparing to the first description of the female genitalia (Przybyłowicz and Wiorek 2023), the specimen is almost identical and differs only in shorter, almost absent ductus bursae, and shorter and wider dorsal pheromone glands (Fig. 5). Still, the taxonomic status of this morphotype requires further study.

Distribution.

(Fig. 16A) Disjunct, in the northern and central east of Madagascar, the Regions of: SAVA (PN de Marojejy), Analamanga (La Mandraka), Alaotra-Mangoro (area of Lakato), and Matsiatra Ambony (Ankafina-Tsarafidy). At around 600–1350 m elevation (possibly up to around 1450 m, since the exact locality in the area of Ankafina-Tsarafidy is unknown). See Przybyłowicz and Wiorek (2023) for further comments on the species distribution.

Figure 16. 

Distributional maps of Tritonaclia. A T. kefersteinii; B T. quinquepunctata; C T. tollini, violet dot – doubtful locality, red dot – collecting locality of the rediscovered type series, see text; D. T. vonifotsy Wiorek, sp. nov., violet dot – unidentified females of T. tollini / T. vonifotsy Wiorek, sp. nov., see text; E. T. stephania; F. T. ombilahy Wiorek & Przybyłowicz, sp. nov., violet dot – doubtful locality, see text.

Ecology.

Attracted to light (UV) at night, there is no explicit data on diurnal activity. Recorded in January, February, August and September. Associated with humid, evergreen lowland and midland forests.

Tritonaclia quinquepunctata Griveaud, 1967

Figures 3D, 4C, 4D, 16B

Tritonaclia quinquepunctata Griveaud 1967 [imprint 1966]: 216, figs 6, 29–32; Viette 1990: 170; Przybyłowicz and Wiorek 2023: fig. 1C.

Material.

10 specimens, including holotype (MNHN - 7 ♂♂; NHMUK - 2 ♂♂; PZBT - 1 ♂) (Table S1).

Holotype.

♂, MNHN, EL65138, by original designation.

Diagnosis.

The most similar species is T. kefersteinii. The diagnostic characters are provided in the determination keys. Tritonaclia quinquepunctata differs in the first distal blotch (m4) located more apically, parallel to the costal margin, and separated from it by only a very narrow ochraceous brown stripe (Fig. 3D). Female unknown.

Distribution.

(Fig. 16B) Endemic to the Tsaratanana massif in northern Madagascar (DIANA Region). Up until now known only from the type locality “below Andohanisambirano”. Here, the species is recorded from two further localities in the southern part of Tsaratanana: Andohananalila and Andohanambatoafo (see discussion). The information about the occurrence of T. quinquepunctata in the Marojejy massif provided in Przybyłowicz and Wiorek (2023) resulted from a mistake and is corrected here. At 1850–2050 m; the elevation of the type locality, in the original labels and description (Griveaud 1967) given at 1900 m, should be corrected to 2050 m (Viette 1967, 1991). The latter is equivalent to a campsite by a seasonally dry lake (see below) at –14.1524, 48.958, where DL camped.

Ecology.

Attracted to light (UV) at night, there is no explicit data on diurnal activity. Recorded in March and December. Associated mostly with humid, midland and montane evergreen forests. The type series was collected in a “matsabory”, a marsh-like open area, periodically drying out and turning into a pond in the rainy season (Viette 1967; Diakonoff 1970), with bamboo thickets nearby (Viette 1967). Such bamboo-dominated habitats are found in Tsaratanana at 2000–2200 m (Goodman et al. 2018), and the species may potentially be associated with this kind of vegetation. It might also dwell in sclerophyllous forest vegetation, occurring in Madagascar above 1300 m on the slopes open to winds (Wells 2003), and in Tsaratanana reaching to 2500 m (2000 m in the other massifs of Madagascar) (Crowley 2004). The occurrence of T. quinquepunctata in ericoid thickets is less probable, as this type of vegetation, although usually commencing around 1800–2000 m elevation (Du Puy and Moat 1996; Crowley 2004), in Tsaratanana occurs above the known elevational range of the species and is currently highly degraded (Goodman et al. 2018).

Tritonaclia tollini (Keferstein, 1870)

Figures 6, 7G, 9, 13, 16C, 17A, B, 18A–C, 19

Glaucopis tollini Keferstein, 1870: 13–14, fig. 3

Tritonaclia melania (Oberthür, 1923) syn. nov.

Naclia melania Oberthür, 1923: 134, pl. DLXVI [566], figs. 4880 [♂], 4881 [♀].

Tritonaclia melania Griveaud 1964: 78, figs. 173–176, pl. I, figs. 47, 48; Viette 1991: 170.

Material.

24 specimens, including lectotype and two paralectotypes of G. tollini, and lectotype of N. melania (ISEA PAS - 3 ♂♂, 2 ♀♀; MLUH-ZNS - 3 ♂♂; MNHN - 5 ♂♂; NHMUK - 6 ♂♂, 1 ♀; PZBT - 4 ♂♂) (Table S1).

Types.

Lectotype (of tollini Keferstein, 1870): 1 ♂; [no label], collection data based on the original description: Madagascar, “Tamatave” [Toamasina]; 02 Aug. 1862; C. Tollin leg.; MLUH-ZNS. — Paralectotypes (of tollini Keferstein, 1870): 2 ♂♂; [no label], collection data based on the original description same as for lectotype; MLUH-ZNS, designated here. —

Lectotype (of melania (Oberthür, 1923):

♂; Madagascar, Karianga [doubtful locality]; “sud de Madagascar, 04.1920, Lamberton” NHMUK, NHMUK010620989, designated here.

Diagnosis.

Diagnostic characters are provided in the determination keys, and in the diagnosis of T. vonifotsy Wiorek, sp. nov. (see below). Males of T. tollini are the most similar to T. vonifotsy Wiorek, sp. nov., and to Melanonaclia toulgoeti – for the details, see diagnosis of the former (below). The females of T. tollini and T. vonifotsy Wiorek, sp. nov. (Fig. 7F–I) appear indistinguishable morphologically, even in the genitalia (but see taxonomic note for the latter), and can currently be told apart undoubtedly only using molecular methods (Fig. 2). Griveaud (1964) described the female of T. melania syn. nov. and indicated the female of “T. tollini” (i.e. T. vonifotsy Wiorek, sp. nov.) to be unknown.

Male genitalia redescription.

(Fig. 9) Tegumen well sclerotised, laterally very narrow, medial portion subtrapezoidally widened towards uncus, anterior margin rounded, not incised; uncus short, multidimensional, at base laterally on each side rounded protrusion densely covered with erect setae, medial portion laterally on both sides with distinct rounded incision, terminal portion bifid, with two distinct, lobate protrusions flattened laterally, opened at around 120 degrees, resembling whale tail, covered with creases and serrate along outer margins, entire lateral and dorsal surface of uncus densely covered with elongate, soft setae; anal cone well developed, membranous , without defined scaphium; vinculum very narrow, well sclerotized, incompletely fused with tegumen; saccus subtriangular, dully terminated, straight; juxta in form of well sclerotised ring, on dorsal side submedially with indistinct, subtriangular protrusion or without any additional structures, ventrolaterally on each side with differently expressed, short, lobe-like protrusion, well sclerotised to entirely membranous; valva elongate, of complex shape, reaching beyond uncus tip, costal margin in basal portion slightly incised, straight, distinctly folded inwards, terminal portion transformed into elongate and narrow, claw-like curved and pointed protrusion; sacculus moderately sclerotised to almost membranous, well sclerotised, basal ⅓ straight, then arched, terminal portion in form of elongate, roundly terminated lobe, not reaching to costal tip, almost membranous, heavier sclerotised only along margin, valva outer surface densely scobinate and covered with easy removable scales; phallus weakly to moderately sclerotised, tubular, bulbous in basal portion, terminal portion narrowed and flat cut; vesica left uneverted to avoid destruction, membranous, with well visible structures, in basal portion with two irregular, heavily sclerotised plaques, and elongate, sinuous and coiled row of around 40 heavily sclerotised, densely arranged tooth-like to spine-like cornuti, touching each other laterally.

