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Subfamily Ennominae Duponchel, 1845


The Ennominae comprise the largest of the six subfamilies of Geometridae with 10,682 described species in approximately 1,100 genera. These represent just under half of all geometrids (Minet & Scoble 1999; Scoble & Hausmann 2007). Their distribution is worldwide, but they are particularly well represented in the New World, especially in tropical areas (e.g. Rindge 1983; Krüger & Scoble 1992; Scoble 1995a; Pitkin 2002, 2005). The Nacophorini (sensu Rindge, 1983) are the largest ennomine tribe in southern Australia; they also are widely distributed in the Americas from southern Canada to Chile and southern Argentina. The tribe is poorly represented in Asia and the temperate regions of the Palaearctic (McQuillan 1986). This pattern suggests that fragmentation of Gondwana may have contributed to the dispersal and present day distribution of the tribe.

Duponchel (1845) first proposed the “Ennomites” as a family-group name. However, Pierce (1918) was the first to synthesise a suprageneric classification of Geometridae based on male genitalia. He distinguished eight tribes in the Palaearctic Ennominae: Ennomini, Macariini, Ourapterygini, Bistonini, Boarmiini, Erannini, Gnophini, and Abraxini. Concurrently, Prout (1915–16) grouped genera now known as ennomines as the Geometrinae [Leach], a title subsequently discarded for the subfamily in favour of Ennominae. Prout used the absence or degeneration of M2 in the hindwing as the defining apomorphy for the group, a character still used as the single distinguishing feature for the subfamily, which throws some doubt on the tribe's status as a natural grouping (Minet & Scoble, 1999). Forbes (1948) was the first to attempt a tribal classification of Nearctic ennomine genera and separated tribes on the basis of male antennal, genitalic, and pupal characteristics. McGuffin (1972, 1977, 1981, 1987), in his synopses of Nearctic ennomines, largely followed the tribal classification of Forbes. Holloway (1994) subsequently rationalised the tribal classification and provided a comprehensive listing of tribal names encompassing all family-group names used by various authors up to that time.

Several recent major revisions of ennomine tribes have contributed substantially to the understanding of the taxonomy of the group. Rindge comprehensively reviewed American Bistonini, Melanophiini (both included in the Boarmiini by Holloway (1994)), Nacophorini, and Lithinini (Rindge 1975, 1983, 1986); Scoble (1995b) reviewed Palyadini (treated as Caberini/Baptini by Pitkin (2002)); and Krüger (2001) and Scoble and Krüger (2002) reviewed the genera of Macariini. The Neotropical ennomines, encompassing 267 genera, were reviewed extensively and tribal classifications relevant to this group reassessed by Pitkin (2002, 2005). Beljaev (2008) redefined and proposed new apormorphies based on genital morphology for the Ennomini.

Despite extensive generic revisions, few recent authors have attempted to propose phylogenetic relationships within the Geometridae at the subfamily or tribal levels. McGuffin (1987) deduced phylogenetic relationships within the Ennominae from inferences on the evolutionary status of various characters of adults and immature stages and also drew on observations by Rindge (1964, 1966, 1967, 1973, 1975) on geometrid characters. He proposed that Macariini and Abraxini should hold a ‘basal’ position, and Ourapterygini as the most derived tribe in the Nearctic fauna, using the Oenochrominae as an outgroup to Ennominae. McGuffin also drew attention to the likely importance of using characters from first instar larvae, pupae, and female genitalia in ascertaining ancient relationships. McGuffin was at odds with Forbes in proposing that the bifid pupal cremaster was the ancestral condition and that a cremaster with a full complement of setae was more derived. This supposition was based on the presence of a bifid cremaster in the Palaearctic Archeiarinae (Archiearis Hübner and Leucobrephos Grote), the subfamily generally considered as possessing the most primitive characters based on tympanal and larval features (Minet & Scoble 1999). Holloway (1993) agreed with Forbes’s original proposition that the bifid cremaster is the more derived state on the basis that the presence of four pairs of cremastral setae is more generally distributed in the family. The bifid cremaster is relatively more common in ennomine genera, but it also is found in the Archiearinae, Scopulini, Oenochrominae s. str., Alsophilini, and a few larentiine tribes. However, pupal characteristics by no means have been surveyed extensively, and the importance of these characters is yet to be substantiated.

