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Mimallonidae Burmeister (mimallonids), sometimes known as "sack-bearer" moths for the
larva A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle. The ...
l case-building behavior, are a family of
Lepidoptera Lepidoptera ( ) is an order (biology), order of insects that includes butterfly, butterflies and moths (both are called lepidopterans). About 180,000 species of the Lepidoptera are described, in 126 Family (biology), families and 46 Taxonomic r ...
containing over 300 named species in 43 genera. These moths are found only in the
New World The term ''New World'' is often used to mean the majority of Earth's Western Hemisphere, specifically the Americas."America." ''The Oxford Companion to the English Language'' (). McArthur, Tom, ed., 1992. New York: Oxford University Press, p. 3 ...
, with most taxa occurring in the
Neotropics The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropics, tropical Ecoregion#Terrestrial, terrestrial ecoregions of the Americas and the entire South American temperat ...
. Adult moths are externally similar to those belonging to some of the other
Macroheterocera The Macroheterocera are a well supported clade of moths that are closely related to butterflies and other macro-moths. Taxonomy The Macroheterocera includes the following superfamilies: * Mimallonoidea – sack bearers (variously included in b ...
families
Bombycoidea Bombycoidea is a superfamily of moths. It contains the silk moths, emperor moths, sphinx moths, and relatives. The Lasiocampoidea The Lasiocampidae are a family of moths also known as eggars, tent caterpillars, snout moths (although this also ...
and
Drepanoidea Drepanoidea is the superfamily of "hook tip moths". See Minet and Scoble (1999) for a comprehensive overview. References * Minet, J. and Scoble, M.J. (1999). The Drepanoid/Geometroid Assemblage. Ch. 17 in Kristensen, N.P. (Ed.). ''Lepidoptera, ...
, and thus have been variously treated as belonging to either one of these or other superfamilies.


Distribution

Mimallonids are restricted to the New World, and are distributed in North America, Central America, South America, and the Caribbean (Cuba and The Bahamas). The vast majority of genera and species are found in 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 ...
regions of the New World, with only five described species from the United States.


Biology

Not much has been published on the natural history of adult Mimallonidae, though most species are thought to be
nocturnal Nocturnality is an animal behavior characterized by being active during the night and sleeping during the day. The common adjective is "nocturnal", versus diurnal meaning the opposite. Nocturnal creatures generally have highly developed sens ...
. At least three species have diurnal males. Young
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 live inside of folded leaves or beneath silken networks, and build portable or semi-portable cases out of
silk Silk is a natural protein fiber, some forms of which can be woven into textiles. The protein fiber of silk is composed mainly of fibroin and is produced by certain insect larvae to form cocoons. The best-known silk is obtained from the coc ...
,
frass Frass refers loosely to the more or less solid excreta of insects, and to certain other related matter. Definition and etymology ''Frass'' is an informal term and accordingly it is variously used and variously defined. It is derived from the ...
, and plant material as they grow. The cases are open on either end and vary from irregular in structure to spindle-shaped. The openings of the cases can be blocked by the head and/or the flattened anal plate of the last segment of the body. The sack-like case-making behavior of the caterpillars have earned them the common name "sack-bearers". Mimallonidae larvae feed on several families of plants, including (but not limited to):
Anacardiaceae The Anacardiaceae, commonly known as the cashew family or sumac family, are a family of flowering plants, including about 83 genera with about 860 known species. Members of the Anacardiaceae bear fruits that are drupes and in some cases produce ...
,
Clusiaceae The Clusiaceae or Guttiferae Juss. (1789) (''nom. alt. et cons.'' = alternative and valid name) are a family of plants including 13 genera and ca 750 species. Several former members of Clusiacae are now placed in Calophyllaceae and Hypericaceae. ...
,
Combretaceae The Combretaceae, often called the white mangrove family, are a family of flowering plants in the order Myrtales. The family includes about 530 species of trees, shrubs, and lianas in ca 10 genera. The family includes the leadwood tree, ''Combre ...
,
Fagaceae The Fagaceae are a family of flowering plants that includes beeches, chestnuts and oaks, and comprises eight genera with about 927 species. Fagaceae in temperate regions are mostly deciduous, whereas in the tropics, many species occur as evergre ...
,
Melastomataceae Melastomataceae is a family of dicotyledonous flowering plants found mostly in the tropics (two-thirds of the genera are from the New World tropics) comprising c. 175 genera and c. 5115 known species. Melastomes are annual or perennial herbs, sh ...
,
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 ...
and
Rubiaceae The Rubiaceae are a family of flowering plants, commonly known as the coffee, madder, or bedstraw family. It consists of terrestrial trees, shrubs, lianas, or herbs that are recognizable by simple, opposite leaves with interpetiolar stipules ...
. Research at the Area de Conservación, Guanacaste, Costa Rica has resulted in the rearing of many species of Mimallonidae, including some from several additional families of plants.


