Nacophorini

Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Panorpida
Cladus: Amphiesmenoptera
Ordo: Lepidoptera
Subordo: Glossata
Cladus: Coelolepida
Cladus: Myoglossata
Cladus: Neolepidoptera
Infraordo: Heteroneura
Cladus: Eulepidoptera
Cladus: Ditrysia
Cladus: Apoditrysia
Cladus: Obtectomera
Cladus: Macroheterocera
Superfamilia: Geometroidea

Familia: Geometridae
Subfamilia: Ennominae
Tribus: Nacophorini
Genera: Achagua – Aconcagua – Acrasia – Aethaloida – Amelora – Amphiclasta – Amphicrossa – Androchela – Animomyia – Anischopteris – Aphantes – Aporoctena – Aragua – Arauco – Archephanes – Argidava – Argyrophora – Asemoprepes – Authaemon – Azuayia – Bradyctena – Canelo – Capusa – Cassythaphaga – Ceratonyx – Charca – Chiricahua – Chlenias – Chlenomorpha – Ciampa – Cidariophanes – Conosara – Corula – Cryphaea – Crypsiphila – Cundinamarca – Cycloprorodes – Dispatha – Dolabrossa – Drepanogynis – Drymoptila – Dysbatus – Epicompsa – Eucopelates – Eutelostolmus – Fisera – Gabriola – Gastrina – Gastrinopa – Harpagocnema – Hasodima – Hebdomophruda – Hemimorina – Hemnypia – Hildalgo – Holochroa – Huapianus – Hypsitropha – Illa – Ischnopteris – Kunanyia – Larophylla – Liometopa – Lophosigna – Lophosticha – Lophothalaina – Loweria – Lychnographa – Lyelliana – Mauna – Megazancla – Melanodes – Microligia – Mictodoca – Middletonia – Mnesampela – Mochlotona – Nacophora – Nazca – Neoteristis – Niceteria – Nisista – Nycterephes – Omaguacua – Oratha – Pachycnemoides – Palleopa – Panhyperochia – Papago – Paralaea – Parexcelsa – Paurocoma – Phaeoura – Philolochma – Plesanemma – Plesiolaea – Prosotera – Pseudomaenas – Rhynchopsota – Rindgeopoya – Rucana – Smyriodes – Stegotheca – Stibaroma – Synzeuxis – Tarma – Terpnostola – Thalaina – Thyrinteina – Wichanraran – Xylodryas – Yermoia
References

Byrne, C.J. & N.-S. Wei 2012: Kunanyia stephaniae gen. nov. & sp. nov. (Lepidoptera: Geometridae: Ennominae): an unusual and rare diurnal moth from the mountains of Tasmania. Zootaxa 3503: 25–46. Abstract: [1].
Ferris, C.D. 2010: A new Geometrid genus and species from Southeastern Arizona (Ennominae:Nacophorini). Journal of the Lepidopterists' Society 64 (3): 147–153.
Herbulot, C. 1992f: Nouveaux Ennominae du Cameroun, du Natal et des Comores (Lepidoptera Geometridae). Bulletin de la Société entomologique de Mulhouse 48(1): 37–39. Reference page.
Krüger, M. 1997: Revision of Afrotropical Ennominae of the Drepanogynis Group 1: The genus Hebdomophruda Warren. Part I (Lepidoptera: Geometridae). Annals of the Transvaal Museum 36 (21): 257–295.
Krüger, M. 2005: New species of geometrid moths from Lesotho (Lepidoptera, Geometroidea). Annals of the Transvaal Museum 42: 19–45.
Krüger, M. 2007: Composition and origin of the geometrid fauna (Lepidoptera) of the Sneuberge, Eastern Cape, with descriptions of new taxa. Annals of the Transvaal Museum 44: 25–66.
Krüger, M. 2009: New Afrotropical Nacophorini in the genera Pachycnemoides Krüger, Argyrophora Guenée, Microligia Warren, Pseudomaenas Prout and Drepanogynis Guenée (Lepidoptera: Geometridae: Ennominae). Annals of the Transvaal Museum, 46: 1–35. abstract only seen
Krüger, M. 2013: Revision of Acrasia Rogenhofer, 1875 and description of Panhyperochia gen. nov., a new genus of putative nacophorine geometrids from South Africa (Lepidoptera: Geometridae: Ennominae). Annals of the Ditsong National Museum of Natural History 3: 99–152. [2] Reference page.
Krüger, M. 2014. A revision of the Mauna Walker, 1865 and Illa Warren, 1914 group of genera (Lepidoptera: Geometridae: Ennominae: Nacophorini). Annals of the Ditsong National Museum of Natural History 4: 77–173. Reference page.
Martins Dias, M. 1998: Taxonomical considerations on the genus Cunginamarca Rindge, 1983 (Lepidoptera: Geometridae). Revista Brasileira de Zoologia 15 (4): 951–958. DOI: 10.1590/S0101-81751998000400013. Full article: [3].
Parra, L.E. & M. Pascual-Toca 2003: Revisión taxonómica de los géneros Oratha Walker 1863 y Hasodima Butler 1882 (Lepidoptera: Geometridae). Revista chilena de historia natural 76 (1): 117–128. Abstract and full article: DOI: 10.4067/S0716-078X2003000100011.
Pitkin, L.M. 2005: Moths of the Neotropical genera Ischnopteris, Stegotheca and Rucana (Lepidoptera: Geometridae: Ennominae). Systematics and Biodiversity 3 (1): 13–96. DOI: 10.1017/S1477200004001616.
Rindge, F.H. 1983: A generic revision of the New World Nacophorini (Lepidoptera, Geometridae). Bulletin of the American Museum of Natural History 175 (2): 147–262.
Rindge, F.H. 1961: A revision of the Nacophorini (Lepidoptera, Geometridae). Bulletin of the American Museum of Natural History 123 (2): 91–153. Full article: [4].
Young, C.J. 2008, Characterisation of the Larvae of Australian Nacophorini Zootaxa 1862: 1–74.
Young, C.J. & P.B. Mcquillan 2001: Redescription of the little-known Australian geometrid moth Palleopa innotata Walker, 1866 (Lepidoptera: Geometridae: Ennominae). Insect Systematics & Evolution 32 (3): 263–278. Abstract: DOI: 10.1163/187631201X00209.

