The Nacophorini are one of the smaller

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...

s of

geometer moth The geometer moths are moths belonging to the family Geometridae of the insect order Lepidoptera, the moths and butterflies. Their scientific name derives from the Ancient Greek ''geo'' γεω (derivative form of or "the earth"), and ''metr ...

s in the

subfamily In biological classification, a subfamily (Latin: ', plural ') is an auxiliary (intermediate) taxonomic rank, next below family but more inclusive than genus. Standard nomenclature rules end subfamily botanical names with "-oideae", and zoologi ...

Ennominae Ennominae is the largest subfamily of the geometer moth family (Geometridae) with some 9,700 described species in 1,100 genera. They are usually a fairly small moths, though some (such as the peppered moth) grow to be considerably large. This sub ...

. They are the most diverse Ennominae of

Australia Australia, officially the Commonwealth of Australia, is a Sovereign state, sovereign country comprising the mainland of the Australia (continent), Australian continent, the island of Tasmania, and numerous List of islands of Australia, sma ...

and are widespread in the

Americas The Americas, which are sometimes collectively called America, are a landmass comprising the totality of North and South America. The Americas make up most of the land in Earth's Western Hemisphere and comprise the New World. Along with th ...

. If the

Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

tentatively placed herein indeed belong here, the distribution of the Nacophorini is distinctly

Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final stages ...

n, with their probable origin either of Australia, South America or even

Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest contine ...

(which was not ice-covered until a few

million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (million years ago). ...

). In

Eurasia Eurasia (, ) is the largest continental area on Earth, comprising all of Europe and Asia. Primarily in the Northern and Eastern Hemispheres, it spans from the British Isles and the Iberian Peninsula in the west to the Japanese archipelago a ...

, they are rare by comparison.Young (2008) Despite the lack of thorough study of this tribe in modern times, as traditionally delimited they are probably nearly

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gro ...

, requiring only a few

to be moved in and out of this group to make it correspond to a

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

; as this involves the

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...

, 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 __NOTOC__ The Azelinini are a tribe of geometer moths in the subfamily Ennominae, with many species in the Southern Hemisphere, particularly in South America. Several species are found in North America as well. These stocky geometer moths seem t ...

Ennomini __NOTOC__ The Ennomini are a tribe of geometer moths in the Ennominae subfamily. They are large-bodied and rather nondescript Ennominae, overall showing many similarities to the closely related Azelinini and Nacophorini.Young (2008) Most have ...

and perhaps the

Caberini Caberini is a tribe of geometrid moths in the family Geometridae. There are at least 50 described species in Caberini. Taxonomic note: *Molecular analyses by Sihvonen et al. (2011) supports separating the tribe Baptini Baptini is a tribe of ge ...

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

s, though some

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

are more delicate. Exceptional among their

, 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

flagellomere Antennae ( antenna), sometimes referred to as "feelers", are paired appendages used for sensing in arthropods. Antennae are connected to the first one or two segments of the arthropod head. They vary widely in form but are always made of one o ...

s 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 The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...

, 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

seta In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. Th ...

e on the third

abdominal The abdomen (colloquially called the belly, tummy, midriff, tucky or stomach) is the part of the body between the thorax (chest) and pelvis, in humans and in other vertebrates. The abdomen is the front part of the abdominal segment of the torso ...

segment, these structures probably serve to distribute

pheromone A pheromone () is a secreted or excreted chemical factor that triggers a social response in members of the same species. Pheromones are chemicals capable of acting like hormones outside the body of the secreting individual, to affect the behavio ...

s, and while the abdominal comb is found in many

, 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 A sex organ (or reproductive organ) is any part of an animal or plant that is involved in sexual reproduction. The reproductive organs together constitute the reproductive system. In animals, the testis in the male, and the ovary in the female, a ...

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 Costa may refer to: Biology * Rib (Latin: ''costa''), in vertebrate anatomy * Costa (botany), the central strand of a plant leaf or thallus * Costa (coral), a stony rib, part of the skeleton of a coral * Costa (entomology), the leading edge of th ...

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 In insect anatomy, the juxta is an organ in the males of most Lepidoptera (i.e. moths and butterflies) that supports the aedeagus, the organ used for reproduction in insects. The juxta is located between the two valvae. Juxta has also been us ...

and can be lobes or spines, small or large, covered in bristles or nude. But unlike the similar-looking but probably

analogous Analogy (from Greek ''analogia'', "proportion", from ''ana-'' "upon, according to" lso "against", "anew"+ ''logos'' "ratio" lso "word, speech, reckoning" is a cognitive process of transferring information or meaning from a particular subject ( ...

structures found in

Ourapterygini The Ourapterygini are one of the large tribes of geometer moths in the subfamily Ennominae. The tribe was described by Charles Théophile Bruand d'Uzelle in 1846. They are particularly plentiful in the Neotropics. Ourapterygini are generally hel ...

these "furcae" are entirely or almost symmetrical in Nacophorini. The Lithinini and the

Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...

n Ennominae traditionally placed in the

Archiearinae Archiearinae is a subfamily of the geometer moth family (Geometridae). It was described by David Stephen Fletcher in 1953. Genera The subfamily contains the following genera: *'' Acalyphes'' Turner, 1926 *'' Archiearides'' D. S. Fletcher, 1953 ...

also have such symmetrical furcae, indicating the close relationship between them and the Nacophorini.

