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 use of the term in English is in the discipline of anthropology. The definition is contested, in part due to conflict ...

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 (derivative form of or "the earth"), and "measure" in referen ...

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 botanical subfamily names with "-oideae", and zo ...

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

. They are the most diverse Ennominae of Australia and are widespread in the Americas. If the African

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

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

Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...

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 (also known as the Antarctic Ocean), it contains the geographic South Pole. ...

(which was not ice-covered until a few

million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...

). In

Eurasia Eurasia ( , ) is a continental area on Earth, comprising all of Europe and Asia. According to some geographers, Physical geography, physiographically, Eurasia is a single supercontinent. The concept of Europe and Asia as distinct continents d ...

, 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 biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

, requiring only a few

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

clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...

; as this involves the

type species In International_Code_of_Zoological_Nomenclature, 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 spe ...

, 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 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 to be close ...

Ennomini __NOTOC__ The Ennomini are a tribe (biology), tribe of geometer moths in the Ennominae subfamily. They are large-bodied and rather undistinguished Ennominae, overall showing similarities to the closely related Azelinini and Nacophorini.Young (20 ...

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 from the Caberini. Research ...

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 A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

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

sensilla A sensillum (plural ''sensilla'') is an arthropod sensory organ protruding from the cuticle of exoskeleton, or sometimes lying within or beneath it. Sensilla appear as small hairs or pegs over an individual's body. Inside each sensillum there are ...

e shaped like stout pegs and

sensillae basiconicae A sensillum (plural ''sensilla'') is an arthropod sensory organ protruding from the cuticle of exoskeleton, or sometimes lying within or beneath it. Sensilla appear as small hairs or pegs over an individual's body. Inside each sensillum there are ...

on the

flagellomere An antenna (plural: antennae) is one of a pair of appendages used for sensing in arthropods. Antennae are sometimes referred to as ''feelers''. Antennae are connected to the first one or two segments of the arthropod head. They vary widely in ...

s or

rami Rami or Ramy may refer to: People Given name In music *Rami (singer), Japanese singer * Rami El-Kaleh (born 1983), Libyan-Irish musician *Rami Jaffee (born 1969), American keyboardist *Rami Kleinstein (born 1962), Israeli singer and composer *Ram ...

. 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 (: thoraces or thoraxes) or chest is a part of the anatomy of 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 di ...

, 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 (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae, ...

e on the third

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

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, also known as a reproductive organ, is a part of an organism that is involved in sexual reproduction. Sex organs constitute the primary sex characteristics of an organism. Sex organs are responsible for producing and transporting ...

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 Boarmiini (also often called ''Cleorini'') are a large tribe (biology), tribe of geometer moths in the Ennominae subfamily. Description and systematics This family is sometimes massively expanded, with the closely related Bistonini, Bupalini ...

, 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 An aedeagus ( or aedeagi) is a reproductive organ of male arthropods through which they secrete sperm from the testes during copulation (zoology), copulation with a female. It can be thought of as the insect equivalent of a mammal's penis, th ...

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 use ...

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

analogous Analogy is a comparison or correspondence between two things (or two groups of things) because of a third element that they are considered to share. In logic, it is an inference or an argument from one particular to another particular, as oppose ...

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 held ...

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

Tasmania Tasmania (; palawa kani: ''Lutruwita'') is an island States and territories of Australia, state of Australia. It is located to the south of the Mainland Australia, Australian mainland, and is separated from it by the Bass Strait. The sta ...

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 th ...

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 ...

s of the Australian and South American 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 ...

, including '' Campomanesia'', ''

Eucalyptus ''Eucalyptus'' () is a genus of more than 700 species of flowering plants in the family Myrtaceae. Most species of ''Eucalyptus'' are trees, often Mallee (habit), mallees, and a few are shrubs. Along with several other genera in the tribe Eucalyp ...

'', ''

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 ( ), also known as the 'guava-pear', 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), nativ ...

s (''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 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 widely ...

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 petal ...

have also been recorded as food plants. Caterpillars of the African species tentatively placed in this tribe have been recorded from

Cunoniaceae Cunoniaceae is a family of 27 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 Tasmania ...

Ericaceae The Ericaceae () are a Family (biology), 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 about 4,250 known species spread acros ...

Fabaceae Fabaceae () or Leguminosae,International Code of Nomen ...

and Thymelaceae. Caterpillars of '' Declana'' from New Zealand, 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 Endemism, endemic to Australasia. They are shrubs, sometimes small trees and usually have small, crowded, rather Aroma compound, aromatic leaves. The flowers are similar to those of p ...

'' 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, Pinyon_pine, piñons, larches, pines and spruces. The family is incl ...

–

larch Larches are deciduous conifers in the genus ''Larix'', of the family Pinaceae (subfamily Laricoideae). Growing from tall, they are native to the cooler regions of the northern hemisphere, where they are found in lowland forests in the high la ...

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. ''World Flora Online'' accepts 134 species-rank taxa (119 species and 15 nothospecies) of pines as cu ...

s (''Pinus'', notably

Monterey pine ''Pinus radiata'' (synonym (taxonomy), syn. ''Pinus insignis''), the Monterey pine, insignis pine or radiata pine, is a species of pine native to the Central Coast (California), Central Coast of California and Mexico (on Guadalupe Island and Ced ...

, ''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, found across southern South America (Chile, Argentina) and east and southeast Australia, New Zealand, New Guin ...

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 Asterids are a large clade (monophyletic group) of flowering plants, composed of 17 orders and more than 80,000 species, about a third of the total flowering plant species. The asterids are divided into the unranked clades lamiids (8 orders) and ...

recorded as food plant of Nacophorini to date.

Systematics

The genera ''

Oratha ''Oratha'' is a genus of moths in the family Geometridae The geometer moths are moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as ...

'', 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 ''Phaeoura'' is a genus of moths in the family Geometridae The geometer moths are moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified ...

'', which includes ''Nacophora'' nowadays, appears to be closer to the

. As this includes the

Phaeoura quernaria ''Phaeoura quernaria'', the oak beauty, is a moth of the family Geometridae. The species was first described by James Edward Smith in 1797. It is found in eastern North America, occurring as far west as east-central Alberta. The habitat consist ...

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

junior synonym In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently. ...

of the Ennomini. The enigmatic genus '' Hoplosauris'', of uncertain placement in the

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

taxonomic 280px, Generalized scheme of taxonomy Taxonomy is a practice and science concerned with classification or categorization. Typically, there are two parts to it: the development of an underlying scheme of classes (a taxonomy) and the allocation ...

and

systematic Systematic may refer to: Science * Short for systematic error * Systematic fault * Systematic bias, errors that are introduced by an inaccuracy inherent to the system Economy * Systematic trading, a way of defining trade goals, risk control ...

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 Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

''Metrocampa'' (at least '' "M." ada'' and '' "M." biplaga'') appear very close to nacophorine genera – to '' Thalaina'', and to ''

Conosara ''Conosara'' is a genus of moths in the family Geometridae The geometer moths are moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified a ...

'' 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 ha ...

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 ...

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

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''). Both the genus and species were first described by ...

'' 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 Di ...

'' and possibly the South American ''

Archiearides Archiearides is a genus of moths in the family Geometridae The geometer moths are moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified as ...

'', traditionally placed in the

, seem to be close relatives of the Australian nacophorines '' Niceteria'' and ''

Paralaea ''Paralaea'' is a genus of moths in the family Geometridae The geometer moths are moth Moths are a group of insects that includes all members of the order Lepidoptera that are not Butterfly, butterflies. They were previously classified a ...

'' 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.

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 Moth tribes