The Nacophorini are one of the smaller tribes of geometer moths in the subfamily Ennominae. They are the most diverse Ennominae of Australia and are widespread in the Americas. If the African genera tentatively placed herein indeed belong here, the distribution of the Nacophorini is distinctly Gondwanan, with their probable origin either of Australia, South America or even Antarctica (which was not ice-covered until a few million years ago). In Eurasia, they are rare by comparison.[1]

Despite the lack of thorough study of this tribe in modern times, as traditionally delimited they are probably nearly monophyletic, requiring only a few genera to be moved in and out of this group to make it correspond to a clade;[1] as this involves the type species, the correct name for this clade might be Lithinini or maybe Campaeini, which are both liable to be eventually merged with the Nacophorini. The Azelinini, Ennomini and perhaps the Caberini are probably their closest living relatives, and a more radical approach to monophyly would be to merge the Nacophorini, Lithinini and possibly the Campaeini into the Ennomini.

Description

Nacophorini are generally robust and quite hairy geometer moths, though some species are more delicate. Exceptional among their subfamily, many have slim wings. They typically rest with the hindwings tucked under the forewings. Nacophorini have long antennae, and most if not all have terminal sensillae shaped like stout pegs and sensillae basiconicae on the flagellomeres or rami. The "horn" between the antenna sockets which is present in many geometer moths is usually exceptionally well developed in the Nacophorini. Some have a crest of thorns on their thorax, and a few have a spine at the tip of their foreleg tibia. The hindleg tibia is usually swollen in males, which also often have a "penciltip" of hairs tucked into a groove. Together with a comb of setae on the third abdominal segment, these structures probably serve to distribute pheromones, and while the abdominal comb is found in many Ennominae, the full set of structures is rarely found outside of the Nacophorini, which usually possess at least a swollen tibia or tibial "pencil", and often both.

Wnile the female genitalia are rather nondescript, there are a number of features of the male genitalia that are usually not exclusive to Nacophorini, but in combination are quite characteristic. Like in most Boarmiini, the valval costa typically has a batch of bristles on its underside near the tip, whereas the harpe or "clasper" of Nacophorini lacks the complex modifications found in Boarmiini. The aedeagus has a pointed tip in almost all members of this tribe, displaying little of the variation found in related geometer moths. The anellus usually has extensions at the side, which extend from the edge of the juxta and can be lobes or spines, small or large, covered in bristles or nude. But unlike the similar-looking but probably analogous structures found in Ourapterygini these "furcae" are entirely or almost symmetrical in Nacophorini. The Lithinini and the Tasmanian Ennominae traditionally placed in the Archiearinae also have such symmetrical furcae, indicating the close relationship between them and the Nacophorini.[1]
Ecology

Larval food plants are mainly Rosidae. Caterpillars of the Australian and South American genera feed predominantly on Myrtaceae, including Campomanesia, Eucalyptus, Eugenia and guavas (Psidium). A rather notorious nacophorine species from the tropical Americas is the Brazilian eucalyptus brown looper (Thyrinteina arnobia), which can be a commercially significant pest in eucalyptus plantations. Faboideae and Mimosoideae have also been recorded as food plants.