Ecology

Larval food plants are mainly

Rosidae Under the International Code of Nomenclature for algae, fungi, and plants (ICN), Rosidae is a botanical name at the rank of subclass. Circumscription of the subclass will vary with the taxonomic system being used; the only requirement being t ...

Caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...

s of the

n and

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...

n genera feed predominantly on

Myrtaceae Myrtaceae, the myrtle family, is a family of dicotyledonous plants placed within the order Myrtales. Myrtle, pōhutukawa, bay rum tree, clove, guava, acca (feijoa), allspice, and eucalyptus are some notable members of this group. All speci ...

, including '' Campomanesia'', ''

Eucalyptus ''Eucalyptus'' () is a genus of over seven hundred species of flowering trees, shrubs or mallees in the myrtle family, Myrtaceae. Along with several other genera in the tribe Eucalypteae, including '' Corymbia'', they are commonly known as euca ...

'', ''

Eugenia ''Eugenia'' is a genus of flowering plants in the myrtle family Myrtaceae. It has a worldwide, although highly uneven, distribution in tropical and subtropical regions. The bulk of the approximately 1,100 species occur in the New World tropics, ...

'' and

guava Guava () is a common tropical fruit cultivated in many tropical and subtropical regions. The common guava ''Psidium guajava'' (lemon guava, apple guava) is a small tree in the myrtle family ( Myrtaceae), native to Mexico, Central America, the ...

s (''Psidium''). A rather notorious nacophorine species from the

tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...

Americas is the Brazilian eucalyptus brown looper ('' Thyrinteina arnobia''), which can be a commercially significant pest in eucalyptus plantations.

Faboideae The Faboideae are a subfamily of the flowering plant family Fabaceae or Leguminosae. An acceptable alternative name for the subfamily is Papilionoideae, or Papilionaceae when this group of plants is treated as a family. This subfamily is wide ...

and

Mimosoideae The Mimosoideae are a traditional subfamily of trees, herbs, lianas, and shrubs in the pea family (Fabaceae) that mostly grow in tropical and subtropical climates. They are typically characterized by having radially symmetric flowers, with petals ...

have also been recorded as food plants. Caterpillars of the

n species tentatively placed in this tribe have been recorded from

Cunoniaceae Cunoniaceae is a family of 27 Genus, genera and about 335 species of woody plants in the order Oxalidales, mostly found in the tropical and wet temperate regions of the Southern Hemisphere. The greatest diversity of genera are in Australia and Ta ...

Ericaceae The Ericaceae are a family of flowering plants, commonly known as the heath or heather family, found most commonly in acidic and infertile growing conditions. The family is large, with c.4250 known species spread across 124 genera, making it th ...

Fabaceae The Fabaceae or Leguminosae,International Code of Nomenc ...

and Thymelaceae. Caterpillars of '' Declana'' from

New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 smaller islands. It is the sixth-largest island count ...

, also tentatively placed in the Nacophorini, have been found on Myrtaceae (eucalyptus, ''

Kunzea ''Kunzea'' is a genus of plants in the family Myrtaceae and is endemic to Australasia. They are shrubs, sometimes small trees and usually have small, crowded, rather aromatic leaves. The flowers are similar to those of plants in the genus '' Lep ...

'' and ''

Leptospermum ''Leptospermum'' is a genus of shrubs and small trees in the myrtle family Myrtaceae commonly known as tea trees, although this name is sometimes also used for some species of ''Melaleuca''. Most species are endemic to Australia, with the greate ...

''),

Pinaceae The Pinaceae, or pine family, are conifer trees or shrubs, including many of the well-known conifers of commercial importance such as Cedrus, cedars, firs, Tsuga, hemlocks, larches, pines and spruces. The family is included in the order Pinales, ...

–

larch Larches are deciduous conifers in the genus ''Larix'', of the family Pinaceae (subfamily Laricoideae). Growing from tall, they are native to much of the cooler temperate northern hemisphere, on lowlands in the north and high on mountains furt ...

es (''Larix''),

pine A pine is any conifer tree or shrub in the genus ''Pinus'' () of the family Pinaceae. ''Pinus'' is the sole genus in the subfamily Pinoideae. The World Flora Online created by the Royal Botanic Gardens, Kew and Missouri Botanical Garden accep ...

s (''Pinus'', notably

Monterey pine ''Pinus radiata'' ( syn. ''Pinus insignis''), the Monterey pine, insignis pine or radiata pine, is a species of pine native to the Central Coast of California and Mexico ( Guadalupe Island and Cedros island). It is an evergreen conifer in the f ...

