Hedylidae, the "American moth-butterflies", is a

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

of insects in the order

Lepidoptera Lepidoptera ( ) or lepidopterans is an order (biology), order of winged insects which includes butterflies and moths. About 180,000 species of the Lepidoptera have been described, representing 10% of the total described species of living organ ...

, formerly representing the superfamily Hedyloidea. They have traditionally been viewed as an

extant Extant or Least-concern species, least concern is the opposite of the word extinct. It may refer to: * Extant hereditary titles * Extant literature, surviving literature, such as ''Beowulf'', the oldest extant manuscript written in English * Exta ...

sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...

of the butterfly superfamily

Papilionoidea Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...

, but a 2014

phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...

analysis has suggested Hedylidae is a subgroup of

, and not a sister group, and are more accurately referred to as butterflies rather than moths. They are represented by a single

Neotropical The Neotropical realm is one of the eight biogeographic realms constituting Earth's land surface. Physically, it includes the tropical terrestrial ecoregions of the Americas and the entire South American temperate zone. Definition In biogeogra ...

genus, ''Macrosoma'', with 35 currently recognized species.

Taxonomy and systematics

Hedylidae were previously treated as a

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 ...

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 suborder Heterocera, but the group is Paraphyly, paraphyleti ...

: Oenochrominae, the "Hedylicae". Prout considered they might even merit treatment as their own family. Scoble first considered them to be a hitherto unrecognised group of butterflies and also suggested Hedylidae might possibly constitute the

of the "true" butterflies (

), rather than of (Hesperioidea + Papilionoidea). Weintraub and Miller argued against this placement (but see). In 1995, Weller and Pashley found that molecular data did indeed place Hedylidae with the butterflies and a more comprehensive study in 2005 based on 57 exemplar

taxa In biology, a taxon (back-formation from ''taxonomy''; : taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular name and ...

, three

genes In biology, the word gene has two meanings. The Mendelian gene is a basic unit of heredity. The molecular gene is a sequence of nucleotides in DNA that is transcribed to produce a functional RNA. There are two types of molecular genes: protei ...

and 99 morphological characters, recovered the genus ''Macrosoma'' as sister to the ("Papilionoidea" + Hesperioidea). However, the most recent

analyses shows that skippers are true butterflies and therefore within the clade Papilionoidea, whereas the hedylids are a sister group that may be closely related to the obtectomeran moths. This is contrary to some earlier studies that had shown both the skippers and hedylids as being nested within the Papilionoidea. Since there are no obvious gaps between supposed species groups, according to basic morphological structure, Scoble (1986) synonymised the five pre-existing genera of Hedylidae (33 of which had been described in ''Phellinodes'') into the single genus ''Macrosoma''. However, a phylogenetic analysis of all ''Macrosoma'' species is still needed.

Nomenclatural notes

zoological nomenclature The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its formal author, t ...

, numerous junior

homonyms In linguistics, homonyms are words which are either; ''homographs''—words that mean different things, but have the same spelling (regardless of pronunciation), or ''homophones''—words that mean different things, but have the same pronunciatio ...

of ''Macrosoma'' (Hübner, 1818) exist, (''Macrosoma'' Leach 1819 (a

reptile Reptiles, as commonly defined, are a group of tetrapods with an ectothermic metabolism and Amniotic egg, amniotic development. Living traditional reptiles comprise four Order (biology), orders: Testudines, Crocodilia, Squamata, and Rhynchocepha ...

), ''Macrosoma'' de Haan 1826 (

Odonata Odonata is an order of predatory flying insects that includes the dragonflies and damselflies (as well as the '' Epiophlebia'' damsel-dragonflies). The two major groups are distinguished with dragonflies (Anisoptera) usually being bulkier with ...

), ''Macrosoma'' Robineau-Desvoidy 1830 (Diptera:

Muscidae Muscidae are a family of flies found in the superfamily Muscoidea. Muscidae, some of which are commonly known as house flies or stable flies due to their synanthropy, are worldwide in distribution and contain almost 4,000 described species i ...

), ''Macrosoma'' Brandt 1835 (

Coelenterata Coelenterata is a term encompassing the animal phyla Cnidaria (corals, true jellies, sea anemones, sea pens, and their relatives) and Ctenophora (comb jellies). The name comes , referring to the hollow body cavity common to these two phyla. Th ...

