Hesperilla, HEWITSON, 1868

Toussaint, Emmanuel F. A., Braby, Michael F., Müller, Chris J., Petrie, Edward A. & Kawahara, Akito Y., 2022, Molecular phylogeny, systematics and generic classification of the butterfly subfamily Trapezitinae (Lepidoptera: Papilionoidea: Hesperiidae), Zoological Journal of the Linnean Society 195, pp. 1407-1421 : 1414-1416

HESPERILLA HEWITSON, 1868

Motasingha Watson, 1893 syn. nov.

Oreisplanus Waterhouse & Lyell, 1914 syn. nov.

Type species: Hesperilla ornata Leach, 1814 (by original designation).

Remarks

Braby (2010) noted that the genera Hesperilla Hewitson, 1868 , Motasingha Watson, 1893 and Oreisplanus Waterhouse & Lyell, 1914 are closely related and probably form a monophyletic group. Currently, 14 species are assigned to Hesperilla , whereas Motasingha and Oreisplanus each contain two species: Motasingha dirphia (Hewitson, 1868) and Motasingha trimaculata (Tepper, 1882) in Motasingha ( Moulds & Atkins, 1986) , and Oreisplanus munionga (Olliff, 1890) and Oreisplanus perornatus (Kirby, 1893) in Oreisplanus ( Braby, 2000) .

Atkins (1984) and Moulds & Atkins (1986) first demonstrated that Motasingha is most closely related to Hesperilla , indicating that it differed chiefly by the form of the apiculus of the antennal club, which is long, stout and blunt rather than long and pointed. However, like Hesperilla , the club is bent before the middle. Other ‘distinguishing’ morphological characters, such as the wing venation, relative length of the third segment of the labial palpus, number of spurs on the hind tibia, shape of the linear patch of sex-scales (androconia), male and female genitalia, and morphology of the larva (which is semi-translucent) and pupa (which is long and cylindrical, with a strongly produced and heavily sclerotized pupal cap) are otherwise similar to Hesperilla . Indeed, Atkins (1984) noted the strong similarity between Motasingha , which at that time was monotypic, containing the single species Motasingha dirphia , and the Hesperilla idothea (Miskin, 1889) species group of Hesperilla (i.e. Hesperilla donnysa , Hesperilla flavescens , Hesperilla idothea and Hesperilla mastersi ) based on comparative illustrations of the wing venation, male androconia and female genitalia.

Braby (2000) noted that Oreisplanus is closely allied to Hesperilla but is distinguished by the absence of male androconia (forewing sex-brand) and the form of the club of the antenna, which is evenly curved, with the apiculus short and blunt. Absence of a character does not diagnose a lineage and, moreover, androconia are likely to be under strong sexual selection. Other characters used to diagnose Oreisplanus are shared with Hesperilla , such as the wing venation, labial palpus, hind tibia, and form of the pupal cap, which is strongly produced and heavily sclerotized, in addition to the abdominal segments of the pupa, which, like Motasingha , contain long stiff setae that, in addition to the cremaster, provide support within the silken shelter. Indeed, A. F. Atkins (pers. comm.) considers the Hesperilla ornata (Leach, 1814) species group (i.e. Hesperilla ornata , Hesperilla picta , Hesperilla hopsoni and Hesperilla crypsargyra ) to be more closely related to Oreisplanus than to other species of Hesperilla ( Braby, 2010) , a phylogenetic pattern which is concordant with our findings. This pattern was also partly evident in an earlier phylogenetic study by Warren et al. (2008), who showed that Oreisplanus perornatus was sister to Hesperilla ornata and that this pair was sister to Hesperilla donnysa , rendering Hesperilla paraphyletic.

In our phylogenetic analysis, the genus Hesperilla was paraphyletic by inclusion of both Motasingha and Oreisplanus . Motasingha was recovered as sister to a clade that included the Hesperilla idothea species group ( Hesperilla donnysa , Hesperilla flavescens , Hesperilla idothea and Hesperilla mastersi ) and the Hesperilla malindeva species group (i.e. Hesperilla malindeva , Hesperilla sarnia , Hesperilla sexguttata , Hesperilla furva and Hesperilla crypsigramma ) ( Atkins, 1978); this clade, in turn, was sister to a clade that included the remaining species of Hesperilla . Oreisplanus was monophyletic with low branch support, nested within a clade comprising the Hesperilla ornata species group (i.e. Hesperilla ornata , Hesperilla picta , Hesperilla crypsargyra and Hesperilla hopsoni ). This clade formed a monophyletic group sister to Hesperilla chrysotricha .

