Genus

Cuphodes Meyrick, 1897. Proc. Linnean Soc. New South Wales 22: 299 (key), 314. Type species: Cuphodes thysanota Meyrick, 1897, by monotypy.

Cyphosticha Meyrick, 1907. Proc. Linnean Soc. New South Wales 32: 49 (key), 61. Type species: Gracilaria (sic) pyrochroma Turner, 1894, by original designation.

Taxonomic history.

The genus was described by Meyrick in 1897 on the basis of a female, C. thysanota from Australia. It was compared with Stathmopoda Herrich-Schäffer, 1853, then included in the family Elachistidae, now included in the family Stathmopodidae (De Prins and De Prins 2011–2025). Meyrick probably confused Stathmopoda for the long and bristly scales on the mid and hindlegs and a similar peculiar posture of the adult. Subsequently Vári (1961) redescribed the genus, arguing that it should be included in the family Gracillariidae, near Cyphosticha Meyrick, 1907. For the description of the male characters, he used a species from India, Phrixosceles trochosticha Meyrick, 1908, the type genus of Phrixosceles Meyrick, 1908 which he considered as a synonym of Cuphodes (Fletcher 1929). For females he used both C. thysanota and P. trochosticha. This synonymy was proposed by Vári (1961) based on the similarity of the wing venation and the bristly hairs on the legs. De Prins et al. (2025), studying the Australian fauna, introduced new characters and considered Cyphosticha as a synonym of Cuphodes. They also moved all Australian species of Phrixosceles to the genus Cuphodes, but left the former genus valid with two species, P. trochosticha Meyrick, 1908 (type genus, India) and P. fibulatrix Meyrick, 1922 (Fiji) (De Prins et al. 2025).

Distribution and species richness.

The genus Cuphodes currently comprises 28 species distributed across Australia, Africa and southeastern Asia (De Prins et al. 2025). The nine species with known biology are leaf miners associated with Ebenaceae, Fabaceae, Putranjivaceae and Rhamnaceae. This total excludes Cuphodes paragrapta (Meyrick, 1915) from South America, which upon examination, does not appear to belong to Cuphodes but to an unknown genus. In addition to the described species, six putative species were reported by Kawakita et al. (2010), comprising leaf miners on Ebenaceae and Rhamnaceae from East Asia (Table 1), whose morphology has not yet been described.

DNA.

Cuphodes has frequently been included in molecular phylogenetic studies. Kawakita (2010) identified it as a putative sister taxon to Epicephala, concluding that it is probably one of the closest relatives of that genus. More recently, De Prins et al. (2025) found that mitochondrial data consistently support the monophyly of the Australian Cuphodes species, recovered as sister to Toowoomba De Prins, Sruoga & Zwick, 2025 + (Parectopa Clemens, 1860 + Polysoma Vári, 1961), albeit with low to moderate statistical support. In contrast, Cuphodes diospyrosella (Issiki, 1957) and C. wisteriella Kuroko, 1982, included in the analyses of Kawahara et al. (2016) and Li et al. (2022), were recovered as sister to all other Ornixolinae. In those studies, Parectopa was placed either very distantly (Kawahara et al. 2016) or was recovered as strongly polyphyletic (Li et al. 2022).

Australian Cuphodes.

The characters reported by De Prins et al. (2025) for Australian Cuphodes are a shining-white backround with a subtle yellow ochreous pattern on the forewing and exceptionally pilose hindlegs, equipped with erected sharply pointed spiniform scales. In male genitalia subscaphium + gnathos strongly developed; costa of valva very strongly sclerotised; subapical part of valva with a hanging semi-round flap or other sclerotisations of different sizes and shapes; transtilla complete but in case of absence the function of transtilla is taken by very long basal apodemes of valvae or sclerotised basal bow of vinculum. In female genitalia apophyses anteriores either absent or rather short; papillae anales strongly fused and deeply nested in the cuticle of segment VIII; antrum / colliculum sclerotised; signa small; bulla seminalis usually situated at anterior margin of segment VII or in segment VII, small with convoluted ductus seminalis.

South African Cuphodes.

Examination of African species treated by Vári (1961), specifically those with known males (C. melanostola (Meyrick, 1918), C. diospyri Vari, 1961, and C. dolichocera Vari, 1961), reveals important differences in the male genitalia compared with the characters reported by De Prins et al. (2025), particularly in the shape of the valva, which is simply rounded apically and lacks notable sclerotization. In the newly described C. spermotrophus sp. nov., two bristles are present at the apex of tegumen (Figs 1, 2). This feature is characteristic of several genera in other subfamilies of Gracillariidae, such as Spulerina (Acrocercopinae) and Spanioptila (Ornixolinae), and is particularly significant within Lithocolletinae. Among Australian Cuphodes, apical tegumen bristles are prominent in the pyrochroma clade, which includes C. pyrochroma (Turner, 1894), C. pandoxa (Turner, 1913), C. callimacha (Meyrick, 1920), but are apparently absent in the other groups. Notably, such bristles are not visible in the South African Cuphodes illustrated by Vári (1961) and are not mentioned in his descriptions. The wing venation does not differ from that of the type species C. thysanota Meyrick, 1897, in particular due to the absence of R 1 and R 2 in the forewing (Figs 4, 5).

Adults of the new South African species exhibit a typical resting posture for Cuphodes: the head is held downward, the abdomen raised and supported by the hindlegs, and the fore- and midlegs extended forward and held close together (Fig. 14) (Kawahara et al. 2016).

Based on these morphological and behavioural traits, we provisionally place spermotrophus sp. nov. in Cuphodes, following Vári’s interpretation, until further molecular or morphological data allow a more definitive placement.