Systematics and evolution

The current consensus, especially in works based on
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
results, consider Mimallonidae to be the sole family of Mimallonoidea. Other phylogenetic studies of all major Lepidoptera lineages support the placement of Mimallonidae as sister to all Macroheterocera, but within the broader clade
Obtectomera The Obtectomera is a clade of macro-moths and butterflies, comprising over 100,000 species in at least 12 superfamilies. Taxonomy The Obtectomera includes the following 12 superfamilies: * Whalleyanoidea Minet, 1991 * Thyridoidea Herrich-Sc ...
. This phylogenetic placement means that mimallonid moths are not more closely related to any one family of Lepidoptera, but are equally related (share a common ancestor with) all members of the extremely diverse clade Macroheterocera. Morphological features, particularly of the larvae, support the uniqueness of this family and distinctness from Macroheterocera. The systematic relationships with Mimallonidae have been subject to two revisions, resulting in two different classification schemes. William Schaus revised the family and named most of the genera, he then separated the genera into two subfamilies: Lacosominae and Mimalloninae. These subfamilies were based on the presence (Lacosominae) or the absence (Mimalloninae) of the
frenulum A frenulum (or frenum, plural: frenula or frena, from the Latin ''frēnulum'', "little bridle", the diminutive of ''frēnum'') is a small fold of tissue that secures the motion of a mobile organ in the body. In human anatomy Frenula on the h ...
. It was later realized that this character varies within genera, and thus was deemed a sympleisiomorphy. Using
molecular phylogenetics Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
, specifically with the technique of anchored hybrid enrichment, the family was reorganized based on the recognition and naming of robustly supported
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, ...
s, which were further strengthened with morphological
apomorphies 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 ...
. The clades which were robustly supported across all analyses were assigned
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 ...
status, resulting in recognition of seven subfamilies in Mimallonidae. Using both molecular and morphological phylogenetics, all 42
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 ...
(including some newly described ones) were assigned to the named clades. The 300+ species of the family, too, have been formally classified according to their phylogenetic arrangement. Recently, many of the species formally placed in the genus ''Cicinnus'' have been transferred to ''Gonogramma'', making ''Gonogramma'' one of the most diverse mimallonid genera.


Subfamilies, tribes, and genera

Below are the named clades of Mimallonidae, with their constituent genera assigned to their subfamily and tribe (wherever applicable). * Zaphantinae ** '' Zaphanta'' Dyar, 1910 * Roelofinae **'' Roelofa'' Schaus, 1928 *Meneviinae **'' Cunicumara'' St Laurent, 2016 **''Menevia'' Schaus, 1928 **'' Tolypida'' Schaus, 1928 *Aurorianinae **'' Auroriana'' St Laurent and C. Mielke, 2016 * Mimalloninae **'' Eadmuna'' Schaus, 1928 ** '' Macessoga'' Schaus, 1928 ** '' Mimallo'' Hübner, 1820 **'' Mimallodes'' St Laurent and Becker, 2020 **'' Tostallo'' St Laurent and C. Mielke, 2016 * Lacosominae * Alheitini ** '' Adalgisa'' Schaus, 1928 ** '' Alheita'' Schaus, 1928 ** '' Arianula'' Herbin, 2012, ** '' Fatellalla'' St Laurent and Kawahara, 2019 ** '' Herbinalla'' St Laurent and Kawahara, 2018 ** '' Tarema'' Schaus, 1896 ** '' Thaelia'' Herbin, 2016 * Lacosomini ** '' Citralla'' St Laurent and Kawahara, 2019 ** '' Lacosoma'' Grote, 1864 ** '' Vanenga'' Schaus, 1928 * Trogopterini ** '' Reinmara'' Schaus, 1928 ** '' Trogoptera'' Herrich-Schäffer,
856 __NOTOC__ Year 856 (Roman numerals, DCCCLVI) was a leap year starting on Wednesday (link will display the full calendar) of the Julian calendar. Events By place Byzantine Empire * March 15 – Emperor Michael III overthrows the reg ...
* Druenticinae * Druenticini: ** '' Druentica'' Strand, 1932 ** '' Lepismalla'' St Laurent and Kawahara, 2019 ** '' Micrallo'' St Laurent and C. Mielke, 2016 ** '' Pamea'' Walker, 1855 ** '' Procinnus'' Herbin, 2016 ** '' Ulaluma'' St Laurent and Kawahara, 2018 * Luramini ** '' Lurama'' Schaus, 1928 ** '' Ulmara'' Schaus, 1928 * Cicinninae * Bedosiini ** '' Bedosia'' Schaus, 1928 ** '' Bedosiallo'' St Laurent and Kawahara, 2018 * Cicinnini **'' Aceclostria'' Vuillot, 1893 ** '' Aleyda'' Schaus, 1928 ** '' Arcinnus'' Herbin, 2016 ** '' Cicinnus'' Blanchard, 1852 **'' Cerradocinnus'' St Laurent, MIelke, and Kawahara, 2020 **'' Gonogramma'' Boisduval, 1872 **'' Euphaneta'' Schaus, 1928 **'' Isoscella'' St Laurent and Carvalho, 2017 **'' Roelmana'' Schaus, 1928 * Psychocampini ** '' Biterolfa'' Schaus, 1928 ** '' Psychocampa'' Grote and Robinson, 1867