Holloway endeavoured to relate tribes on the basis of apomorphic adult and larval states and assigned Bornean genera to established tribes on this basis. Later, Holloway (1997) postulated a phylogeny for the family, based mainly on Oriental groups but supplemented with some representatives from other geographical areas. He proposed that characters of the Archiearinae are ancestral within the family, and Alsophilinae and Oenochrominae are the next most “primitive” subfamilies, forming a sister group with the rest of the major subfamilies. Here, the Nacophorini are nested within Ennominae as a sister group to Azelinini and Odontoperini on the basis of larval characters. Interestingly, Beljaev (2006) postulates that Nacophorini fall within the Odontoperini based on characters of the male genitalia.

More recently molecular information has offered new interpretations of relationships within the Geometridae (Abraham et al. 2001, Abraham 2002, Young 2006a, Yamamoto and Sota 2007 and Sihvonenen et al. 2011). Abraham et al. were unable to establish monophyly of the Ennominae from their analysis. Young (2006a) reiterated this result using an expanded group of mainly Australian taxa using 28S D2 and EF 1-α. Yamamoto and Sota (2007), provided a more comprehensive molecular study using five gene fragments from both the mitochondrial and nuclear genomes and 69 ingroup species representing the seven known geometrid subfamilies. However only Japanese species were represented in the analysis. This study also concluded that the Ennominae are polyphyletic and included the Alsophilinae in the sub-family, as determined by both Abraham et al. and Young. The morphological characters that support placement of Alsophila in the Boarmiini were discussed more fully in Young (2006a). Sihvonen et al (2011) conducted the most comprehensive molecular tphylogenetic study to date using eight genes and 150 taxa representing the global fauna. They found that their data. supported most higher order relationships in the family. As before, the Ennominae were found to be relatively recently derived, but, unlike previous studies, the monophyly of the tribe was well-supported. Again, Alsophiliniae fell inside the Ennominae.

The Australian Ennominae were revised by Meyrick (1892) and by Turner (1917, 1919) and many new species were subsequently described by Turner in numerous papers. Both adult and immature stages, in particular larvae, of many southern Australian ennomine species were comprehensively described by McFarland (1988).There has never been a world review of this subfamily; however Young (2006) provided a phylogeny of the Australian Ennominae in context with the global fauna using both molecular and morphological data (Young 2006a, 2008). Young (2006b) also described the eggs of 85 Australian ennomines but, as found in other phylogenetic research, was unable to determine unifying ovoid characters for the group. She also described 67 Australian Ennominae and related taxa using 43 larval characters partly to contextualise the Australian taxa with the world fauna (Young 2008b).

The genus Cleora Curtis was reviewed by D.S. Fletcher (1953, 1967), and in a series on the Australian Nacophorini Thalaina Walker was revised by McQuillan (1981), Mnesampela Guest by McQuillan (1985), Paralaea Guest by McQuillan et al. (2001), Palleopa by Young & McQuillan (2001) and Archephanes by Young & McQuillan (2003). A broad survey, including many of the genera found in northern Australia, was published on the Borneo fauna by Holloway (1993). The tribal classification adopted here follows Forum Herbulot (2003). Beljaev (2009) has susbsequently proposed an alternative classification of the tribes of the Ennominae based primarily on male genitalia.


General References

Abraham, D., Ryrholm, N., Wittzell, H., Hollway, J.D., Scoble, M.J. & Lofstedt, C. 2001. Molecular Phylogeny of the Subfamilies in Geometridae (Geometroidea: Lepidoptera). Molecular Phylogenetics and Evolution 20: 65–77

Beljaev, E. 2008. A new concept of the generic composition of the geometrid moth tribe Ennomini (Lepidoptera, Geometridae) based on functional morphology of the male genitalia. Entomological Review (English translation of Entomologicheskoe Obozrenie) 88(1): 50-60

Beljaev, E. 2009. Tentative tribal system of Ennominae based on current family-group names. Forum Herbulot. (Accesssed 30 May 2012)

Beljaev, E.A. 2006. A morphological approach to the Ennominae phylogeny (Lepidoptera: Geometridae). Spixiana (Munich) 29: 215–216

Duponchel, P.A.J. 1845. Catalogue méthodique des Lépidoptères d'Europe. Paris.

Fletcher, D.S. 1953. A revision of the genus Carecomotis (Lep. Geometridae). Annals and Magazine of Natural History 12 6: 100-142, pls 3-6

Fletcher, D.S. 1967. A revision of the Ethiopian species and a check list of the world species of Cleora (Lepidoptera: Geometridae). Bulletin of the British Museum (Natural History) Entomology Suppl. 8: 1-119, pls 1-14

Forbes, W.T.M. 1948. Lepidoptera of New York and neighbouring states Part II. Geometridae, Notontidae, Sphingidae, Lymantriidae. Memoirs of the Cornell University Agricultural Experiment Station No. 274

Hausmann, A. (ed.) 2003. The Forum Herbulot world list of family group names in Geometridae , 11 pp., with pdf of update version of 12.6.2007. (Acessed 30 May 2012)

Holloway, J.D. 1993. The Moths of Borneo. Family Geometridae, Subfamily Ennominae. Kuala Lumpur : Malayan Nature Society Part 11 309 pp., 593 figs, 19 pls.