), ''Macrosoma'' Hope 1837 (

Coleoptera Beetles are insects that form the Taxonomic rank, order Coleoptera (), in the superorder Holometabola. Their front pair of wings are hardened into wing-cases, elytra, distinguishing them from most other insects. The Coleoptera, with about 40 ...

), ''Macrosoma'' Lioy 1864 or 1865 (Diptera:

Sarcophagidae Sarcophagidae () are a family (biology), family of fly, flies commonly known as flesh flies. They differ from most flies in that they are Ovoviviparity, ovoviviparous, opportunistically depositing hatched or hatching maggots instead of eggs on c ...

), ''Macrosoma'' Hammer 1979 (

Acarina Mites are small arachnids (eight-legged arthropods) of two large orders, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as ea ...

: Oribatidae). To add to this potential confusion in lists of names, there exist two junior homonyms of ''Hedyle'' Guenée, 1857: ''Hedyle'' Bergh, 1895 (

Opisthobranchia Opisthobranchs () is a now informal name for a large and diverse group of specialized complex gastropods which used to be united in the subclass Opisthobranchia. That taxon is no longer considered to represent a monophyletic grouping. Euopisth ...

, "Acochlidioidea",

Hedylopsidae Hedylopsis is a genus of sea slugs, marine gastropod mollusks within the clade Acochlidiacea. Taxonomy Hedylopsidae has been classified within the superfamily Hedylopsoidea in the taxonomy of Bouchet & Rocroi (2005). Sensu Schrödl & Neuss ...

; currently placed in the genus ''

Hedylopsis Hedylopsis is a genus of sea slugs, marine gastropod mollusks within the clade Acochlidiacea. Taxonomy Hedylopsidae has been classified within the superfamily Hedylopsoidea in the taxonomy of Bouchet & Rocroi (2005). Sensu Schrödl & Neusse ...

'' Thiele, 1931), and ''Hedyle'' Malmgren 1865 (a

polychaete Polychaeta () is a paraphyletic class of generally marine Annelid, annelid worms, common name, commonly called bristle worms or polychaetes (). Each body segment has a pair of fleshy protrusions called parapodia that bear many bristles, called c ...

worm). The sea slug family name "Hedylidae Bergh, 1895" (type species ''Hedyle weberi'' Bergh, 1895) is thus also invalid.

Morphology and identification

The eggs of hedylid moths have an upright configuration and are variable in shape: in ''Macrosoma inermis'' they are particularly narrow and spindle-shaped,Scoble, M.J. (1990a). A catalogue of the Hedylidae (Lepidoptera: Hedyloidea), with descriptions of two new species. ''Entomologica Scandinavica'', 21: 113-119. resembling those of some

Pieridae The Pieridae are a large family (biology), family of butterflies with about 76 genera containing about 1,100 species, mostly from Afrotropical realm, tropical Africa and Indomalayan realm, tropical Asia with some varieties in the more northern ...

, and in the case of ''M. tipulata'' they are more barrel-shaped, like certain

Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species ha ...

. The

larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...

e resemble (probably convergently) those of

Apaturinae The Apaturinae are a subfamily of butterflies that includes many species commonly called emperors. Description Strikingly-coloured, with cryptic underwing. A distinguishing character of the subfamily is the green proboscis. Agathina emperor ( ...

. Adult hedylids resemble geometer moths. They share many morphological and genetic characteristics with both the superfamilies

and the

Hesperioidea Skippers are a group of butterflies placed in the family Hesperiidae within the order Lepidoptera (moths and butterflies). They were previously placed in a separate superfamily, Hesperioidea, but have since been placed in the superfamily Papilion ...

. The

abdomen 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 ...

is very long and slim, like many

butterflies of the subfamilies

Ithomiinae Ithomiini is a butterfly tribe in the nymphalid subfamily Danainae. It is sometimes referred to as the tribe of clearwing butterflies or glasswing butterflies. Some authors consider the group to be a subfamily (Ithomiinae). These butterflies are ...

and

Heliconiinae The Heliconiinae, commonly called heliconians or longwings, are a subfamily of the brush-footed butterflies (family (biology), family Nymphalidae). They can be divided into 45–50 genera and were sometimes treated as a separate family Heliconi ...

, hence the name of one ''Macrosoma'' species "''heliconiaria''". Unlike other butterflies, however, the antennae are un-

clubbed ''Clubbed'' is a 2008 British drama film about a 1980s factory worker who takes up a job as a club doorman, written by Geoff Thompson and directed by Neil Thompson. Plot In 1984, Danny - a lonely factory worker intimidated by life - is battere ...

