ZYGAENIDAE AND THEIR INTERNAL

Yen, Shen-Horn, Robinson, Gaden S. & Quicke, Donald L. J., 2005, The phylogenetic relationships of Chalcosiinae (Lepidoptera, Zygaenoidea, Zygaenidae), Zoological Journal of the Linnean Society 143 (2), pp. 161-341 : 272

MONOPHYLY OF ZYGAENIDAE AND THEIR INTERNAL RELATIONSHIPS

Compared with earlier studies using incomplete taxon sampling and far fewer characters, the phylogenetic trees obtained during the present study are largely incongruent with previous hypotheses of zygaenid relationships. The new arrangements differ in the following aspects.

The Procridinae , either as a monophyletic group (EW) or as a paraphyletic one (SAW), are the most basal lineage of Zygaenidae . Inclusion of Janseola was first documented by Vári et al. (2002) who referred to an unpublished comment by Alberto Zilli. Although a clade that comprises Janseola and Inouela + Homophylotis nigra is recovered in the present study, the most closely related genus to Janseola is probably Tascia Walker, 1856 [type species: T. chrysotelus Walker, 1856 , now called T. finalis ( Walker, 1854) ] from southern Africa. This relationship is based on the features of valval shape, articulated stick-like appendages derived from conjunction between valval bases and transtilla. The genus Saliunca , placed in ‘Callizygaenini’ by Alberti (1954), is placed in the Procridinae , and therefore the Callizygaenini sensu Alberti is paraphyletic.

Monophyly of the Zygaeninae is supported in the present study, but it is not suggested to be the sister group of Chalcosiinae because the clade comprising Callizygaeninae + Cleoda is more closely related to the clade which comprises Heteropan and Chalcosiinae .

Monophyly of the Callizygaeninae (presently comprising Callizygaena and Procotes ) + Cleoda is strongly supported. However, the characters of ‘ Cleoda syntomoides’ are in fact based on a speculated ‘marriage’ between the ‘male’ and ‘female’ specimens. This species was described by Walker (as Doclea syntomoides ) based on a male specimen with ‘greenish forewing from Malakka’. However, the ‘ type specimen’ deposited in the Hope Entomological Collection of Oxford University Museum is in fact a female, which exhibits completely different body size and wing patterns (yellow wings, see Fig. 9 ) from the features described by Walker. This ‘ type specimen’ was probably used for the figure of the ‘ Docleopsis sulaensis’ in Swinhoe (1892: t.2, f.10), but it is not conspecific with the true Docleopsis sulaensis Jordan, 1907 from Sula archipelago.

We were unable to locate the ‘male type’ of Syntomoides in Oxford University Museum, although two male specimens were found in the Natural History Museum in London. One of them may have been the model for the illustration of Syntomoides in Seitz’s ‘ Gross-Schmetterlinge der Welt ’ ( Jordan, 1907: 16, t. 2e). The characters and collection locality (‘Malakka’) agree well with the description by Walker so we suspect that they are the ‘real’ Syntomoides . To examine whether the male and the suspected female were conspecific, we conducted a test analysis in which the male and female were treated as different terminal taxa. However, they still formed a monophyletic group, which clustered with Callizygaena ada . Therefore we tentatively associated the data sets of the male Syntomoides and the speculated female into one data set for all the analyses. Cleoda and Callizygaena are grouped with the characters of the multi-forked scales on the chaetosemata and wings. It is still uncertain whether Cleoda can be placed in Callizygaeninae because information on the immature stages is not available. The male androconial organ of Cleoda is homoplasious with those of Harrisina and Pryeria , and the extended female ovipositor is distinct from those of the species placed in Callizygaena and Procotes .

Heteropan and Chalcosiinae always form a sistergroup relationship based on 27 synapomorphies (EW/ SAW). Of these, seven characters are unique to this clade: patagia strongly sclerotized and sac-like (70: 4), patagia more developed than parapatagia (72: 2), T1 of larva with retractile cervical gland (354: 2), thoracic spiracles not concealed by ‘spiracle access’ (393: 1), terga 3–8 with multi-rowed spinulets (394: 3), spinulets occupying the whole terga of the pupa (395: 2), and A10 of pupa with an elevated transverse tuberculate band (396: 1). The three taxa of Heteropan are grouped by 27 apomorphies (EW/SAW), but of these only 56: 2 (lateral margin of flagellomeres ridged and undulated, 57: 2 (back–bone of flagellomere cristate or keeled along the whole antenna) are unique.

Monophyly of Chalcosiinae is supported by 24 (EW) or 23 (SAW) characters. Of these, only two characters, scales on patagia bending downwards (81: 1), subdorsal and lateral syndeses located posterior to front margin of tergum 2 (178: 3), are unique. Tarmann (1992c), Epstein et al. (1999) and Naumann et al. (1999) suspected that the hindwing-abdominal androconial organ, developed ovipositor and absence of accessory glands in female genitalia provided the synapomorphies of Chalcosiinae . However, the present study shows that only two uncontradicted characters relevant to the scent organs – a bundle of long hair bristles inserting into the pleural pouch (337: 3) and presence of ‘normal type’ of pleural pouch (338: 4) – support the monophyly of Chalcosiinae . Presence of elongated ovipositor occurs inconsistently in the majority of the studied taxa, so this trait is not seen as a synapomorphy of the subfamily. In addition, 403: 5 (mandibular lobe long, with sclerotization from sub-basal to apical part), an uncontradicted character relevant to the chemical defence system, also supports the monophyly of Chalcosiinae .

Figures

Figure 9. Ingroup (part) and outgroups. From top to bottom, left to right. Row I: (Ingroup) Chalcosiopsis variata, Heteropan sp. (left), Heteropan appendiculata (right), Inouela sp., Doclea syntomoides, (outgroups from here) Callizygaena ada (Callizygaeninae). Row II: Artona gracilis, Clelea formosana, Theresimima ampellophaga (Procridinae), Dianeura jacksoni (Anomoeotidae). Row III: Himantopterus fuscinervis (Himatopteridae), Janseola titaea (placed in Heterogynidae by Scoble, 1992), Burlacena sp. (Zygaenidae, subfamily unassigned), Saliunca sp. (Procridinae). Row IV: Harrisina sp., Pollanisus viridipulverulenta, Adscita statices, Illiberis pruni (Procridinae), Staphylinochrous sp. (Anomoeotidae). Row V: Pryeria sinica, Zygaena fausta (Zygaeninae), Heterogynis sp. (Heterogynidae), Lactura dives (Lacturidae), Phauda sp. (Phaudinae).