Distribution.

(Fig. 16C) Northeast Madagascar, mostly around Antongil Bay, in the Regions of: SAVA (Binara, and at Ambavony and Antafononana rivers), Analanjirofo (Lake Amparahibe, south to Antanambe and Anove), and Atsianana (locus typicus “Tamatave” = Toamasina). At around 0–850 m elevation. The lectotype of T. melania syn. nov., labelled as collected in Karianga (SE Madagascar, Region Atsimo-Atsinanana), originates from Charles Lamberton’s materials (“Sud de Madagascar, Reçu de M. Lamberton en avril 1922”), and as such should be treated with caution, considering the general species’ range and the other known cases of Lamberton’s unreliable specimens from “Southern Madagascar” (see Wiorek et al. 2021, 2026). Nevertheless, this locality requires confirmation in light of a similar, disjunctive distribution of T. vonifotsy Wiorek, sp. nov. (see below).

Ecology.

Active by day (specimens provided with detailed label data were caught between 11:00 AM and 5:00 PM, Table S1), there is no data on nocturnal activity or light attraction. Specimens undoubtedly representing the species (i.e. males and DNA barcoded females) were recorded from November to March, in July and August. For information about the phenology of indistinguishable female specimens of T. tollini / T. vonifotsy Wiorek, sp. nov., unavailable for further identification, see the latter species below. Moths are associated with humid, evergreen lowland forest, including riparian habitats. The single specimen from Binara was collected in a gallery forest, in an area generally covered with deciduous, seasonally dry, western forest.

Taxonomic note.

• Hampson (1898) apparently did not examine the original specimens of Keferstein (1870), and most probably did not know where they were deposited. The description in the Catalogue (Hampson 1898) suggests specimens of the species later described as Melanonaclia toulgoeti to be mistaken there for T. tollini, additionally with Melanonaclia lugens (Oberthür, 1893) treated as its synonym (therein as “Thyrosticta tollini ab. 1 lugens”, see also Wiorek et al. 2026). Griveaud (1964) certainly did not know the locality of T. tollini type specimens, as indicated by a question mark (Griveaud 1964: 76). Despite that, he “utilised” the name tollini for the specimens with yellow hindwing medial blotch, even though the said blotches are directly described as “gray” in the original description (Keferstein 1870) – perhaps because the specimens with gray ones had been known to P. Griveaud as Tritonaclia melania syn. nov., described by Oberthür (1923). The Lepidoptera collection of Georg A. Keferstein is currently deposited in the Zentralmagazin Naturwissenschaftlicher Sammlungen at Martin-Luther-Universität Halle-Wittenberg in Halle (Saale), Germany (Schneider and Steinheimer 2018). The original series of T. tollini was rediscovered there by MW in 2022, and here the lectotype and paralectotypes are designated (Fig. 6A–C, F–H). The specimens perfectly match the original description and illustration in Keferstein (1870) (Fig. 6E). They have no labels, but the collection data can be extracted from the first part of Keferstein’s (1870) paper, a logbook of C. Tollin’s expedition to Madagascar. According to that, Tollin arrived at Toamasina [at the time Tamatave] in May 1862, and in the next months conducted excursions in the area. There is no information about him moving to more distant regions of Madagascar, and indeed Viette (1963) indicates that Tollin collected in Foulepointe and Vohidotra villages, both in Toamasina Province. Thus, the specimens were collected there on August 2nd, 1862 [“on August 2nd a new species of the genus Glaucopis”]. All three specimens are males and have well-visible, extending genitalia (Fig. 9F, G), identical to T. melania (Griveaud 1964: figs 174, 175). Thus, we treat Tritonaclia melania syn. nov. as a junior subjective synonym of T. tollini. Consequently, the specimens with a yellow hindwing blotch, interpreted by Griveaud (1964) as “T. tollini”, belong to a separate species Tritonaclia ombilahy Wiorek & Przybyłowicz, sp. nov., described below. • Griveaud (1964) indicated the type series of T. melania syn. nov. to consist of the male holotype and female allotype. However, the original description by Oberthür (1923) is based on one male and three females, with no direct designation of holotype. The single male specimen in question, housed in NHMUK (NHMUK010620989; Oberthür 1923: pl. DLXVI, fig. 4880), has only a “type” label. Thus, here we designate that specimen as the lectotype of T. melania syn. nov.. In the collection of NHMUK, we found only one female (NHMUK013325655), which might have belonged to the original series of one male and three females, and the fate of the remaining two specimens is unknown; they were not found in the NHMUK supplementary collection. Additionally, in the same collection, we found a single, old and worn male specimen (NHMUK016251182), of “Thyrosticta” tollini (i.e. following the generic classification by Hampson 1898), correctly identified as such in Paul Mabille’s handwriting (Horn and Kahle 1935–1937). The specimen is provided with two additional labels indicating it was subsequently obtained by Charles Oberthür in 1923, perhaps after Mabille had passed away in April that year, and eventually deposited in NHMUK in 1927. Therefore, we suppose that Oberthür was unaware of Keferstein’s (1870) description of T. tollini, and he received the specimen after he described T. melania syn. nov. • The male specimen DL_2983 from Binara, in the tree forms a branch sister to all other conspecific individuals (Fig. 2). Although that northernmost locality is relatively remote from the species’ main range (Fig. 16C) and differs in the type of vegetation, we did not find any differences in the genitalia of the specimen. The taxonomic identity of the Binara population requires further studies.

Tritonaclia vonifotsy Wiorek, sp. nov.

https://zoobank.org/8C013D3E-0155-4552-A92A-1258FF4B362F
Figures 7A–F, 10, 14, 16D, 17C, 17D, 18D–F, 20 Specimens of the species in Griveaud 1964: 76–78, figs. 177–179, pl. I, fig. 46, were interpreted as Tritonaclia tollini, subsequently in Viette 1991: 170.

Type material.