Holloway, J.D. 1997. The moths of Borneo Part 10: Family Geometridae, subfamilies Sterrhinae and Larentiinae. Malayan Nature Journal 51: 1-242

Inoue, H. 1992. Geometridae. pp. 111-129 in Heppner, J.B. & Inoue, H. (eds). Lepidoptera of Taiwan. Checklist. Gainsville : Association for Tropical Lepidoptera Vol. 1(2) xlix 276 pp.

Krüger, M. 2001. A revision of the tribe Macariini (Lepidoptera: Geometridae: Ennominae) of Africa, Madagascar and Arabia. Bulletin of the Natural History Museum, London (Zoology) 70: 1–502

Krüger, M. & Scoble, M.J. 1992. Neotropical red-brown Ennominae in the genera Thysanopyga Herrich-Schäffer and Perissopteryx Warren (Lepidoptera: Geometridae). Bulletin of the British Museum (Natural History) Entomology 61: 77–148

McFarland, N. 1988. Portraits of South Australian geometrid moths. Kansas : Allen Press, Lawrence iv+400 pp., frontispiece.

McGuffin, W.C. 1972. Guide to the Geometridae of Canada (Lepidoptera) II. Subfamily Ennominae. 1. Memoirs of the Entomological Society of Canada 86: 1–159

McGuffin, W.C. 1977. Guide to the Geometridae of Canada (Lepidoptera) II. Subfamily Ennominae. 2. Memoirs of the Entomological Society of Canada 101: 1–191

McGuffin, W.C. 1981. Guide to the Geometridae of Canada (Lepidoptera) II. Subfamily Ennominae. 3. Memoirs of the Entomological Society of Canada 117: 1–153

McGuffin, W.C. 1987. Guide to the Geometridae of Canada (Lepidoptera) II. Subfamily Ennominae. 4. Memoirs of the Entomological Society of Canada 138: 1–182

McQuillan, P.B. 1981. A review of the Australian moth genus Thalaina (Lepidoptera: Geometridae: Ennominae). Transactions of the Royal Society of South Australia 105: 1-23

McQuillan, P.B. 1985. A taxonomic revision of the Australian autumn gum moth genus Mnesampela Guest (Lepidoptera: Geometridae, Ennominae). Entomologica Scandinavica 16: 175-202

McQuillan, P.B. 1986. Trans-Tasman relationships in the highland moth (Lepidoptera) fauna. pp. 263-276 in Barlow, B.A. (ed.). Flora and Fauna of Alpine Australasia. Ages and Origins. East Melbourne : CSIRO vii+543 pp.

McQuillan, P.B., Young, C.J. & Richardson, A.M.M. 2001. A revision of the Australian moth genus Paralaea Guest (Lepidoptera: Geometridae: Ennominae). Invertebrate Taxonomy 15: 277–317

Meyrick, E. 1892. Revision of Australian Lepidoptera. V. Proceedings of the Linnean Society of New South Wales 2-n.s. 6(4): 581-678

Minet, J. & Scoble, M.J. 1998. The drepanoid/geometroid assemblage. pp. 301-320 in Kristensen, N.P. (ed.). Lepidoptera, Moths and Butterflies. Volume 1: Evolution, Systematics, and Biogeography. Handbuch der Zoologie/Handbook of Zoology. Volume IV Arthropoda: Insecta Part 35. i-x. Berlin : Walter de Gruyter Vol. 1(35). [year of publication not confirmed]

Pierce, F.M. 1918. Genitalia of the British Geometridae. Liverpool : The Northern Publishing Company, Ltd.

Pitkin, L.M. 2002. Neotropical ennomine moths: a review of the genera (Lepidoptera: Geometridae). Zoological Journal of the Linnean Society 135: 121–401

Pitkin, L.M. 2005. Moths of the Neotropical genera Ischnopteris, Stegotheca and Rucana (Lepidoptera: Geometridae: Ennominae). Systematics and Biodiversity 3(1): 13–96

Prout, L.B. 1915-1916. Geometrinae. pp. 303-413 in Seitz, A. The Macrolepidoptera of the World, The Palaearctic Geometrae. Stuttgart : Alfred Kernen Vol. 4.