Importance to humans

Mimallonidae have been reported as pests of some economically significant crops. In Surinam, ''Mimallo amilia'' has been noted as a pest of
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 ...
('' Psidium gaujava''). In
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, ''M. amilia'' is a known pest of eucalyptus (''
Eucalyptus urophylla ''Eucalyptus urophylla'', commonly known as Timor white gum, Timor mountain gum, popo or ampupu, is a species of eucalypt native to islands of the Indonesian Archipelago and Timor. It is also common in other countries with humid and subhumid tr ...
'') and ''Psychocampa callipius'' a pest of
cashew The cashew tree (''Anacardium occidentale'') is a tropical evergreen tree native to South America in the genus ''Anacardium'' that produces the cashew seed and the cashew apple accessory fruit. The tree can grow as tall as , but the dwarf cult ...
(''Anacardium occidentale''). In addition to some Mimallonidae species being regarded as pests, others have been considered as potential biological controls of invasive plants. Species belonging to the genus ''Druentica'' have been considered as potential control agents of ''
Miconia calvescens ''Miconia calvescens'', the velvet tree, miconia, or bush currant, is a species of flowering plant in the family Melastomataceae. It is native to Mexico and Central and South America and it has become one of the world's most invasive species. M ...
'' (where it is invasive throughout the
Pacific Islands Collectively called the Pacific Islands, the islands in the Pacific Ocean are further categorized into three major island groups: Melanesia, Micronesia, and Polynesia. Depending on the context, the term ''Pacific Islands'' may refer to one of se ...
, including
Hawaii Hawaii ( ; haw, Hawaii or ) is a state in the Western United States, located in the Pacific Ocean about from the U.S. mainland. It is the only U.S. state outside North America, the only state that is an archipelago, and the only stat ...
and
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 ''
Clidemia hirta ''Miconia crenata'', (syn. ''Clidemia hirta''), commonly called soapbush, clidemia or Koster's curse, is a perennial shrub. It is an invasive plant species in many tropical regions of the world, creating serious damage. Description ''Miconia cre ...
'' in Hawaii. ''Aceclostria mus'' has been evaluated as a potential control agent of the
Brazilian pepper tree ''Schinus terebinthifolia'' is a species of flowering plant in the cashew family, Anacardiaceae, that is native to subtropical and tropical South America. Common names include Brazilian peppertree, aroeira, rose pepper, broadleaved pepper tree, ...
(''Schinus terebinthifolius'') in the United States.


References


Sources

*''Firefly Encyclopedia of Insects and Spiders'', edited by Christopher O'Toole, , 2002
Natural History Museum Lepidoptera genus database


External links



Page of images of Mimallonidae from Area de Conservación Guanacaste,
Costa Rica Costa Rica (, ; ; literally "Rich Coast"), officially the Republic of Costa Rica ( es, República de Costa Rica), is a country in the Central American region of North America, bordered by Nicaragua to the north, the Caribbean Sea to the no ...
. {{Taxonbar, from1=Q18536944, from2=Q952639 Lepidoptera superfamilies Macroheterocera