Holotype: ♂; Madagascar, Makira Ankirindro; 15.2904 S, 49.5474 E; 636 m a.s.l.; 16 Jan. 2003; D.C. Lees leg.; by cuisine; ISEA PAS, DL_1864, genitalia slide S553, hindwing venation slide S554; BOLD Process ID SYNMD008-26, GenBank COI sequence accession number: PX777880. — Paratypes: • 1 ♂; Madagascar Est, Ambodivoanio; 16.1816 S, 49.6631 E; 50 m a.s.l.; Aug. 1965; A. Peyrieras leg.; “Tritonaclia tollini Keferstein ♂ P. Griveaud det. 1966”; MNHN, MNHN, EL74622, préparation ♂ no. 155; • 1 ♂; [collection data unknown]; “acheté à M. Goudot, 1839”, “Tritonaclia tollini Keferstein ♂ P. Griveaud det.”; MNHN, EL74623, slide P. Griveaud No. 286; • 3 ♂♂; “Madagascar Est, s/préf. Vondrozo, forêt Madiorano, 30 km ouest Vondrozo” [30 km W of Vondrozo, Madiorano Forest]; 22.8065 S, 47.043 E, 750 m a.s.l.; Apr. 1973; A. Peyrieras, A. Rakotoarisolo leg.; MNHN, EL74620, EL74621, EL74624–préparation ♂ no. 156; • 1 ♂; same locality and collector as for holotype; 11 Jan. 2003; NHMUK, NHMUK010918661 / DL1816 / DL-SE11, BMNH(E) 2019-70; BOLD Process ID SYNMD003-26, GenBank COI sequence accession number: MK158587 • 1 ♀; Makira, Ankirindro, ridge, SA6675; 15.2931 S, 49.5472 E, 675 m a.s.l.; 10 Mar. 2003; D.C. Lees leg.; collecting time 2:40 pm; NHMUK, NHMUK010918662 / DL1863, BMNH(E) 2019-70, genitalia slide NHMUK010316589, BOLD Process ID SYNMD005-26, GenBank COI sequence accession number: MK158588. — The type series comprises all known specimens undoubtedly belonging to the species (Table S1).

Other material.

17 indistinguishable females of T. tollini / T. vonifotsy Wiorek, sp. nov., unavailable for further determination with DNA barcoding (MNHN - 7 ♀♀; NHMUK - 3 ♀♀; PZBT - 7 ♀♀) + 1 ♀ record from iNaturalist (Table S1).

Locus typicus.

Madagascar, Makira Ankirindro.

Diagnosis.

The diagnostic characters are provided in the determination keys. The most similar species, Tritonaclia tollini, differs in the colouration of the male hindwing medial blotch (Fig. 17). In males of both species, the outer margin of the forewing is slightly incised, and in spread specimens it tends to crease between the veins CuA2 and 1A+2A (Figs 6A–J, 7A–E, 18A), deepening the incised appearance, and making it similar to Melanonaclia toulgoeti (Wiorek et al. 2026). However, the latter species lacks the hindwing colour medial blotch and terminal bifurcation of the vein 1A+2A, both characteristic for T. tollini and T. vonifotsy Wiorek, sp. nov. (Fig. 18A, B). Females of T. tollini and T. vonifotsy Wiorek, sp. nov. appear morphologically indistinguishable (Fig. 7F–I), but see the taxonomic note below.

Figure 17. 

Wing patches of modified, scent scales in Tritonaclia. A T. tollini hindwing upperside; B T. tollini forewing underside; C T. vonifotsy Wiorek, sp. nov. hindwing upperside; D T. vonifotsy Wiorek, sp. nov. forewing underside.

Figure 18. 

Wing venation, A–C Tritonaclia tollini, D–F T. vonifotsy Wiorek, sp. nov. A Male forewing, frame indicates the area magnified in B; B terminal bifurcation of vein 1A+2A; C male hindwing, note the complete fusion of veins Sc+Rs+M1 and presence of vein 3A.; D male hindwing, slide ISEA PAS, S554, frames indicate areas magnified in E and F; E terminal bifurcation of veins Sc+Rs+M1; F basal, vestigial bifurcation of 3A.

Male description.

(Fig. 7A–E, 18D–F) Head. Proboscis well developed, ochraceous yellow; head entirely brown to dark brown, almost black; ventroterminal portion of postocular area provided with tuft of protruding scales; labial palpus 3 segmented, porrect, around ½ protruding beyond frons, 1st palpomere with elongate scales directed downwards, 2nd palpomere 2× longer than 1st one, 3rd palpomere of the length of 1st one; antennae entirely brown, piliform, ventrally densely covered with short erect setae, additionally each antennomere on both margins with pair of distinct setae, one longer and arched, one shorter and straight; Thorax. Patagia inner side brown to almost black, medially with elongate scales, lateral side with pure with blotch of regular scales; tegulae entirely brown to almost black, terminal portion with tuft of long, piliform scales; thorax entirely brown to almost black; Legs. Entirely brown to almost black, in some specimens with admixture of single, creamy to white scales on frontal surface of hindleg coxa, and sometimes also on midleg coxa; arolium present; Foreleg. Tibia of ⅔ of femur length, epiphysis present, of ⅔ of tibia length, ochraceous yellow; Midleg. Tibia with one pair of brown terminal spurs; Hindleg. One pair of brown terminal spurs, and one pair of additional spurs in ¾ of tibia length, shorter than terminal ones; Abdomen. Entirely brown to almost black, tergites 4th–6th lateroterminally with well developed, pure white blotch, in some specimens admixture of white scales also on 7th tergite; sternites 2nd–6th medioterminally with differently expressed, well developed, pure white blotch, sometimes with admixture of single creamy scales, sternite 7th medially with only small white dot, sternite 8th without markings, with deep U-shaped incision, reaching to around ½ of 8th tergite length; lateral portion of abdomen unavailable; valvae distinctly protruding from abdomen, outer surfaces densely covered with scales concolorous with abdomen; Forewing. Subtriangular, hind margin rounded, without well-defined tornus; backround brown to almost black, with 5 or 4 pure white, semi-transparent blotches of different size: one or two in medial portion and three in distal portion; first medial blotch (m2) subsquare to subtrapezoidal, in medioterminal portion of DC, between from R and DC hind margin (touching both); second medial blotch (m3) medially between base of CuA2, and 1A+2A, in form of subtriangular to rounded dot, very small and indistinct, up to completely absent; first distal blotch (m4) subsquare to suboval; second distal blotch (m5) relatively small, rounded to irregularly suboval; third distal blotch (m6) the largest one, always much larger than m5, elongate suboval to subtrapezoidal; wing underside as upperside, but basal and medial portion covered with pale area of dull-white regular scales, along tornus and hind margin reduced in size; medially above 1A+2A elongate, suboval field of pale yellow-creamy scales, terminated before CuA2; 1A+2A in terminal portion distinctly bifurcated; Hindwing. Suboval to irregularly subtrapezoidal, reaching to ½ of forewing length, costal margin in medial portion arched and widened, outer margin slightly sinuous, medially with indistinct incision; background along outer and hind margins brown to almost black, medial part covered with dull-white, in DC with additional, elongate, irregular blotch of no well-defined margins, covered with regular, elongate, pale yellow-creamy scales; basal blotch in basodistal portion of wing, along DC hind margin and CuA2, separated from basal part of terminal margin by narrow stripe of brown scales, heart-shaped, slightly semi-transparent, covered with pure white scales of slightly reduced size; first distal blotch beyond outer margin of DC, most often hardly visible on upperside; second distal blotch suboval, of moderate size, medially between CuA1 and M2+M3, covered with slightly reduced, pure white scales; underside with entirely brown to almost black background, basal blotch as on upperside, covered with mixture of regular and slightly reduced pure white scales, first distal blotch well defined, suboval, covered with regular and reduced pure white scales, beyond DC outer margin and below Sc+Rs+M1 (touching); second distal blotch as on upperside, with mixture of regular and reduced pure white scales; additionally, in some specimens above terminal portion of M2+M3 additional, indistinct dot of a few white scales; M2 and M3 fused in around ½ of their length, DC outer margin convex, Sc+Rs+M1 completely fused, but in some specimens slightly bifurcated in the terminal portion; 3A present, in some specimens in basal portion with vestigial, lateral arm.