, but rather

filiform Filiform, thread or filament like, can refer to: * Filiform, a common term used in botany to describe a thread-like shape *Filiform, or filiform catheter, a medical device whose component parts or segments are all cylindrical and more or less un ...

or bipectinate. Unlike the

, in which they had been placed by the geometer expert L.B.Prout, hedylids lack tympanic organs at the base of the abdomen, but have them on the wings (see under

Behaviour Behavior (American English) or behaviour (British English) is the range of actions of Individual, individuals, organisms, systems or Artificial intelligence, artificial entities in some environment. These systems can include other systems or or ...

). Unlike other butterflies, however (except the unique case of the remarkable Australian skipper butterfly '' Euschemon rafflesia'', whose males possess these structures), the single-spined

frenulum A frenulum or frenum (: 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 human ...

and

retinaculum A retinaculum (: retinacula) is a band of thickened deep fascia around tendons that holds them in place. It is not part of any muscle and primarily functions to stabilize tendons. The term retinaculum is Neo-Latin, derived from the Latin verb ''re ...

are not lost or reduced in males, except in three ''Macrosoma'' species where there is no functional wing coupling system. The retinaculum is always lost in females, and the frenulum may be

vestigial Vestigiality is the retention, during the process of evolution, of genetically determined structures or attributes that have lost some or all of the ancestral function in a given species. Assessment of the vestigiality must generally rely on co ...

. The family have been fully catalogued and illustrated in an identification guide.Scoble, M.J. (1990b). An identification guide to the Hedylidae (Lepidoptera: Hedyloidea). ''Entomologica Scandinavica'', 21: 121-158.

Butterfly-like characteristics of Hedylidae

#" Mesoscutum" with "secondary line of weakness" near median "notal" wing process, as in some representatives of Papilionidea and Hesperioidea (potentially unique butterfly character; # Mesothoracic

aorta The aorta ( ; : aortas or aortae) is the main and largest artery in the human body, originating from the Ventricle (heart), left ventricle of the heart, branching upwards immediately after, and extending down to the abdomen, where it splits at ...

with horizontal chamber, as in other butterflies (not Papilionidae), but as also in

Cossidae The Cossidae, the cossid millers or carpenter millers, make up a family of mostly large miller moths. This family contains over 110 genera with almost 700 known species, and many more species await description. Carpenter millers are Lepidopter ...

; # Metathoracic "furca" resembling a blunt arrowhead; this a variable but potentially unique character in butterflies; #Second ''median plate'' of

forewing Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindwi ...

base lying partly under the base of

vein Veins () are blood vessels in the circulatory system of humans and most other animals that carry blood towards the heart. Most veins carry deoxygenated blood from the tissues back to the heart; exceptions are those of the pulmonary and feta ...

"1A+2A", unlike the configuration in moths; #"Postspiracular bar" on first abdominal segment; #Female genitalic "anterior apophyses" reduced; #Male genitalia relatively "deep" dorso-ventrally; #Abdomen curved (especially in males), as in papilionoids; #Abdominal first tergal segment is strongly "pouched" (Scoble 1986; as also in

Thyatirinae The Thyatirinae, or false owlet moths, are a subfamily of the moth family Drepanidae with about 200 species described. Until recently, most classifications treated this group as a separate family called Thyatiridae. Taxonomy References * * ...

moths; # "Precoxal" sulcus joining "marginopleural" sulcus; #Male Foreleg pretarsus lost, thus fused into two elements as in

nymphalid The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species have ...

butterflies, with the mid and hindlegs used for perching, but apparently redeveloped in hesperiids; #Egg upright, spindle-shaped and ribbed as in some Pieridae (e.g. the orange tip butterfly), some other butterflies, and as in some moth groups also; #

Larva A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect development such as insects, some arachnids, amphibians, or cnidarians typically have a larval phase ...

with "anal comb", as in some Hesperioidea (not however Megathyminae) and Pieridae, but not in other Papilionoidea except one species (and also independently in

Tortricidae The Tortricidae are a family of moths, commonly known as tortrix moths or leafroller moths, in the order Lepidoptera. This large family has over 11,000 species described, and is the sole member of the superfamily Tortricoidea, although the genu ...

), that is used for propulsion of

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 ...

away from the caterpillar; #Caterpillar with horn-like processes and a "bifid" tail as in many Nymphalidae; #Caterpillar with "secondary

setae 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 ...

