Phylogenetic relationships of the geometroid lepidopterans (: Cimeliidae, Epicopeiidae, Sematuridae, Drepanidae, Uraniidae, Geometridae)

Eugene A. Beljaev

Beljaev, E. A. (2009): Phylogenetic relationships of the geometroid lepidopterans (Lepidoptera: Cimeliidae, Epicopeiidae, Sematuridae, Drepanidae, Uraniidae, Geometridae). Pp. 134-136 in: Hausmann, A. (ed.): Proceedings of the fifth Forum Herbulot 2008. Global strategies for plotting geometrid biodiversity in web-based databases (Munich, ZSM, 24-28 June 2008). – Spixiana 32/1: 134-136 Dr. Eugene A. Beljaev, Institute of Biology and Soil Science, Vladivostok, Russia; e-mail: [email protected]

Advancement in the taxonomy of geometrid Warr. are also transferred into the Desmobathrinae at subfamily and tribal levels seems set on a path of as a basally derived (possibly paraphyletic) group comparison between morphological and molecular (genera marginales, g.m.). The tribe Eumeleini and, phylogenetic analysis. This study investigates the tentatively, the genus Celerena Wlk. are included in morphological phylogeny of families and subfamilies the subfamily Geometrinae. The tribe Ametridini is of geometroid lepidopterans (consisting of Geometr- associated with the subfamily . idae, Uraniidae s.l., Drepanidae s.l., Epicopeiidae and The proposed relationships of the families and Sematuridae), with the Cimeliidae as probable sister subfamilies of geometroid lepidopterans coincide family to this group. Main efforts were given on the with or, at least, do not contradict the modern determination of probable ancestral apomorphies of molecular phylogeny of these taxa in most nodes, the families and subfamilies, and on the establish- including the proposed sister relationship of the ment of monophyletic groups. Sematuridae and Epicopeiidae. The most marked A combination of the results of independent differences concern the grouping of the Uraniidae morphological analyses of basal abdominal seg- subfamilies and the position of the ‘Alsophilinae’. ments and male genitalia (including musculature), A detailed substantiation of the phylogeny is given confirms the sister relationship between Geometridae in Beljaev (2008). Selected apomorphies for most and Uraniidae and the more remote relationship debatable nodes are shown in Fig. 1. with Drepanidae. Phylogenetic relationships of The work was supported by the grants of Russian Drepaninae, Thyatirinae and Cyclidiinae remain Foundation for Basic Researches No 07-04-00882-a unresolved. The assumed branching of subfamilies and Far Eastern Branch of Russian Academy of Sci- in Uraniidae (Uraniinae (Auzeinae (Epipleminae + ence No 06-III-A-06-150. Microniinae))) does not support current theories on Apomorphies* their relationships. Epicopeiidae and Sematuridae, 1. Muscles m2 arising from tegumen dorsolaterally and probably, form a sister pair branching earlier than remotely from median ridge of tegumen. the radiation of tympanote geometroid families. The 2. Labides joined by their ventral margin with dorso- family Cimeliidae is proposed as phylogenetically medial margin of the base of valvellae. closest to the complex of geometroid families. 3. Labides with ventromedial process articulated with A new phylogeny of geometrid subfamilies is lateral margins of the juxta dorsomediad of base of hypothesized, based on the examination of structures valvellae. of the proximal segments of the abdomen, tibial 4. Valvellae separated dorsally from lateral margins of androconial apparatus of males and genitalia. The the juxta. basic division of geometrid subfamilies into two 5. Muscles m3 attached basally to the sclerotized fold large monophyletic lineages – the geometrine and between vinculum and sacculi. 6. Valvellae placed almost ventrad of juxta and flattened larentiine lineages - is confirmed. In the geometrine dorsoventrally. lineage, the branching of subfamilies in the sequence 7. Aedeagus basally ankylosed with caulis. (Archiearinae (Ennominae (Desmobathrinae (Oeno- 8. Anterolateral processes of second sternite modified chrominae s. str. + Geometrinae)))) is postulated. into tympanal organs. The larentiine lineage includes the sister subfamilies 9. Labides arising from basal part of the ventral margin Larentiinae and Sterrhinae. The taxa ‘Alsophilinae’, of costula. ‘Orthostixinae’ and ‘Cheimoptenini’ are included into the core group of Desmobathrinae (genera * For Sematuridae and Epicopeiidae, apomorphies are centrales, g.c.). The genera Abraxaphantes Warr., Di- defi ned relative to the basally derived position of chromodes Gn., Epidesmia D. & Westw. and Heteralex Deuveia in the latter family (Minet, 2002).