Male genitalia description.

(Fig. 10) Tegumen well sclerotised, moderately wide, in medial portion subtriangular projected towards uncus, both margins with narrow, rib-like sclerotisation all along; uncus multidimensional, of variable length, from elongate, with subtriangular basal portion, reaching to valva tip, to short and reaching to ⅔ of valva length, basally always provided with subconical, curved lateral protrusion densely covered with short erect setae, terminal portion Y-shaped bifid, with two distinct, lobate protrusions flattened laterally and opened at around 90 degrees, in terminal portion creased, with serrate outer margin; terminal half of uncus laterally covered with elongate, erect setae; anal cone well developed, subconical, originating at basal portion on uncus, with very indistinct scaphium formed by moderate sclerotisation of dorsal wall, sometimes absent; vinculum narrow, moderately to well sclerotized, incompletely fused with tegumen; saccus subconical, dully terminated, projected slightly anteriorly; juxta well sclerotised, in form of ring, lateroventrally provided with differently expressed lobes, entirely or partially sclerotised, dorsoposteriorly provided with two straight, spine-like pointed protrusions of moderate length, of equal or different lengths, ventral portion of juxta distinctly bent posteriorly and touching basal portion of valvae; valva elongate, of complex shape, reaching at least uncus tip, costal margin straight, distinctly folded inwards, terminal portion transformed into spatula-like, slightly curved and roundly to subtriangularly terminated protrusion; sacculus well sclerotised, basal ⅓ arched, terminated with straight, slightly narrowing and pointed protrusion directed inwards and reaching to around ½ of valva length and width, terminal portion lobate, almost membranous, heavier sclerotised only along margin, roundly terminated and almost reaching costal tip; valva outer surface densely scobinate and covered with easy removable scales; phallus well sclerotised, moderately short and broad, slightly irregularly curved, terminal portion narrowed, cylindrical and flat cut; vesica left uneverted to avoid destruction, but with well visible structures, in basal portion with two irregular, heavily sclerotised lobes, and elongate, sinuous row of around 25–30 heavily sclerotised, densely arranged tooth-like to spine-like cornuti, touching each other laterally;

(Re)description of the female of T. tollini and T. vonifotsy Wiorek, sp. nov.

(Fig. 7F–I) Head. (including eye size), thorax, legs, and abdomen colouration and pattern as in the male, but see the comment on the foreleg coxa in the taxonomic note below. Forewing. Subtriangular, slightly more elongate and with more prominent tornus than in male; background brown to almost black, with five pure white, semi-transparent blotches of different size, arranged as in male, second medial blotch (m3) differently expressed, round to suboval but always present; third distal (m6) slightly larger than in male; underside as upperside, with no additional colour blotches; 1A+2A in terminal portion bifurcated as in male; Hindwing. Shape and size similar to male, uniformly brown to almost black, with three or four differently expressed, pure white, semi-transparent blotches, and no additional marks or blotches of different colour; basal blotch as in male; first distal blotch beyond outer margin of DC, variable in shape and size, from small round dot to elongate, S-shaped; second distal blotch in form of small dot between bifurcating M2+M3, in some specimens absent; third distal blotch as in male, between CuA1 and M2+M3; underside as upperside, but basal and first distal blotches slightly enlarged; venation as in male, in a single female vein 3A asymmetrically developed, bifurcated in left hindwing.

(Re)description of female genitalia.

(Figs 13, 14) Anal papilla subtrapezoidal, densely scobinate and covered with elongate, erect, soft setae of different lengths; pseudopapilla narrow, membranous, reaching to valva terminal margin, densely scobinate; apophysis posterioris slightly longer than anal papilla, well sclerotized, needle-like, pointed; apophysis anterioris slightly shorter, needle-like, originating from subtriangular lobe; dorsal pheromone glands in form of two narrow, membranous, densely entangled and furcate tubes; ventral pheromone glands absent; lamella postvaginalis well defined, subtrapezoidal, terminal margin densely scobinate, medioterminally with shallow, rounded incision, lateroterminally on each side with differently expressed, rounded protrusion covered with small grain-like sculpture and elongate, erect setae, absent in some cases; sternite 7th medially with deep, U-shaped incision; ostium bursae distinct, wide, rounded to suboval, antrum short, cup shaped; tergite 7th enlarged, of the length of sternites 7th+8th, tergite 8th reduced to narrow plaque, with heavier sclerotised, rib-like basal margin, laterally fused with lateral arms of lamella postvaginalis and forming subtriangular base of apophysis anterioris; ductus bursae tubular, of uniform width, well sclerotised and distinctly convoluted rightwards into three coils; Cervix bursae of around ½–⅓ of corpus bursae length, of width of ductus bursae, slightly curved, entirely membranous with dense longitudinal plicae; corpus bursae suboval, membranous, with dense minute scobinations and moderately wide longitudinal plicae;. Signum bursae located medially, in form of two horizontal, 8-shaped, well sclerotised plaques arranged longitudinally in ⅓ and ½ of corpus bursae length, covered with short, subconical, spine-like protrusions; ductus seminalis in form of narrow, membranous tube originating from lateral diverticulum located at base of corpus bursae, of shape and plication similar to cervix bursae and parallel to it.

Etymology.

From Malagasy vonifotsy, meaning “pale yellow”, referring to the colouration of the male hindwing medial blotch.

Distribution.

(Fig. 16D) Mostly in the northeast of Madagascar, in the Analanjirofo Region (Ankirindro and Ambodivoanio). Three male specimens (Fig. 7D, E, 10C–E) from Madiorano Forest in the area of Vondrozo, Ihorombe Region, collected by A. Peyrieras and A. Rakotoarisolo in April 1974, although distinctly distant from the remaining localities, leave little doubt. At 50–750 m elevation. All remaining, unidentified females of T. tollini / T. vonifotsy Wiorek, sp. nov., unavailable for further identification, were collected or observed in the northeast part of the island, in the Regions of SAVA (at Antafononana River), Analanjirofo (from Ambodivoangy and Fampanambo, south to Antanambe), and Atsinanana (Forêt d’Analalava). At around 25 to max. 1000 m elevation (the highest possible point of Antampona). Associated with humid, evergreen lowland forest.

Ecology.

Specimens undoubtedly belonging to the species (males and one DNA barcoded female) were collected in January, March, April and August. Additionally, the unidentified, non DNA-barcoded females of T. tollini / T. vonifotsy Wiorek, sp. nov. were recorded in January, February, April, June, July and November.

Taxonomic note.