, ''P. radiata'') and

coast Douglas-fir ''Pseudotsuga menziesii'' var. ''menziesii'', commonly known as Coast Douglas-fir, Pacific Douglas-fir, Oregon pine, or Douglas spruce, is an evergreen conifer native to western North America from west-central British Columbia, Canada southward t ...

(''Pseudotsuga menziesii'') –, and

southern beech ''Nothofagus'', also known as the southern beeches, is a genus of 43 species of trees and shrubs native to the Southern Hemisphere in southern South America (Chile, Argentina) and Australasia (east and southeast Australia, New Zealand, New Gui ...

es (Nothofagaceae). In addition, they were found on a species of ''

Olearia ''Olearia'', most commonly known as daisy-bush, is a genus of flowering plants belonging to the family Asteraceae, the largest of the flowering plant families in the world. Olearia are found in Australia, New Guinea and New Zealand. The genus i ...

'', the only

euasterid In the APG IV system (2016) for the classification of flowering plants, the name asterids denotes a clade (a monophyletic group). Asterids is the largest group of flowering plants, with more than 80,000 species, about a third of the total floweri ...

recorded as food plant of Nacophorini to date.

Systematics

The genera '' Oratha'', usually placed in the Nacophorini, is in many aspects suspiciously similar to '' Pero'' of the

and '' Rhinodia'' of the

. It is liable to be moved to either of these tribes. '' Phaeoura'', which includes ''Nacophora'' nowadays, appears to be closer to the

. As this includes the

'' Phaeoura quernaria'', Lithinini which might warrant merging with the Nacophorini would then supersede the name "Nacophorini", which would become a

junior synonym The Botanical and Zoological Codes of nomenclature treat the concept of synonymy differently. * In botanical nomenclature, a synonym is a scientific name that applies to a taxon that (now) goes by a different scientific name. For example, Linna ...

of the Ennomini. The enigmatic genus ''

Hoplosauris ''Hoplosauris'' is a genus of moths in the family Geometridae The geometer moths are moths belonging to the family Geometridae of the insect order Lepidoptera, the moths and butterflies. Their scientific name derives from the Ancient Greek '' ...

'', of uncertain placement in the

, is in some respects intermediate between the Nacophorini and the Ennomini. While the

taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...

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 In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...

''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

apomorph In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...

ies of the male genitalia, and their

caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder Sym ...

s are also very similar. It is unknown whether the somewhat more distinct Campaeini would warrant inclusion in this group too. The

Tasmania ) , nickname = , image_map = Tasmania in Australia.svg , map_caption = Location of Tasmania in AustraliaCoordinates: , subdivision_type = Country , subdi ...

n genera ''

Acalyphes ''Acalyphes'' is a monotypic moth genus in the family Geometridae. Its single species, ''Acalyphes philorites'', is found in Australia, where it feeds on pencil pine (''Athrotaxis cupressoides ''Athrotaxis cupressoides'', is also known as penci ...

'' and ''

Dirce Dirce (; , , modern Greek , meaning "double" or "cleft") was a queen of Thebes as the wife of Lycus in Greek mythology. Family Dirce was a daughter of the river-gods Achelous or Ismenus, or of Helios. Mythology After Zeus impregnated Dir ...

'' and possibly the South American ''

Archiearides Archiearides is a genus of moths in the family Geometridae The geometer moths are moths belonging to the family Geometridae of the insect order Lepidoptera, the moths and butterflies. Their scientific name derives from the Ancient Greek ''geo ...

'', traditionally placed in the

, 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

which are tentatively assigned to the Nacophorini.

Selected genera

As numerous ennomine genera have not yet been assigned to a tribe,See references in Savela (2008) the genus list should be considered preliminary. Thalaina clara

As noted above, ''

'', ''

'' and possibly ''

'' would seem to need moving here from the

, and at least some Lithinini and perhaps Campaeini seem to belong here too.

Footnotes

References

* (2007): Butterflies and Moths of the World, Generic Names and their Type-species
''Nacophora''
Retrieved 22 July 2008. * (2008)

Version of 8 March 2008. Retrieved 21 July 2008. * (2008): Characterisation of the Australian Nacophorini using adult morphology, and phylogeny of the Geometridae based on morphological characters. ''

Zootaxa ''Zootaxa'' is a peer-reviewed scientific mega journal for animal taxonomists. It is published by Magnolia Press (Auckland, New Zealand). The journal was established by Zhi-Qiang Zhang in 2001 and new issues are published multiple times a week. ...

'' 1736: 1-141
PDF abstract and excerpt
{{Taxonbar, from=Q12062272