", as in Pieridae; #

Ventral Standard anatomical terms of location are used to describe unambiguously the anatomy of humans and other animals. The terms, typically derived from Latin or Greek roots, describe something in its standard anatomical position. This position prov ...

proleg A proleg is a small, fleshy, stub structure found on the ventral surface of the abdomen of most larval forms of insects of the Order (biology), order Lepidoptera, though they can also be found on larvae of insects such as symphyta, sawflies. In ...

"crochet" hooks not forming a complete circle, unlike configuration in hesperiids and papilionoids; #

Pupa A pupa (; : pupae) is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their life cycle, the stages th ...

affixed to the substrate via a silken girdle around the 1st abdominal segment, like in Pieridae (as also in some

, especially the subfamily

Sterrhinae Sterrhinae is a large subfamily of geometer moths (family Geometridae) with some 3,000 described species, with more than half belonging to the taxonomically difficult, very diverse genera, ''Idaea'' and ''Scopula'' (Hausmann, 2004; Sihvonen, 2005 ...

(in which the girdle is around the abdomen), but lost in many

); #

l cocoon lost, as in papilionoids, and a few other groups of Lepidoptera; #"Temporal cleavage line" lost in the pupa (as in papilionoids).

Distribution

Hedylidae range in

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

south from central

Mexico Mexico, officially the United Mexican States, is a country in North America. It is the northernmost country in Latin America, and borders the United States to the north, and Guatemala and Belize to the southeast; while having maritime boundar ...

and in

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

through the

Amazon Amazon most often refers to: * Amazon River, in South America * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon (company), an American multinational technology company * Amazons, a tribe of female warriors in Greek myth ...

from southern

Peru Peru, officially the Republic of Peru, is a country in western South America. It is bordered in the north by Ecuador and Colombia, in the east by Brazil, in the southeast by Bolivia, in the south by Chile, and in the south and west by the Pac ...

(where there are a full 26 species, up to 12 at a single site: to central

Bolivia Bolivia, officially the Plurinational State of Bolivia, is a landlocked country located in central South America. The country features diverse geography, including vast Amazonian plains, tropical lowlands, mountains, the Gran Chaco Province, w ...

and southwestern

Brazil Brazil, officially the Federative Republic of Brazil, is the largest country in South America. It is the world's List of countries and dependencies by area, fifth-largest country by area and the List of countries and dependencies by population ...

). In the

Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered around the Caribbean Sea in the Atlantic Ocean, North Atlantic Ocean, mostly overlapping with the West Indies. Bordered by North America to the north, Central America ...

, they also occur in

Cuba Cuba, officially the Republic of Cuba, is an island country, comprising the island of Cuba (largest island), Isla de la Juventud, and List of islands of Cuba, 4,195 islands, islets and cays surrounding the main island. It is located where the ...

Jamaica Jamaica is an island country in the Caribbean Sea and the West Indies. At , it is the third-largest island—after Cuba and Hispaniola—of the Greater Antilles and the Caribbean. Jamaica lies about south of Cuba, west of Hispaniola (the is ...

, and

Trinidad Trinidad is the larger, more populous island of the Republic of Trinidad and Tobago, the country. The island lies off the northeastern coast of Venezuela and sits on the continental shelf of South America. It is the southernmost island in ...

Behaviour

Hedylids are attracted to artificial lights, but occasionally some species can be found flying by day. Thus, they may be involved in some

mimicry In evolutionary biology, mimicry is an evolved resemblance between an organism and another object, often an organism of another species. Mimicry may evolve between different species, or between individuals of the same species. In the simples ...

complexes with

(e.g. the female only of ''Macrosoma lucivittata''). A few species are white and resemble pierid butterflies (e.g. ''Macrosoma napiaria''). Based on a study of ''Macrosoma heliconiaria'', it has been found that hedylids have tympanic organs on their

forewings Insect wings are adult outgrowths of the insect exoskeleton that enable insect flight, insects to fly. They are found on the second and third Thorax (insect anatomy), thoracic segments (the mesothorax and metathorax), and the two pairs are often ...

for hearing apparently homologous to the "Vogel's organ" in some

that would help them evade bats at night. They have been shown to exhibit typical moth evasive behaviour towards bats such as erratic spiralling movements and dives. The resting posture is often at a curious angle, with the thorax tilted and the posterior edge of the hindwings nearly touching the substrate (Scoble, 1986). The larvae which lack the prominent horns in the first

instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...

tend to rest on the

midrib A primary vein, also known as the midrib, is the main vascular structure running through the center of a leaf. The primary vein is crucial for the leaf’s efficiency in photosynthesis and overall health, as it ensures the proper flow of material ...

of the leaf and often skeletonise leaves or at either side produce an untidy patchwork of holes. The elegant pupa is attached by a cremaster and silken girdle and sometimes resembles a bird dropping.