Rindge, F.H. 1964. A revision of the genera Melanolophia, Pherotesia and Melanotesia (Lepidoptera). Bulletin of the American Museum of Natural History 126(3)

Rindge, F.H. 1967. A revision of the neotropical species of the moth genus Glena (Lepidoptera, Geometridae). Bulletin of the American Museum of Natural History 127(3)

Rindge, F.H. 1973. A revision of the North American species of the genus Pseudoboarmia (Lepidoptera, Geometridae). American Museum Novitates 2514: 1–27

Rindge, F.H. 1975. A revision of the New World Bistonini (Lepidoptera Geometridae). Bulletin of the American Museum of Natural History 156(2)

Rindge, F.H. 1983. A generic revision of the New World Nacophorini (Lepidoptera: Geometridae). Bulletin of the American Museum of Natural History 175: 147–262

Rindge, F.H. 1986. Generic descriptions of New World Lithinini (Lepidoptera: Geometridae). American Museum Novitates 2838: 1–68

Rindge, F H. 1966. A revision of the of the moth genus Anacamptodes (Lepidoptera, Geometridae). Bulletin of the American Museum of Natural History 132: 175–244

Scoble, M.J. 1995a. The Lepidoptera: Form, Function and Diversity. Oxford : Oxford University Press.

Scoble, M.J. 1995b. A review of the moth tribe Palyadini with the description of a new genus (Geometridae: Ennominae). Systematic Entomology (20): 35–58

Scoble, M.J. & Krüger, M. 2002. A review of the genera of Macariini with a revised classification of the tribe (Geometridae: Ennominae). Zoological Journal of the Linnean Society 134: 257-315

Scoble M.J. & Hausmann A. 2007. Online list of valid and available names of the Geometridae of the World. iBOL. species_checklists.php (Accessed 30 May 2012)

Sihvonen, P., Mutanen, M., Kaila, L., Brehm, G., Hausmann, A. & Staude, H.S. 2011. Comprehensive molecular sampling yields a robust phylogeny for geometrid moths (Lepidoptera: Geometridae). PLoS ONE (Public Library of Science) 6(6): e20356 [1-11]

Turner, A.J. 1917. Revision of the Australian Lepidoptera. VI. Proceedings of the Linnean Society of New South Wales 42: 304-336, 344-390 [3 Oct. 1917]

Turner, A.J. 1919. Revision of Australian Lepidoptera. VI (Third instalment). Proceedings of the Linnean Society of New South Wales 44: 258-310, 383-413 [2 Oct. 1919]

Yamamoto, S. & Sota, T. 2007. Phylogeny of the Geometridae and the evolution of winter moths inferred from a simultaneous analysis of mitochondrial and nuclear genes. Molecular Phylogenetics and Evolution 44: 711–723

Young, C.J. 2006. Molecular relationships of the Australian Ennominae (Lepidoptera: Geometridae) and implications for the phylogeny of the Geometridae from molecular and morphological data. Zootaxa 1264: 1-147

Young, C.J. 2006b. Descriptions of the eggs of some southern Australian Geometridae (Lepidoptera). Zootaxa 1287: 294 pp.

Young, C.J. 2008. Characterisation of the Australian Nacophorini using adult morphology, and phylogeny of the Geometridae based on morphological characters. Zootaxa 1736: 141 pp.

Young, C.J. 2008b. Characterisation of the Larvae of Australian Nacophorini. Zootaxa 1862: 1-74

Young, C.J. & McQuillan, P.B. 2001. Redescription of the little-known Australian geometrid moth Palleopa innotata Walker, 1866 (Lepidoptera: Geometridae: Ennominae). Insect Systematics and Evolution 32: 263–278

Young, C.J. & McQuillan, P.B. 2003. Redescription and life-cycle of Archephanes zalosema Turner, 1926 (Lepidoptera: Geometridae: Ennominae), a specialist on Winteraceae. Insect Systematics and Evolution 34: 81–94


History of changes

Note that this list may be incomplete for dates prior to September 2013.
Published As part of group Action Date Action Type Compiler(s)
21-Feb-2022 GEOMETROIDEA 04-Dec-2021 MODIFIED Dr Cathy Byrne (TMAG) Di Moyle (TMAG)
21-Feb-2022 GEOMETRIDAE 02-Jun-2014 MODIFIED Dr Federica Turco (QM)
21-Feb-2022 28-Jun-2012 MODIFIED
12-Feb-2010 (import)