134 Drepanidae Uraniidae Geometridae

Cimeliidae Sematuridae Epicopeiidae (Deuveia) Epicopeiidae Thyatirinae Cyclidiinae Drepaniinae Uraniinae Auzeinae Epipleminae Microniinae Archiearinae Ennominae Desmobathrinae (g.c.) Desmobathrinae (g.m.) str. Oenochrominae s. Geometrinae (Celerena) Geometrinae Larentinnae Sterrhinae () Sterrhinae

25- 27 31-32

24 28-30

22-23 37-38 39-40

14-15 20-21

13 16-19 33-36

10-12

5-7 8-9

3-4

1-2

Fig. 1. Phylogenetic cladogram of the families and subfamilies of geometroid lepidopterans.

10. Tympanal case joined with anterior margin of the 19. Signum on the corpus bursae as entire, well-sclero- second abdominal sternite; wall of anterior apodeme tized, rounded, dentate and moderately concave of the sternite lying opposite tympanum is formed plate. by wall of tympanal case. 20. In males setal patches on third abdominal sternite 11. Labides as lobe-like process placed between the consist of strong needle-like setae. dorsobasal angle of valva and juxta, dorsally sepa- 21. Apical part of the air canal of ansa possesses rather rated from subcostal fold of valvula and have flex- long thinned portion, with almost parallel sided, ible connection with costula. apex of canal capitate. 12. Valvellae reduced to small setaceous plates (cristae), 22. Uncus medainly free of connection with tegumen widely fused with juxta and sacculi. and articulated with bases of the gnathos arms 13. Dorsal wall of tympanal case fused with sternite; air only. channel into the cavity of anterior apodeme of the 23. Valva with large, prominent medial lobe of valvula. sternite located between basal plate of the case and 24. Tympanal organs without lobe-like cardolacina. wall of corpus of the case. 25. In males third abdominal sternite with pair of lon- 14. Labides shaped as membranous setaceous lobes. gitudinally elongated setal patches.

15. Saccus absent. 26. Tegumen wide, muscles m4 attached to tegumen. 16. Male third abdominal sternite with a pair of lateral 27. Costa separated from valva at apex. patches of long setae protruding over scales covering 28. Uncus at base with wide median “window” inter- abdomen. sected by longitudinal band of sclerotization. 17. Base of the gnathos arms deeply wedged between 29. Juxta is short and wide, concave at the base, more or uncus and tegumen so that lateral sides of the uncus less semilunar.

base widely contact the base of gnathos. 30. Muscles m3 basally attached to the sacculi. 18. Costula is in form of bilobate hemitranstilla; muscle 31. Scales with large content of green pigment geover-

m4 attached to the dorsal lobe of hemitranstilla, din.

mediad of muscle m2 and both muscles crossed. 32. Male hind tibiae with distal process.