List of species

This list of species is largely based on

phenetic In biology, phenetics (; ), also known as taximetrics, is an attempt to classify organisms based on overall similarity, usually with respect to Morphology (biology), morphology or other observable traits, regardless of their phylogeny or evoluti ...

characters.

Biology and host plants

The life history of ''Macrosoma heliconiaria'' was originally described from plants of ''Byttneria aculeata'' in

. This was a historical breakthrough into the biology of hedylids. In this study, Kendall commented notably "I thought the larvae might represent a satyr species, but when the first larva pupated I was sure it was a pierid. The first adult emerged as a complete surprise. The pupa...is secured by girdle and cremaster, not unlike a pierid". ''Macrosoma cascaria'' was later also reared on this plant in

Panama Panama, officially the Republic of Panama, is a country in Latin America at the southern end of Central America, bordering South America. It is bordered by Costa Rica to the west, Colombia to the southeast, the Caribbean Sea to the north, and ...

. More life histories are now known. From these data, known host plants span a wide range of (according to the

APG II system The APG II system (Angiosperm Phylogeny Group II system) of plant classification is the second, now obsolete, version of a modern, mostly Molecular phylogenetics, molecular-based, list of systems of plant taxonomy, system of plant taxonomy that ...

)

rosid The rosids are members of a large clade (monophyletic group) of flowering plants, containing about 70,000 species, more than a quarter of all angiosperms. The clade is divided into 16 to 20 orders, depending upon circumscription and classificatio ...

dicotyledonous The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, ...

plants, including the

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

Myrtales The Myrtales are an order of flowering plants in the malvid clade of the rosid group of dicotyledons. Well-known members of Myrtales include: myrtle, pōhutukawa, bay rum tree, clove, guava, acca (feijoa), allspice, eucalyptus, crape myrtles, ...

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 ...

(

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 ...

Miconia ''Miconia'' is a genus of flowering plants in the glory bush family, Melastomataceae, native to warm temperate to tropical regions of the Americas. The species are mostly shrubs and small to medium-sized trees up to 15 m tall. The generic name ...

'', ''

Conostegia ''Miconia'' is a genus of flowering plants in the Tibouchina, glory bush family, Melastomataceae, native to warm temperate to Tropics, tropical regions of the Americas. The species are mostly shrubs and small to medium-sized trees up to 15 m ta ...

'', and ''

Ossaea ''Ossaea'' is a genus of flowering plants in the family Melastomataceae. There are about 90 species distributed from Mexico to South America and the Caribbean The Caribbean ( , ; ; ; ) is a region in the middle of the Americas centered arou ...

''), the eurosid I

Malpighiales The Malpighiales comprise one of the largest Order (biology), orders of flowering plants. The order is very diverse, with well-known members including willows, Viola (plant), violets, aspens and Populus, poplars, Euphorbia pulcherrima, poinsett ...

families Family (from ) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictability, structure, and safety as ...

Euphorbiaceae Euphorbiaceae (), the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of Euphorbia, the type genus of the family. Most spurges, such as ''Euphorbia paralias'', ar ...

('' Croton''), and

Malpighiaceae Malpighiaceae is a family of flowering plants in the order Malpighiales. It comprises about 73 genera and 1315 species, all of which are native to the tropics and subtropics. About 80% of the genera and 90% of the species occur in the New World ( ...

(''

Byrsonima ''Byrsonima'' is one of about 75 genera in the Malpighiaceae, a family of flowering plants in the order Malpighiales. In particular in American English, they are known as locustberries (which also may refer specifically to '' Byrsonima coccolobif ...

''), the eurosid II

orders Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * H ...

Sapindales Sapindales is an order of flowering plants. Well-known members of Sapindales include citrus; maples, horse-chestnuts, lychees and rambutans; mangos and cashews; frankincense and myrrh; mahogany and neem. The APG III system of 2009 includ ...

Rutaceae The Rutaceae () is a family (biology), family, commonly known as the rueRUTACEAE
in Bo ...

(''

Zanthoxylum ''Zanthoxylum'' is a genus of about 250 species of deciduous and evergreen trees, shrubs and climbers in the Family (biology), family Rutaceae that are native to warm temperate and Subtropics, subtropical areas worldwide. It is the type gen ...