Caterpillars of the African species tentatively placed in this tribe have been recorded from Cunoniaceae, Ericaceae, Fabaceae and Thymelaceae. Caterpillars of Declana from New Zealand, also tentatively placed in the Nacophorini, have been found on Myrtaceae (eucalyptus, Kunzea and Leptospermum), Pinaceae – larches (Larix), pines (Pinus, notably Monterey pine, P. radiata) and coast Douglas-fir (Pseudotsuga menziesii) –, and southern beeches (Nothofagaceae). In addition, they were found on a species of Olearia, the only euasterid recorded as food plant of Nacophorini to date.[1]
Systematics

The genera Oratha, usually placed in the Nacophorini, is in many aspects suspiciously similar to Pero of the Azelinini and Rhinodia of the Caberini. It is liable to be moved to either of these tribes.

Phaeoura, which includes Nacophora nowadays, appears to be closer to the Ennomini. As this includes the type species Phaeoura quernaria,[2] Lithinini which might warrant merging with the Nacophorini would then supersede the name "Nacophorini", which would become a junior synonym of the Ennomini. The enigmatic genus Hoplosauris, of uncertain placement in the Ennominae, is in some respects intermediate between the Nacophorini and the Ennomini.

While the taxonomic and systematic questions are in need of thorough study, the situation regarding the Lithinini is more clear. Their genus Idiodes and some species formerly placed in the paraphyletic Metrocampa (at least "M." ada and "M." biplaga) appear very close to nacophorine genera – to Thalaina, and to Conosara and Corula, respectively. The Lithinini and the Nacophorini share the same apomorphies of the male genitalia, and their caterpillars are also very similar. It is unknown whether the somewhat more distinct Campaeini would warrant inclusion in this group too.

The Tasmanian genera Acalyphes and Dirce and possibly the South American Archiearides, traditionally placed in the subfamily Archiearinae, seem to be close relatives of the Australian nacophorines Niceteria and Paralaea and would probably need to be moved to the present tribe. And finally, there are some African genera as well as Declana from New Zealand which are tentatively assigned to the Nacophorini.[1]
Selected genera

As numerous ennomine genera have not yet been assigned to a tribe,[3] the genus list should be considered preliminary.
Clara's satin moth, Thalaina clara

Achagua
Aconcagua
Acrasia (tentatively placed here)
Aethaloida
Amelora
Amphiclasta
Amphicrossa
Androchela
Animomyia
Anischnopteris
Aphantes
Aphilopota (tentatively placed here)
Aporoctena
Archephanes
Argidava
Argyrophora (tentatively placed here)
Authaemon
Betulodes
Bradyctena
Callioratis (tentatively placed here)
Canelo
Capusa
Cassythaphaga
Ceratonyx
Charca
Chiricahua[4]
Chlenias
Chlenomorpha
Chrysomima
Ciampa
Cidariophanes
Conosara
Corula
Cryphaea
Crypsiphila
Cycloprorodes
Declana (tentatively placed here)
Dectochilus
Diptychis (tentatively placed here)
Dolabrossa
Drepanogynis (tentatively placed here)
Drymoptila
Dysbatus
Epicompsa
Erilophodes
Fisera
Furcatrox
Gabriola
Gastrina
Hasodima
Hebdomophruda (tentatively placed here)
Hemimorina
Hemnypia
Holochroa
Illa (tentatively placed here)
Ischnopteris
Kunanyia[5]
Lackrana
Mauna (tentatively placed here)
Melanodes
Microligia (tentatively placed here)
Mictodoca
Mnesampela
Mochlotona
Nazca
Niceteria
Nisista
Oratha (probably belongs in Azelinini or Caberini)
Pachycnemoides (tentatively placed here)
Palleopa
Papago
Paralaea
Parexcelsa
Phaeoura (probably belongs in Ennomini, includes Nacophora)
Plesanemma
Pseudomaenas (tentatively placed here)
Rhynchopsota
Rucana
Smyriodes
Stibaroma
Thalaina
Thyrinteina
Veniliodes (tentatively placed here)
Yermoia
Zerenopsis (tentatively placed here)

As noted above, Acalyphes, Dirce and possibly Archiearides would seem to need moving here from the Archiearinae, and at least some Lithinini and perhaps Campaeini seem to belong here too.
Footnotes

Young (2008)
Pitkin & Jenkins (2007)
See references in Savela (2008)
"Nacophorini Report". Integrated Taxonomic Information System. Retrieved 2018-04-27.

Byrne, C. J. and N. Wei. (2012). Kunanyia stephaniae gen. nov. & sp. nov. (Lepidoptera: Geometridae: Ennominae): an unusual and rare diurnal moth from the mountains of Tasmania. Zootaxa 3503 25-46.

References
Wikimedia Commons has media related to Nacophorini.

Pitkin, Brian & Jenkins, Paul (2007): Butterflies and Moths of the World, Generic Names and their Type-species - Nacophora. Retrieved 22 July 2008.
Savela, Markku (2008): Lepidoptera and Some Other Life Forms: Ennominae. Version of 8 March 2008. Retrieved 21 July 2008.
Young, Catherine J. (2008): Characterisation of the Australian Nacophorini using adult morphology, and phylogeny of the Geometridae based on morphological characters. Zootaxa 1736: 1-141. PDF abstract and excerpt