Tritonaclia vonifotsy Wiorek, sp. nov. exhibits the above-described distinct differences in the male uncus length (Fig. 10D–F). The specimen from the remote, southernmost locality of Madiorano Forest has an uncus of intermediate length (Fig. 10E). This character requires further research, including molecular examination, which was limited in our study. The rounded, setaceous protrusions of the female lamella postvaginalis seem to be present in T. vonifotsy Wiorek, sp. nov. and absent in T. tollini (Figs 13A–C, 14B). However, we had only a single DNA barcode-confirmed female genitalia preparation available for each of the two species, whilst the character in question is potentially prone to destruction during dissection. Further, both species seem to stably differ in the very indistinct admixture of pale creamy to white scales in the proximomedial part of the foreleg coxa, allegedly present in both sexes of T. tollini and absent in T. vonifotsy Wiorek, sp. nov., including the three females DNA barcoded in our study (not shown). Although potentially the only morphological character distinguishing the females of both species, it is a very subtle trait requiring further research on a longer series of DNA barcoded specimens. Therefore, here we refrain from interpreting it as a diagnostic character.

Tritonaclia stephania (Oberthür, 1923)

Figures 8A–C, 11, 16E

Naclia stephania Oberthür, 1923: 135, pl. DLXVI [566], fig. 4882;

Tritonaclia stephania Griveaud 1964: 79, figs 185–187, pl. I, fig. 49; Viette 1991: 170; Wiorek et al. 2021: 20, figs. 2B, 3G, 5F.

Materials.

43 specimens, including holotype (ISEA PAS - 1 ♂; MNHN - 21 ♂♂; NHMUK - 1 ♀; PZBT - 19 ♂♂, 1 ♀) (Table S1).

Holotype.

♀, NHMUK, NHMUK010620988, by monotypy (in Griveaud 1964 erroneously as male lectotype).

Diagnosis.

Superficially, the most similar species is Tritonaclia ombilahy Wiorek & Przybyłowicz, sp. nov., described below. The diagnostic characters are provided in the determination keys. — Male genitalia were redescribed in Wiorek et al. (2021).

Distribution.

(Fig. 16E) Longitudinal range along the central east of Madagascar, in the Regions of Alaotra-Mangoro (Vallée d’Ivelona, Ambatondrazaka, Forêt Ambohiboatavo), Analamanga (RS d’Ambohitantely, La Mandraka), Vakinankaratra (Ampolomita), Matsiatra Ambony (Ankafina-Tsarafidy), Ihorombe (Forêt Vakoany). At around 1000–1600 m elevation.

Ecology.

Attracted to light at night, there is no information about diurnal activity. Collected from September to March, in May and July. Associated with evergreen, humid midland forest.

Taxonomic note.

Griveaud (1964) indicated the type specimen of T. stephania (NHMUK010620988), deposited in NHMUK (at the time BMNH), to be a male lectotype, but in 2021 ŁP identified it to be a female. Further, there is no information about the number or sex of the type series specimens in the original description by Oberthür (1923). For the five other Syntomini species described in Oberthür (1923), numbers of specimens larger than one are specifically indicated, and generally congruent with the materials currently housed in NHMUK (verified by MW, but see T. melania syn. nov. above). This, by analogy, implies that the abovementioned specimen should have been interpreted as a holotype by monotypy. The only superficially most similar specimen in the collection (NHMUK013702406) belongs to Tritonaclia ombilahy Wiorek & Przybyłowicz, sp. nov. described below. It can be excluded as the potential second specimen from the original series (alleged “paralectotype”), since in the original description Oberthür (1923) directly indicates the small size of T. stephania, whilst T. ombilahy Wiorek & Przybyłowicz, sp. nov. is distinctly larger. • The female “neallotype” of T. stephania (MNHN, EL65139, genitalia slide by P. Griveaud, No. 291), designated by Griveaud (1964), is distinctly larger than the male (Fig. 8A, G). In 2022, ŁP and MW found in MNHN three male specimens with a body size similar to that female, collected in 1970 and 1971, thus after Griveaud’s (1964) “Amatidae” monograph. These specimens turned out to represent a new species, Tritonaclia ombilahy Wiorek & Przybyłowicz, sp. nov., described below. Based on the distinct and discrete wingspan difference between both species, with almost no intraspecific variation, we assign the female “neallotype” of T. stephania to T. ombilahy Wiorek & Przybyłowicz, sp. nov. • During the visit to PZBT in 2023, ŁP found a single female specimen of T. stephania in poor condition (not shown). The specimen, as well as the female holotype in NHMUK, was unavailable for dissection, and the female genitalia of the species remain unknown.

Tritonaclia ombilahy Wiorek & Przybyłowicz, sp. nov.

https://zoobank.org/9C0B6CC5-6A35-45E7-84EC-0EA7492D9BD1
Figs 8D–G, 12, 15, 16F Specimens of the species in Griveaud 1964: 80, fig. 184, pl. I, fig. 50 were identified as Tritonaclia stephania.

Type material.

Holotype: ♂; “Madagascar Est, N.O. de Manantenina, Chaines Anosyennes, massif nord” [NW of Manantenina, Anosyennes Mountains, northern massif]; 24.1134 S, 47.108 E, 1050 m. a.s.l.; 22–29 Nov. 1971; P. Griveaud leg.; “Museum Paris, Madagascar Centre mission C.N.R.S.R.C.P. no. 225”; MNHN, EL74649, préparation ♂ no. 157; Paratypes: • 1 ♂; same collection data as for holotype; MNHN, EL74647; • 1 ♂; Ambohimanakana, Manambato, Anova [=Anove]; 16.6055 S, 49.798 E, 10 m a.s.l.; 26 Oct.–4 Nov. 1970; P. Viette, P. Griveaud leg.; “Museum Paris, Madagascar Centre mission C.N.R.S.R.C.P. no. 225”; MNHN, EL74648; • 1 ♀; Madagascar, Fianarantsoa [doubtful locality]; [collection date and collector unknown]; “ex Lamberton 1922”; originally designated by P. Griveaud (1964) as “neallotype” of Tritonaclia stephania; MNHN, EL65139, slide P. Griveaud No. 291; • 1 ♀; Madagascar, Fianarantsoa [doubtful locality]; [collection date and collector unknown]; “ex Lamberton 1922”; “Ex Oberthür Coll., Brit. Mus. 1927-3”; NHMUK, NHMUK013702406. — The type series comprises all known specimens of the species (Table S1).

Locus typicus.

NW of Manantenina, Anosyennes Mountains, northern massif.

Diagnosis.

Superficially, the most similar but distinctly smaller species is Tritonaclia stephania. The diagnostic characters are provided in the determination keys.

Description.

