''Theritas'' is a

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

of gossamer-winged butterflies (

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 ...

Lycaenidae Lycaenidae is the second-largest family (biology), family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide, whose members are also called gossamer-winged butterflies. They constitute about 30% of ...

) found in the

Neotropics 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 biogeog ...

. Among the

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 ...

Eumaeini The Eumaeini are a tribe of gossamer-winged butterflies (family Lycaenidae). They are typically placed in the subfamily Theclinae, but sometimes considered a separate subfamily Eumaeinae. Over 1,000 species are found in the Neotropical realm S ...

of its

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 ...

Theclinae The subfamily Theclinae is a group of butterflies, often referred to as hairstreaks, with some species instead known as elfins or by other names. The group is part of the family Lycaenidae, the "gossamer-winged butterflies". There are many tropi ...

, it is usually placed in the group around the genus '' Atlides''. In particular, it seems most closely related to ''

Arcas In Greek mythology, Arcas (; Ancient Greek: Ἀρκάς) was a hunter who became king of Arcadia. He was remembered for having taught people the arts of weaving and baking bread and for spreading agriculture to Arcadia. Family Arcas was the ...

''.Bálint ''et al.'' (2007), Brower (2007)

Description

''Theritas'' species are large among the

but altogether still smallish butterflies. They typically have a pennant-like hindwing "tail" which projects outwards at an approximate

right angle In geometry and trigonometry, a right angle is an angle of exactly 90 Degree (angle), degrees or radians corresponding to a quarter turn (geometry), turn. If a Line (mathematics)#Ray, ray is placed so that its endpoint is on a line and the ad ...

from the squared-off edge of the tapered tornal area. A smaller tail is located slightly more forward on the hindwing. The upperwings of these butterflies are largely or entirely colored brilliant blue. But these traits – as opposed to the androconia pouches discussed below – are also found in several other Eumaeini, which have been included in this genus but do not really appear to belong here. Like ''

'', they have

androconia The external morphology of Lepidoptera is the physiological structure of the bodies of insects belonging to the order Lepidoptera, also known as butterflies and moths. Lepidoptera are distinguished from other orders by the presence ...

(scent scales) on the hindwing upperside near the base of the cubital vein; these are

autapomorph In phylogenetics, an autapomorphy is a distinctive feature, known as a derived trait, that is unique to a given taxon. That is, it is found only in one taxon, but not found in any others or outgroup taxa, not even those most closely related to ...

ically tucked away in a flat deep pouch in space 1b between the wing veins. Some species also have androconia on the forewings, but it needs to be clarified whether these are indeed ''Theritas

sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...

''. The hindwing underside is brownish in these butterflies, with a subbasal line running between the subcosta and the anal vein branch of the cubital vein. This genus has some

sexual dichromatism Sexual dimorphism is the condition where sexes of the same species exhibit different morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecious species, which consis ...

in that the males show green

iridescence Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear gradually to change colour as the angle of view or the angle of illumination changes. Iridescence is caused by wave interference of light in microstru ...

of the blue upperwing color. By comparison, ''Arcas'' has golden-green hindwing undersides dusted with black, with a black band running roughly parallel to the distal wing margin, longer tails, and lacks sexual dichromatism. The female genitals have a long, straight and tubular genital duct with a plain anal end in the type species '' T. mavors'', but variation within the genus is insufficiently studied.Bálint ''et al.'' (2006, 2007)

Species

There have been 2 major competing

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 ...

schemes for this genus in the late 20th century. The traditional view delimits the genus loosely, with about 30 species. A more recent proposal restricts the genus to 6-8 valid species, which all possess the typical

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 and can be divided into three species groups or

superspecies In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...

:Bálint ''et al.'' (2007) ''mavors'' group * '' Theritas kalikimaka'' (Clench, 1944) (sometimes included in ''T. mavors'') * '' Theritas lotis'' (Goodson, 1945) (sometimes included in ''T. mavors'') * '' Theritas mavors'' Hübner, 1818 ''paupera'' group * '' Theritas drucei'' (Lathy, 1926) * '' Theritas gozmanyi'' (Bálint, Wojtusiak, Kerstész & Biró, 2007) * '' Theritas harrietta'' (Weeks, 1901) * '' Theritas paupera'' (C. Felder & R. Felder, 1865) ''triquetra'' group * '' Theritas triquetra'' (Hewitson, 1865) In 2019, phylogenetic research (Martins 2019) was published that reclassified ''Theritas'' as a small, widespread Neotropical genus containing these five species: * '' Theritas mavors'' Hübner, 1818 - Mexico to central Brazil * '' Theritas triquetra'' (Hewitson, 1865) - Brazil Atlantic region * '' Theritas paupera'' (C. Felder & R. Felder, 1865) - Venezuela to southern Peru * '' Theritas harrietta'' (A.G. Weeks, 1901) - southern Peru to Bolivia * '' Theritas drucei'' (Lathy, 1926) - Brazil This research included seven genera of the Atlides Section of the tribe Eumaeini: ''

'', '' Atlides'', ''

Brangas ''Brangas'' is a genus of hairstreaks in the butterfly family Lycaenidae, found in Mexico, Central America, and South America. There are about 16 described species in ''Brangas''. Species These species belong to the genus ''Brangas'': * '' Bran ...