135 33. Male hind tibia with hairpencil, directed towards 40. Discal spots on the wings have light nucleus, differ- medial secretory zone on the second abdominal ent in colour from groundcolour of wings. sternite. 34. Scoloparial dilatation of ansa is formed by dilation Beljaev E. A. 2008. Phylogenetic relationships of the of the ansa air canal. family and subfamilies of geometrid moths (Lepi- 35. Gnathos basally fused with tegumen and uncus. doptera: Geometridae). Chteniya pamyati pamyati 36. Subanal plate fused with median part of gnathos. N. A. Kholodkovskogo [Lectures in Memorian of 37. In males, tibial hairpencil composed of thickened N. A. Kholodkovsky] 60. 238 pp. – Zool. Inst. Russ. springy hairs, medial groove on hind tibia and mar- Acad. Sci. St.-Petersburg. (In Russian with English ginosternal processes on second abdominal sternite summary). absent. Minet J. 2002. The Epicopeiidae: phylogeny and a 38. Labides separated distally from costula by narrow redefinition, with the description of new taxa membranous fold. (Lepidoptera: Drepanoidea) – Annls. Soc. ent. Fr.

39. On forewing, vein M1 relatively very long and section (N.S.) 38 (4): 463-487. between point of branching from Rs stem and discal vein is longitudinally orientated.

Evolutionary Relationships within the Australian Geometrinae – Research Update

Catherine J. Young

Young, C. J. (2009): Evolutionary Relationships within the Australian Geometri- nae – Research Update. Pp. 136 in: Hausmann, A. (ed.). Proceedings of the fifth Forum Herbulot 2008. Global strategies for plotting geometrid biodiversity in web- based databases (Munich, ZSM, 24-28 June 2008). – Spixiana 32/1: 136 Dr. Catherine J. Young, School of Geography and Environmental Studies, Uni- versity of Tasmania, Locked Bag 78, GPO Hobart, 7001, Tasmania; e-mail: [email protected]

This ongoing study examines the relationships and Geometrinae is uncertain, but is evident between Australian geometrine genera and the rela- from Rhodopsin data; tionships between these groups and other geometrid – Dysphanini is a separate tribe from the Geome- subfamilies. Geometrinae are strongly represented trinae on Rhodopsin data only; in Australia and this includes many genera of the – Chlorocoma is not monophyletic as ‘Prasinocyma’ Pseudoterpnini, the so called robust or grey Geo- semicrocea most likely belongs to Chlorocoma, metrinae. Very little has been published on the whereas Chlorocoma cadmaria does not; systematics of this group apart from the important – Anomogenes is possibly an ennomine and not a and comprehensive work by Pitkin et al. (2007) on geometrine. the Pseudoterpnini. Previous research into the higher To further clarify these relationships, molecular data order relationships within geometrid subfamiles has will be analysed using Bayesian inference. Morpho- shown that their may be a sister group relationship logical data, based mainly on adult characters, but between the Geometrinae and the Oenochrominae also some immature characters, will be analysed to s. str. based on both morphological and molecular further understand relationships. data (Young 2006, Yamamoto & Sota 2007). The Pseu- Pitkin, L. M., Han, H. X. & James, S. 2007. Moths of the doterpnini are also interesting in an evolutionary tribe Pseudoterpnini (Geometridae: Geometrinae): context because they have been postulated as being a review of the genera. Zool. Journal Linn. Soc. basally derived. In this study, data from two genes, 150: 343-412. 28SD2 and LW Rhodopsin were used to construct Yamamoto, S. & Sota, T. 2007. Phylogeny of the Ge- phylogenies for geometrine species and representa- ometridae and the evolution of winter moths tives of other geometrid subfamiles. Preliminary inferred from a simultaneous analysis of mitochon- results show that: drial and nuclear genes. Molecular Phylogenetics – The Pseudoterpnini most likely do not form a and Evolution 44: 711-723. separate clade within the Geometrinae; Young, C. J. 2006. Molecular relationships of the Aus- – The Pseudoterpnini are most likely not basally tralian Ennominae (Lepidoptera: Geometridae) and implications for the phylogeny of the Geo- derived; metridae from molecular and morphological data. – A sister relationship between Oenochrominae Zootaxa 1264: 1-147.

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