(Fig. 8D–G) Head. Entirely pale brown, proboscis well developed, ochraceous yellow; labial palpus 3 segmented, elongate, porrect and slightly curved upwards, with around ½ of the length protruding beyond frons; 1st palpomere short, ventrally covered with elongate scales, 2nd palpomere around 2× longer than 3rd palpomere; antennae piliform, ventrally densely covered with short erect setae, additionally each antennomere on both margins with pair of setae, one elongate and arched, another one much shorter and straight; Thorax. Patagia pale brown, with elongate scales, laterally with small dot of clear white regular scales; tegulae entirely pale brown, with elongate scales, and piliform scales in terminal portion; mesothorax and metathorax entirely pale brown; Legs. Entirely pale brown, tarsal claw with additional tooth; Foreleg. Tibia ⅔ of femur length, epiphysis present, of ⅔ of tibia length, tibia and 1st tarsomere together slightly longer than femur; behind coxa paintbrush-like tuft of fine, elongate, creamy scales. Midleg. Tibia with one pair of pale brown terminal spurs; Hindleg. Tibia with one pair of pale brown terminal spurs, and one pair of additional spurs in ¾ of tibia length, of around ⅔ of length of terminal ones; Abdomen. Pale brown, tergites 4th–6th lateroterminally with differently expressed clear white to pale creamy marks on each side, from fully developed blotch to admixture of a few scales; in some specimens white scales present also on tergite 7th; sternites – 6th medioterminally with differently expressed clear white blotch, in terminal segments usually reduced to a few scales; in form of scales admixture; 8th sternite reduced; abdomen laterally unavailable; valvae protruding and well visible, outer surfaces covered with pale brown scales; Forewing. Subtriangular, slightly elongate; background pale brown, with 5 white blotches, two in medial and three in distal wing portion, of various shape and size, all semi-transparent, covered with smaller and scarce scales; first medial blotch (m2) small, subtriangular to suboval, in terminal portion of DC, from R (touching), terminated in half of DC width, in some specimens distinctly reduced, almost absent, in form of small, but always semi-transparent dot; second medial blotch (m3) medially between CuA2 and 1A+2A, smallest one, always in form of indistinct dot; first distal blotch (m4) of moderate size, suboval to round; second distal blotch (m5) always relatively small, rounded to elongate, in some specimens very indistinct; third distal blotch (m6) always the largest one, rounded to irregularly subtrapezoidal; Underside as upperside, but in medial and hind portions almost entirely covered with small, scarce, grain-like scales, giving entire wing slight semi-transparent appearance; hind margin along basal portion with tuft of piliform scales; retinaculum subcostale present, narrow and elongate; Hindwing. Oval, elongate, reaching to ⅔ of forewing length; background pale brow with 1 or 2 white semi-transparent blotches of different shape and size; basal blotch from wing base, between DC hind margin and basal portion of wing hind margin, irregularly pear-shaped, slightly incised in basal and terminal portion along vein 1A+2A, separated from wing margin by very narrow pale brown stripe; distal blotch between CuA1 and M3, in form of relatively small, suboval dot, in some specimens very indistinct and reduced up to just a few scales visible only under microscope; underside as upperside; M2 and M3 fused in basal portion but at different length, Sc+Rs+M1 completely fused;

Description of male genitalia.

(Fig. 12) Tegumen moderately sclerotised, arms laterally narrow, submedially provided with irregularly subtrianagular, slightly curved lateral lobe covered with tuft of elongate, erect setae; uncus basally broad, then distinctly narrowed, rod-like, arched downwards, in terminal portion slightly widened laterally and dully terminated, dorsally with indistinct, rounded incision, ventrally with sharp horizontal edge; vinculum narrow, moderately sclerotized, incompletely fused with tegumen; saccus indistinct, in form of subtriangular plaque; juxta particularly well developed and heavily sclerotised, in form of slightly suboval ring, ventrally connected with membrane provided with two moderately sclerotised, shield-like plaques, dorsolaterally armed with prominent, sinuous, bull-horn-shaped protrusions directed posteriorly, asymmetrical, right one slightly longer and reaching to around ½ of uncus length; valva elongate, of complex shape, not reaching beyond uncus, costal margin distinctly folded inwards, in basal portion shallowly incised, terminal portion transformed into well sclerotised, claw-like, pointed protrusion, sacculus well sclerotised, distinctly arched, in terminal portion provided with membranous, lobe-like, densely scobinate protrusion, roundly terminated, not reaching to valva tip, covered with elongate erect setae; phallus short and broad, narrowed in terminal half, coecum penis rounded; vesica left uneverted to avoid destruction, but with well visible structures, basal portion with two heavily sclerotised, subtriangular plaques with elongate and slightly twisted terminal portion, along vesica sinuous row of 15 distinct, heavily sclerotised subconical teeth flattened dorsoventrally and leaning one on another;

Female genitalia.

Figure 15 were described and illustrated as T. stephania in Griveaud (1964: 80, fig. 184).

Etymology.

From Malagasy ombilahy meaning “bull”, after the shape of the dorsal spines of the male juxta, resembling the horns of zebu cattle commonly bred in Madagascar.

Distribution.

(Fig. 16F) Mountain massifs of southeastern Madagascar: Andringitra, at the border of the Regions Matsiatra Ambony and Ihorombe, and the Anosyenne Mountains, at the border of the Regions Anosy and Atsimo-Atsinana. Fianarantsoa (Region Matsiatra Ambony), as the collecting locality of two specimens from C. Lamberton, is doubtful (see similar cases above, and Wiorek et al. 2026). At 1050–1650 m elevation.

Ecology.

Specimens were collected in October and November. Associated with evergreen, humid midland forest.

3.4. Scent scales patches of Tritonaclia tollini and T. vonifotsy Wiorek, sp. nov.

Males of Tritonaclia tollini and T. vonifotsy Wiorek, sp. nov. have characteristics, suboval patches of transformed scales on the forewing underside and hindwing upperside, distinct from the pale colouration of the surrounding areas (Fig. 17). Compared to the regular, suboval scales of the wing background, the transformed ones are elongate (approximately 2–3× longer), more densely arranged (i.e. the wing membrane is invisible), and subrectangular in the terminal portion (Figs 17, 19, 20). In each species, the general morphology of the scales of both fore- and hindwing patches is similar, but the forewing scales seem to be shorter. In T. vonifotsy Wiorek, sp. nov., the hindwing scales are around 250–325 μm long. The forewing scales are around 225 μm long, in the basal portion spindle-like, widened, and narrowed around the terminal ⅔, in the terminal portion subrounded to subrectangular with rounded corners (Fig. 20). In T. tollini, hindwing scales are around 425 μm long, and forewing scales are around 300 μm long, more spatula-like, with the basal portion regularly rounded, and without distinct narrowing towards the terminal portion (Fig. 19). In both species, the outer surface of the lower lamina of a scale (‘adwing’) is creased, with indistinctly marked longitudinal ridges but with no windows (Fig. 20B, C (larger scale), 19C). The outer surface of the upper lamina (‘abwing’) is covered with distinct, parallel, longitudinal ridges. Interridge areas seem to be broader in T. vonifotsy Wiorek, sp. nov. than in T. tollini (around 1.8–2.6 μm vs. 1.5–1.6 μm, respectively). In both species, the interridge area is covered with transverse, arched, differently anastomosing and fusing microribs (flutes), surrounding subrounded windows, varying in shape and size, and partially closed with perforated, membranous structures. The windows seem to be more numerous and larger in T. vonifotsy, Wiorek sp. nov. than in T. tollini (Figs 19B, 19D, 20D, 20E). The internal cavity of the scales is open, formed by mesh-like arranged trabeculae (Fig. 20B, E, F).

Figure 19. 

SEM visualisation of male wing scent (androconial) scales in Tritonaclia tollini. A, B Upper lamina of forewing underside scale (in A basal part of scale in bottom right); C, D upper lamina of hindwing upperside scale (in C basal part of scale in bottom right).

Figure 20. 

SEM visualisation of male wing scent (androconial) scales in Tritonaclia vonifotsy Wiorek, sp. nov. A Upper lamina of forewing underside scale (basal part in top right); B inner structure of forewing underside scale; C hindwing upperside scale, upper lamina in top left (basal part of scale bottom left), lower lamina in bottom right (basal part of scale top right); D magnification of upper lamina of hindwing upperside scale; E cross section of hindwing upperside scale; F magnification of inner structure of hindwing upperside scale.

