Manulea Wallengren, 1863

Macià, Ramon, Ylla, Josep, Gastón, Javier, Huertas, Manuel & Bau, Josep, 2022, The species of Eilema Hübner, [1819] sensu lato present in Europe and North Africa (Lepidoptera: Erebidae: Arctiinae: Lithosiini), Zootaxa 5191 (1), pp. 1-87 : 16-17

Genus Manulea Wallengren, 1863

Manulea Wallengren, 1863 , Wiener entomologische Monatschrift, 7: 145, 146.

Type species: Lithosia gilveola Ochsenheimer, 1810 (a junior synonym of Phalaena palliatella Scopoli, 1763 ), by subsequent designation of Moore (1878).

Diagnosis.

Imago. Forewings narrow with nearly straight costal margin and a pale line along it; externally resembles many other groups of Eilema sensu lato. In some species, androconial scales are present on the costal margin of the forewing underside.

Male genitalia. Uncus cylindrical, moderate in width, with a small hooked end; valvae oval or contracted apically, sacculus process ending with and inwardly curved hook, not fused with the valve ending (cucullus); clasper absent; juxta with poorly developed lateral apical processes; vinculum trapezoidal, membranous with a horizontal base, with sharply sclerotised edges; aedeagus thick, globular and large with an apical sclerotised fascia in the form of a spine turned backwards and two cornuti, spike like, of different size.

Female genitalia. In the female genitalia, the main characteristic is the large appendix bursae in comparison to the corpus bursae.

Remarks. Manulea was treated as different from Eilema by Edwards (1996) but was based on Australian species that are significantly different from the type species of both genera.

The genus was characterized and compared with Eilema in detail by Dubatolov & Zolotuhin (2011). No autapomorphies can be recognized, but in the male genitalia, the combination of the following features is characteristic for the genus: (1) the vinculum is trapezoidal and membranous, with thin and weakly sclerotised arms. (2) the valvae has a long and wide dorsal part with thin costa lacking processes; (3) the sacculus is relatively narrow and has an elongate saccular process, well separated from the dorsal part of the valve (cucullus), (4) the aedeagus vesica broad and bearing few robust cornuti. A similar structure of the vinculum is also known in several other genera, namely the Eurasian Eilema , Tarika , Collita , and the African Lepidilema , Dimorphilema , Syntemnilema and Lophilema , but all these genera differ from each other by their valvae structures.

Molecular data. Within the Eilema / Manulea group, the genus Manulea constitutes a clade with moderate Bayesian support (PP=0.61), with Manulea complana and Manulea pseudocomplana forming an internal wellsupported clade (PP=0.93). Genetic TN distances between Manulea species are, in all cases, below 5% (1.21– 4.99%) while showing higher distances with Eilema caniola (5.43–5.98%). RESL Cluster Sequences Analysis (Ratnasingham & Herbert, 2013) returned 5 clusters corresponding to the four existing Manulea species plus an additional cluster including the samples belonging to Manulea iberica . Most of the COI sequences were assigned with high confidence (>98% match) to a pre-existing BIN for the corresponding species. Only the clusters coinciding with the samples of Manulea costalis (Zeller, 1847) and Manulea iberica ( Mentzer, 1980) , with no previously published records, did not match any existing BINs. It is also worth to mention that Maximum Likelihood tests performed independently for each genetic marker (data not shown) returned highly compatible tree topologies between markers and compared with the Bayesian Inference tree.

The analysis of published sequences attributed to Manulea bicolor (Grote, 1864) , Manulea dorsalis (Walker, 1866) , Manulea replana (Lewin, 1805) , Manulea ussurica (Daniel, 1954) , Manulea japonica (Leech,1889) and Manulea sp. shows that they are all more closely related to other genera ( Setema , Katha or others) than to Manulea , with no specimens falling within the Manulea clade ( Fig. 120 ). This can be interpreted as a clear example of the current lack of consistency of this genus in the sense that the type species is not representative of all the species that have been and continue to be placed in it. There is no doubt that the assemblage of species referred to Manulea needs a major revision.

Taxa included

Manulea complana ( Linnaeus, 1758)

Manulea iberica ( Mentzer, 1980)

Manulea pseudocomplana ( Daniel, 1939)

Manulea palliatella ( Scopoli, 1763)

Manulea costalis (Zeller, 1847)

Figures

FIGURE 120. Phylogenetic tree representation obtained by Bayesian Inference of the extended dataset including the COI Barcode sequences published in BOLD as Eilema or Manulea genus and the sequences obtained in this study. The combined data from the mitochondrial marker COI and the nuclear markers ArgK and DDX23 have been run in three different partitions in BEAST2. Posterior probability values are displayed beside the nodes. For clarity, the branches under the species or subspecies nodes have been collapsed indicating the number of specimens for each taxon next to the countries of collection (AU: Austria; CRO: Croatia; CYP: Cyprus; CZR: Czech Republic; DE: Denmark; FI: Finland; Fr: France; GE: Germany; GR: Greece; IT: Italy; MAC: Macedonia; NL: Netherlands; NOR: Norway; PAK: Pakistan; RU: Russia; SP: Spain; UK: United Kingdom). Branches in grey color and font correspond to clades not reviewed in the current work. In all other cases, font colour indicates distinct clades as discussed in the text: Green: Eilema genus; Light blue: Manulea genus; Dark blue: lutarella group from Indalia genus; Yellow: uniola group from Indalia genus; Orange: predotae group from Indalia genus; Red: Nyea genus; Violet: Setema genus; Black: remaining species under study and outgroup species (Nudaria mundana).