'') and more commonly

Malvales The Malvales are an Order (biology), order of flowering plants. As circumscribed by Angiosperm Phylogeny Group, APG II-system, the order includes about 6000 species within nine Family (biology), families. The order is placed in the eurosids II, w ...

Malvaceae Malvaceae (), or the mallows, is a family of flowering plants estimated to contain 244 genera with 4225 known species. Well-known members of economic importance include Theobroma cacao, cacao, Cola (plant), cola, cotton, okra, Hibiscus sabdariffa, ...

tribes 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 ...

Bombacoideae Bombacoideae is a subfamily of the mallow family, Malvaceae. It contains herbaceous and woody plants. Their leaves are alternate, commonly palmately lobed, with small and caducous stipules. The flowers are hermaphroditic and actinomorphic; the c ...

(''

Ochroma ''Ochroma pyramidale'', commonly known as balsa, is a large, fast-growing tree native to the Americas. It is the sole member of the genus ''Ochroma'', and is classified in the subfamily Bombacoideae of the mallow family Malvaceae. The tree is fa ...

''),

Malvoideae Malvoideae is a botanical name at the rank of subfamily, which includes in the minimum the genus ''Malva''. It was first used by Burnett in 1835, but was not much used until recently, where, within the framework of the APG System, which unites th ...

(''

Hampea ''Hampea'' is a genus of flowering plants in the family (biology), family Malvaceae. They are trees native to Mexico, Central America, and Colombia. There are about 21 species. Species include: * ''Hampea appendiculata'' * ''Hampea breedlovei'' ...

'' and also ''

Hibiscus ''Hibiscus'' is a genus of flowering plants in the Malva, mallow family, Malvaceae. The genus is quite large, comprising List of Hibiscus species, several hundred species that are Native plant, native to warm temperate, Subtropics, subtropical ...

'',

Byttnerioideae Byttnerioideae is a Family (biology), subfamily of the flowering plant family Malvaceae. ''Byttneria'' is the type genus for the subfamily. ''Byttneria'' is now treated as synonym of ''Ayenia''. Tribes and genera Four tribes are recognised by t ...

(''Byttneria aculeata'', ''

Theobroma ''Theobroma'' is a genus of flowering plants in the mallow family, Malvaceae. It was previously classified as a member of Sterculiaceae, which has been incorporated into Malvaceae to make it monophyletic. It contains roughly 20 species of smal ...

'') and

Grewioideae Grewioideae is a subfamily of the family Malvaceae and was first described by Hochreutiner. The group is named after its type genus, ''Grewia'', which is named for the English scientist Nehemiah Grew (1641–1712). It contains a number of gener ...

(''

Luehea ''Luehea'' is a genus of trees in the family Malvaceae. Its native range stretches from Mexico to southern tropical America and Cuba. It is native to the countries of; Argentina, Belize, Bolivia, Brazil, Colombia, Costa Rica, Cuba, Ecuador, El S ...

''). The "green lizard caterpillar" ''Macrosoma tipulata'' attacks an economically important local fruit tree "Cupuaçu" (''

Theobroma grandiflorum ''Theobroma grandiflorum'', commonly known as cupuaçu, also spelled cupuassu, cupuazú, cupu assu, or copoazu, is a tropical rainforest tree related to cacao. Native and common throughout the Amazon basin, it is naturally cultivated in the jung ...

'') in

and can defoliate saplings; the biology of this species has been studied and illustrated in some detail. The larva of this species lives about 15 days in 5 instars, the pupal stage lasts about 7 days and the adult lives about 10 days. ''M. tipulata'' and many other species can be found as adults through most of the year.

DNA sequences

A few species have been

sequenced In genetics and biochemistry, sequencing means to determine the primary structure (sometimes incorrectly called the primary sequence) of an unbranched biopolymer. Sequencing results in a symbolic linear depiction known as a sequence which succi ...

for the

mitochondrial A mitochondrion () is an organelle found in the cells of most eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...

genes "cytochrome oxidase I", and "ND1" and

nuclear gene A nuclear gene is a gene whose DNA sequence is located within the cell nucleus of a eukaryotic organism. These genes are distinguished from extranuclear genes, such as those found in the genomes of mitochondria and chloroplasts, which reside ou ...

s "Wingless" and "Ef-1", including ''Macrosoma semiermis''. Some species are currently being barcoded.

Cited literature

Sources

External links