4. Discussion

Our molecular results confirm not only the monophyletic character of Tritonaclia in the arrangement proposed here, but also that the genus is not closely related to the other lineages of black-and-white Madagascan Syntomini, represented in the tree. Tritonaclia is placed as sister to the clade formed by Dubianaclia Griveaud, 1964, Thyrosticta Hampson, 1898 (in the shape revised in Wiorek et al. 2026) and Mauricenaclia Wiorek, 2026. This topology is congruent with, and supplements the results presented in Wiorek et al. (2026: fig. 6A, B). In that study, the black-and-white taxa represented here by single samples do not form a monophyletic group but are placed in different clades across the tree. This raises questions about the evolution of wing and body patterns in Madagascan Syntomini. To address these questions, it is crucial to infer a robust and possibly complete phylogenetic tree of the group. A reconstruction of the ancestral states of particular characters could help to address the issue in the future, especially since the existence of mimicry rings within Syntomini has been suggested (Wiorek et al. 2026).

Among the six species of Tritonaclia, only T. tollini and T. vonifotsy Wiorek, sp. nov. are typically lowland, occurring up to around 750–850 m elevation. The remaining four species are associated mostly with higher, midland and lower montane environments, with T. quinquepunctata being the most “montane” and reaching up to 2050 m.

Of those studied, there are four main mountain massifs of Madagascar which exhibit well-developed vegetational belts: Tsaratanana and Marojejy in the north, Ankaratra in the centre, and Andringitra in the south (Crowley 2004; Clark et al. 2025). The first two seem to be the most important areas of Syntomini biodiversity in the island in general; the last two are not or no longer contiguous. Amongst these, Tsaratanana is the most remote, difficult to access, and poorly studied mountain region of Madagascar (Clark et al. 2025), despite a relatively long history of exploration and a series of scientific expeditions conducted (Viette 1967; Albignac et al. 1970).

This is also reflected in prior uncertainty regarding the locality of the two southern collecting places of T. quinquepunctata in Tsaratanana: Andohanambatoafo and Andohananalila. Significant numbers of new species were described from both, not only in Lepidoptera, but also e.g. Coleoptera (Basilewsky 1985). However, their exact locality has remained unknown, also after the period of activity of French entomologists on the island (e.g. Krüger 2001). Apparently, both localities were visited only once, by Pierre Soga, during the expeditions to Tsaratanana in December 1966 and March 1967 (Viette 1967; Lacroix and Viette 1998; Viette 1991). Unfortunately, there are no logbooks, nor even a comprehensive list of Soga’s collecting localities and trips (see Wiorek et al. 2026). The prefix andohana, present in the names, literally means “at the head of”, and in Malagasy geographical names can also mean “the limit of” (L. Randriantseheno, pers. comm.). More precise information about Andohananalila is provided in Albignac et al. (1970: 91). It is located “above the river Ananalila, NW of Mangindrano”, and the elevation 1850 m corresponds with the local highest point above (“andohana”) the river’s headwaters. Andohanambatoafo is an analogous case, being located over the headwaters of the river Ambatoafo, NE of Mangindrano, on the map in Albignac et al. (1970: Carte 2), spelt as “Amboatoafo”. This place is apparently located close to the botanical collecting point in the area (e.g. Johnson and Murray 2020), spelt as “vallon d’Ambatohafo” [“valley of Ambatohafo”]. Therefore, both localities in question are located on the sides of the southern approach path to the Tsaratanana summit that passes through the village of Mangindrano (Albignac et al. 1970).

Tsaratanana massif is known to harbour a unique fauna of Syntomini, of which all species, including T. quinquepunctata, are local endemics. It comprises a group of Maculonaclia species: M. bicolorata Griveaud, 1967, M. flamea Griveaud, 1967, M. matsabory Griveaud, 1967, M. obscura Griveaud, 1967, M. petrusia Griveaud, 1967, and also Soganaclia roedereri Griveaud, 1971, S. tsaratananae Griveaud, 1971, and Tsirananaclia formosa Griveaud, 1973. Based on the genitalia illustrated in Griveaud (1967), all Maculonaclia species listed above are most probably closely related to each other and may derive from local speciation events. Additionally, in the region of Tsaratanana occurs Fletcherinia decaryi Griveaud, 1964, the only Madagascan syntomine belonging to the “Amata clade” of the tribe, thus not closely related to the Madagascan radiation, and apparently deriving from a separate episode of island colonisation (Griveaud 1964; Przybyłowicz et al. 2019).

In contrast to Tsaratanana, the neighbouring Marojejy massif is one of the richest and relatively well-explored biodiversity hotspots in Madagascar (Goodman et al. 2023a). This applies also to Lepidoptera, which were collected there in many different localities at different elevations (Viette 1991; see also Wiorek et al. 2026), including the expeditions of DL. The total number of Syntomini recorded there so far is 23 species (Griveaud 1964; Wiorek et al. 2026). This number nearly equals a quarter of the Syntomini species count for Madagascar, and, as far as we know, it is the highest number of species known from a single area in the island. Additionally, eight of them are considered local endemics: Maculonaclia truncata Griveaud, 1964, Stictonaclia marojejyensis Griveaud, 1964, S. subflava Griveaud, 1964, Privatenaclia seguyi (Griveaud, 1964), P. triangulifera (Griveaud, 1964), and Tsirananaclia sucini Griveaud, 1964 (Griveaud 1964; Wiorek et al. 2026).

Furthermore, the southernmost located Andringitra massif is the only locality for the sympatric occurrence of T. stephania and T. ombilahy Wiorek & Przybyłowicz, sp. nov. In total, six Syntomini species are known from that massif (Griveaud 1964, 1974; Wiorek et al. 2026), including two local endemics: Tsarafidynia blanci Griveaud, 1974 and Micronaclia imaitsia Griveaud, 1964.

Only in the fourth “main” massif, Ankaratra, no species of Tritonaclia – nor, to our best knowledge, any Syntomini species – have ever been reported. Skippernaclia ankaratra (Griveaud, 1964) does not apparently occur in the massif, but rather in localities around it (Wiorek et al. 2026). This seems to result rather from negligence in studies of the region, except for single studies of Microlepidoptera (Viette 1956), rather than absence of the group, since, e.g., Tritonaclia stephania occurs in RS d’Ambohitantely in the Central Highlands, located around 140 km N from Ankaratra (Wiorek et al. 2021). The fauna of Ankaratra is generally poorly studied, and its vegetation is already highly degraded, with little natural forest left (Goodman et al. 2018).

Therefore, further analyses based on a dated phylogeny of Madagascan Syntomini and possibly complete distributional data are necessary to explore the evolutionary history of the group, and the role of the mountain massifs of Madagascar in their diversification, in the context of mechanisms proposed so far (Vences et al. 2009).