Given the well-supported phylogenetic relationships among the three genera uncovered in the present study, and the lack of clear synapomorphies by which to distinguish Motasingha and Oreisplanus , we synonymize Motasingha and Oreisplanus under Hesperilla s.l., as foreshadowed by Braby (2010) more than a decade ago. Thus, according to our phylogeny, the nomenclature, new combinations and synonyms recognized for the revised 18 species of Hesperilla s.l. and their constituent subspecies are as follows:

Hesperilla chrysotricha ( Meyrick & Lower, 1902)

Hesperilla chrysotricha chrysotricha ( Meyrick & Lower, 1902)

Hesperilla chrysotricha cyclospila ( Meyrick & Lower, 1902)

= Hesperilla chrysotricha leucospila Waterhouse, 1927

= Hesperilla chrysotricha plebeia Waterhouse, 1927

= Hesperilla chrysotricha leucosia Waterhouse, 1938

= Hesperilla chrysotricha lunawanna L.E. Couchman, 1949

= Hesperilla chrysotricha naua L.E. Couchman, 1949

Hesperilla ornata (Leach, 1814)

Hesperilla ornata ornata (Leach, 1814)

Hesperilla ornata monotherma (Lower, 1907)

Hesperilla picta (Leach, 1814)

Hesperilla perornatus Kirby, 1893

Hesperilla munionga Olliff, 1890

Hesperilla munionga munionga Olliff, 1890

Hesperilla munionga larana (Couchman, 1962) comb. nov.

Hesperilla crypsargyra (Meyrick, 1888)

Hesperilla crypsargyra crypsargyra (Meyrick, 1888)

= Hesperilla crypsargyra lesouefi Tindale, 1953

Hesperilla crypsargyra binna Johnson & Wilson, 2005

Hesperilla hopsoni Waterhouse, 1927

Hesperilla dirphia Hewitson, 1868

Hesperilla trimaculata Tepper, 1882

Hesperilla trimaculata trimaculata Tepper, 1882

= Hesperilla quadrimaculata Tepper, 1882

Hesperilla trimaculata dilata (Waterhouse, 1932) comb. nov.

= Motasingha dirphia dea Waterhouse, 1932

Hesperilla trimaculata occidentalis Moulds & Atkins, 1986 comb. nov.

Hesperilla donnysa Hewitson, 1868

Hesperilla donnysa donnysa Hewitson, 1868

= Hesperilla donnysa diluta Waterhouse, 1932

= Hesperilla donnysa delos Waterhouse, 1941

= Hesperilla donnysa icaria Waterhouse, 1941

= Hesperilla donnysa patmos Waterhouse, 1941

= Hesperilla donnysa samos Waterhouse, 1941

Hesperilla donnysa aurantia Waterhouse, 1927

Hesperilla donnysa galena Waterhouse, 1927

Hesperilla donnysa albina Waterhouse, 1932

Hesperilla flavescens Waterhouse, 1927

= Hesperilla donnysa flavia Waterhouse, 1941

Hesperilla idothea (Miskin, 1889)

Hesperilla idothea idothea (Miskin, 1889)

= Telesto dispar Kirby, 1893

Hesperilla idothea clara Waterhouse, 1932

Hesperilla mastersi Waterhouse, 1900

= Hesperilla mastersi marakupa L.E. Couchman, 1965

Hesperilla malindeva Lower, 1911

= Hesperilla malindeva dagoomba Johnson & Valentine, 1994

Hesperilla sarnia Atkins, 1978

Hesperilla sexguttata Herrich-Schäffer, 1869

= Toxidia sexguttata sela Waterhouse, 1932

Hesperilla furva Sands & Kerr, 1973

Hesperilla crypsigramma ( Meyrick & Lower, 1902)

Larvae of Hesperilla specialize on Cyperaceae , especially the genus Gahnia , with one species (H e s p e r i l l a t r i m a c u l a t a) a l s o f e e d i n g o n Haemodoraceae .