'', '' Denivia'', '' Lucilda'', '' Pseudolycaena'', and ''Theritas''. Additionally, the genera ''Margaritheclus'' and ''Dabreras'' were classified as synonyms of ''Lucilda'' and ''Brangas''.

Provisionally placed here

A number of species are usually placed in ''Theritas'', but apparently will have to be split off if the genus is to be a

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 ...

group. '' "T." anna'' for example might be closer to ''

'' and '' Evenus'' or even belong in the latter, while '' "T." deniva'' and some allies could just as well be closer to '' Micandra'' and ''

Penaincisalia ''Penaincisalia'' is a genus of butterflies in the family Lycaenidae. It contains the following species, all found in the Neotropical realm: partial list * ''Penaincisalia alatus'' Druce 1907 * ''Penaincisalia anosma'' Draudt 1921 * ''Penaincisa ...

'' than to ''Theritas'' proper. All in all, it is not unlikely that some of the genera nowadays considered

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. ...

s of ''Theritas'' turn out to be valid. Several of the proposed genera are

monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...

, and thus perhaps least likely to be valid. One of these has been argued to be ''

nomen nudum In Taxonomy (biology), taxonomy, a ''nomen nudum'' ('naked name'; plural ''nomina nuda'') is a designation which looks exactly like a scientific name of an organism, and may have originally been intended to be one, but it has not been published ...

'' per Article 13.1 of the ICZN Code, because an appropriate genus description was not given; the matter has been submitted to the

ICZN 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 ...

for discussion. That all nonwithstanding, species provisionally placed in ''Theritas'' are: * '' Theritas acontius'' (Goodson, 1945) * '' Theritas adamsi'' (Druce, 1909) (close to ''"T." deniva''?) * '' Theritas anna'' (Druce, 1907) (

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 ...

of ''Aveexcrenota''; including ''A. mimianna'') * '' Theritas arene'' (Goodson, 1945) * '' Theritas augustinula'' (Goodson, 1845) * '' Theritas augustula'' (Kirby, 1877) * '' Theritas boliboyeri'' (Bálint & Wojtusiak, 2006) (later ''Lucilda boliboyerus'' (Bálint & Wojtusiak, 2006) per Martins et al. 2019. * '' Theritas chalumna'' (Schaus, 1902) * '' Theritas crines'' (Druce, 1907) (type species of ''"Lucilda"'') * '' Theritas curitibaensis'' (Johnson, 1992) * '' Theritas danaus'' (C. & R.Felder,

865 __NOTOC__ Year 865 ( DCCCLXV) was a common year starting on Monday of the Julian calendar. Events By place Europe * King Louis the German divides the East Frankish Kingdom among his three sons. Carloman receives Bavaria (with more l ...

(type species of ''Margaritheclus'') * '' Theritas deniva'' (Hewitson, 1874) (type species of ''Denivia'') * '' Theritas hemon'' (Cramer,

775 __NOTOC__ Year 775 ( DCCLXXV) was a common year starting on Sunday of the Julian calendar. The denomination 775 for this year has been used since the early medieval period, when the Anno Domini calendar era became the prevalent method in Europe ...

(close to ''"T." deniva''?) * '' Theritas lisus'' (Stoll, 1790) * '' Theritas margaritacea'' (Draudt, 1919) * '' Theritas monica'' (Hewitson, 1867) * '' Theritas phegeus'' (Hewitson, 1865) * '' Theritas theocritus'' (Fabricius, 1793) * '' Theritas viresco'' (Druce, 1907) In addition, there are at least three more undescribed species placed here: one from

Santa Catarina state Santa Catarina () is one of the 27 federative units of Brazil. It is located in the centre of the country's Southern region. It is bordered to the north by the state of Paraná, to the south by the state of Rio Grande do Sul, to the east by t ...

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 ...

and two from the

Andes The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the List of longest mountain chains on Earth, longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range ...

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 ...

. It is unknown whether they are indeed ''Theritas'' ''

''. ''T. drucei'' and ''T. triquetra'' as well as some species of the disputed assemblage are known from Santa Catarina or its surroundings, while the rest of the ''paupera'' group but also many other related

are found in the Andes.Bálint ''et al.'' (2007), and see references in Savela (2011)

Footnotes

References

* (2006): Comments on the systematics and natural history of ''Aveexcrenota'', a genus of rare Andean eumaeine Lycaenidae (Lepidoptera). '' Acta Zool. Acad. Sci. Hung.'' 52(3): 331–352
PDF fulltext
* (2007): The description of ''Theritas gozmanyi'' from the Andes and its spectroscopic characterization with some notes on the genus (Lepidoptera: Lycaenidae: Eumaeini). '' Acta Zool. Acad. Sci. Hung.'' 53(Supplement 1): 211–224
PDF fulltext
* (2008): Tree of Life Web Project &ndash
''Theritas''
Version of 2008-MAR-24. Retrieved 2009-FEB-18. * (2011): Markku Savela's Lepidoptera and some other life forms &ndash

Version of December 24, 2011. Retrieved April 1, 2012. {{Taxonbar, from=Q7783003 Eumaeini Lycaenidae of South America Lycaenidae genera Taxa named by Jacob Hübner