The species covered in this paper have interesting phylogeographies, as far as they have been sampled. Tritonaclia comprises three pairs of sister species: T. kefersteinii + T. quinquepunctata, T. tollini + T. vonifotsy Wiorek, sp. nov., and T. stephania + T. ombilahy Wiorek & Przybyłowicz, sp. nov., all characterised by different distributional patterns. The distribution of T. kefersteinii was initially discussed in Przybyłowicz and Wiorek (2023), with the suggestion of climate change driven shifts of vegetational ranges, which widened the distribution of vegetation currently associated with higher elevations, potentially facilitating the species’ dispersal. Tritonaclia kefersteinii is a more lowland species, thus, in suitable conditions, it could become more widespread, ranging from Marojejy to Ankafina-Tsarafidy and at present disjunctively distributed. In contrast, T. quinquepunctata is a higher-elevation species, restricted to the massif of Tsaratanana. Intriguingly, although the massifs of Marojejy and Tsaratanana are close to each other and still connected by the forest corridor referred to as COMATSA (Marojejy-Anjanaharibe-Sud-Tsaratanana: Rabearivony et al. 2015), T. kefersteinii does not occur in Tsaratanana, but its range is extended longitudinally. Tritonaclia stephania is one of the most widely distributed Tritonaclia species, occurring in most of the central east of Madagascar. In contrast, its sister species, T. ombilahy Wiorek & Przybyłowicz, sp. nov., is known from two localities in the southeast of the island: Andringitra and the Anosyenne Mountains. Being located in the southernmost part of the “eastern” belt of humid forests, these places are close to the transition zone with subarid biotopes of southern Madagascar (Moat and Smith 2007) and have lower annual precipitation (1477 mm in Andringitra vs. 2066 mm in Ranomafana: Goodman et al. 2018). These localities are also characterised by the presence of some “southern” faunistic elements, such as the Ring-Tailed Lemur (Goodman et al. 2023b). Thus, we hypothesise that T. ombilahy Wiorek & Przybyłowicz, sp. nov. may be more widespread and present also, e.g. in the “lost forest” of Ivohiboro, a neglected yet highly diverse forest patch in SE Madagascar (Otero Jimenez et al. 2023). Conducting formal niche modelling may help to address the issue properly in the future. Such analyses have already indicated precipitation as the key factor explaining insect (Coleoptera) distribution in southern Madagascar (Kamiński and Raś 2012).

Tritonaclia tollini and T. vonifotsy Wiorek, sp. nov. are sympatric species, occurring mostly around the Antongil Bay. The male wing patches of scent (androconial) scales are reported in these species for the first time, not only in Madagascan Syntomini, but – to the best of our knowledge – in the whole tribe. In SEM visualisation, we did not find on the scales any structures resembling pheromone transfer particles (PTPs, see Kristensen and Simonsen 2003). Nevertheless, the mesh-like microstructure of the scales is almost identical to that of scent scales of Rapala dioetas (Hewitson, 1863), Lycaenidae (Lou et al. 2021). Also, the arrangement of the wing patches in the “hindwing upperside–forewing underside” system, as well as the general shape of the scales and size difference between the scent and “regular” background scales, are known in other lepidopterans, especially Papilionoidea – e.g. Nymphalidae (Costanzo and Monteiro 2007), including Heliconiini (Darragh et al. 2017), Pieridae (Nobre et al. 2021), Hesperiidae (Pan et al. 2022) and Lycaenidae (Robbins and Busby 2015). In Arctiinae, male scent structures seem to be associated mostly with the abdomen and genitalia (Conner and Iyengar 2016), but wing structures are also known, e.g. in the Neotropical genus Parascepsis Dognin, 1923 (Grados et al. 2020). In that genus, however, only the forewing underside is provided with a scent patch, whilst the hindwing in the corresponding area has a field of spicules, suggesting involvement of some mechanical interaction in pheromone release. The male pheromones in moths have various roles in the mating behaviour (see Delle-Vedove et al. 2014), and also the fore- and hindwing androconia may have different functions (Costanzo and Monteiro 2007), which requires further research in Tritonaclia.

Generally, the evolution of male secondary sexual organs, like scent patches, is hypothesised to support or even increase species diversification rate (Costanzo and Monteiro 2007; Robbins and Busby 2015; Pereira Martins et al. 2019). Evolutionary gain of such structures may be supported in sympatric speciation, or when species become secondarily sympatric (Robbins et al. 2012), as already observed in Lycaenidae (Pereira Martins et al. 2019). This can also be the case with T. tollini and T. vonifotsy Wiorek, sp. nov., considering their nearly perfectly sympatric ranges, the origins of which still demand examination. Further, in mimetic and sympatric Heliconius Kluk, 1780 species, male pheromones were found to play the key role in maintaining the reproductive barrier (González-Rojas et al. 2020). Considering that T. tollini and T. vonifotsy Wiorek, sp. nov. are also morphologically very similar, including superficially indistinguishable females, the male pheromones could have similar functions in this pair of species.

5. Conclusions

Our results contribute to a better understanding of the biodiversity of the unique lineage of Syntomini tiger moths endemic to Madagascar. The genus Tritonaclia is confirmed to be monophyletic, and its systematics is reviewed based on morphological and molecular data, which are mutually congruent. We describe two new species and place one in synonymy, so that the genus now comprises six species. Several of its species are montane, and the diversity of Syntomini across the main mountain massifs in Madagascar is discussed, underlining the need for further studies of the role of mountainous areas in the diversification of the group. We also report wing scent scales in two Melanonaclia species – structures found for the first time in the entire tribe Syntomini. Further research on the function of these scales is needed, as an intriguing aspect of the chemical ecology of Madagascan Syntomini, whose chemistry remains completely uncharacterised.

6. Acknowledgements

We are grateful to MICET (Antananarivo, Madagascar) staff – especially Tiana Vololona and Benjamin Andriamihaja for arranging research permits, drivers for logistic support, students, guides and porters for inevitable help with fieldwork. We thank Steven M. Goodman, Ravo Eddy Nirina Rakotonandrasana (PN de Isalo) and Sambatrarijaona Andrianambinina (PN d’Andringitra) for help in geolocalising some collecting localities, Hendrik Müller and Hans-Joachim Händel (MLUH-ZNS), Alberto Zilli (Rome), Geoff Martin, Alessandro Giusti (NHMUK), Rodolphe Rougerie, Joël Minet (MNHN) for support during our visits to collections, and Beata Babicz (ISEA PAS Library) for providing hard to find literature, and Irena Brunarska (Jagiellonian University), who operated the SEM microscope. MW is grateful to Michał Grzyska and Marie Fontaneau for help with logistics during his visits to Paris and linguistic support with French texts, Lovasoa Randriantseheno (Institut Pasteur de Madagascar) for consultation of Malagasy place names, and Pavel Matos-Maravi (BC CAS) for discussion of molecular analyses. ŁP thanks Nicole Rasoamanana for hosting during the 2023 visit to PZBT.

The research was funded by the National Science Centre, Poland, Grant No. 2018/29/B/NZ8/00186. MW acknowledges funding from French Government Scholarship (SSHN), and SYNTHESYS+ Research Infrastructure Action under the H2020–EU.1.4.1.2, project number: 823827.

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Supplementary material

Supplementary material 1 

Tables S1–S3

Wiorek M, Lees DC, Wahlberg N, Przybyłowicz Ł (2026)

Data type: .zip

Explanation notes: Table S1. Label data of the specimens included in the study. — Table S2. GenBank accession numbers of sequences used in the study. — Table S3. Collecting localities of specimens included in the study.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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