Denivia K. Johnson, 1992

Martins, Ananda Regina P., Duarte, Marcelo & Robbins, Robert K., 2019, Phylogenetic Classification of the Atlides Section of the Eumaeini (Lepidoptera, Lycaenidae), Zootaxa 4563 (1), pp. 119-134 : 130-133

Denivia K. Johnson, 1992

type species: Thecla deniva Hewitson

Denivia consists of fifteen Neotropical species and one species provisionally placed here. Some lowland species are common in museum collections, such as D. hemon ( Fig. 20 ), and some are rare, such as D. acontius . Similarly, some montane species are common in museum collections, such as D. monica , and some are relatively rare, such as D. curitabaensis . Denivia lisus and D. hemon are widespread, occurring from Mexico to southern Brazil. Within Denivia , the female of D. hemon is the only one that is entirely brown dorsally and ventrally, so that it resembles females of Theritas .

Denivia in the results is characterized by a membranous area in the female ductus bursae ( Table 1, Figs. 18– 19). Although the occurrence of this character is homoplastic, Denivia is monophyletic in all analyses. There is no evident way to divide the genus into better-supported genera. The original description of Denivia ( Johnson 1992) included species that are currently placed in Denivia, Lucilda , and Atlides , but most of the species listed below have not previously been placed in Denivia .

The most conspicuous trait of Denivia is the dorsal forewing scent patch in which neutral-colored androconia are intermixed with iridescent blue wing scales ( Fig. 20 ). According to Martins et al. (2018) ( Fig. 8), this trait evolved within Denivia and was lost evolutionarily once.

Denivia deniva (Hewitson, 1874) (Thecla) , type locality: Brazil

maggae K. Johnson, 1992, type locality: Argentina

grava Bálint, K. Johnson & Kroenlein, 1998 , type locality: Brazil (SP), new synonym

Distribution and Habitat. Argentina and Southern Brazil from MG and ES to SC in lowland and lower montane wet forest.

Synonymy. Denivia grava was mistakenly placed as a synonym (Robbins 2004) of the following species. The jagged hindwing outer margin of the holotype is the reason for its placement here.

Denivia curitabaensis ( K. Johnson, 1992) (Radissima) , type locality: Brazil (PR), New Combination Distribution and Habitat. We have seen three museum specimens of this species from the plateau in central PR and the mountains of SP. It is recorded further south in RS ( Giovenardi et al. 2008; Iserhard et al. 2010),

but we have not seen the specimens on which these records are based. Denivia curitabaensis is a poorly known species that may be easily confused with D. deniva . It differs in lacking the jagged hindwing outer margin of D. denivia .

Denivia augustinula (Goodson, 1945) (Thecla) , type locality: Costa Rica, New Combination

augustinula (Strand, 1916) (Thecla) , type locality: Costa Rica

described as Thecla augustula ab.? augustinula .

Distribution and Habitat. Mexico to northern Colombia in wet lower montane forest, but at lower elevations to the north.

Denivia theocritus (Fabricius, 1793) (Hesperia) , type locality: "Indiis", an incorrect type locality, New Combination

nepia ( Godman & Salvin, 1887) ( Thecla ), type locality: Guatemala; Nicaragua; Panama

Distribution and Habitat. Mexico to western Ecuador in lowland forest. Specimens from eastern Colombia and Ecuador have larger spots on the ventral wings.

Denivia augustula (W.F. Kirby, 1877) (Thecla) , type locality: Costa Rica, repl. name, New Combination augustinus (Butler & H. Druce, 1872) ( Mithras ), type locality: Costa Rica, preocc. (not Westwood, 1852) (ICZN Art. 59.3)

Distribution and Habitat. Costa Rica and Panama to western Ecuador in lower montane forest.

Denivia arene (Goodson, 1945) (Thecla) , type locality: Peru, New Combination

Distribution and Habitat. Ecuador and Peru in montane forest.

Denivia monica (Hewitson, 1867) (Thecla) , type locality: Venezuela, New Combination

crinella (Strand, 1918) ( Thecla ), type locality: Ecuador

described as Thecla crines var.? crinella.

electryon (Goodson, 1945) ( Thecla ), type locality: Ecuador; Peru; Colombia

Distribution and Habitat. Venezuela to Bolivia in lower montane and montane forest.

Denivia hemon (Cramer, 1775) (Papilio) , type locality: Surinam, New Combination

acmon (Cramer, 1775) ( Papilio ), type locality: Surinam

aemon (Cramer, 1776) ( Papilio ), type locality: Surinam, missp.

gispa (Hewitson, 1865) ( Thecla ), type locality: Brazil ([PA])

callirrhoe (Goodson, 1945) ( Thecla ), type locality:?

Distribution and Habitat. Everywhere in Neotropical continental forests below 1,000 m elevation.

Denivia phegeus (Hewitson, 1865) (Thecla) , type locality: Brazil (BA); Amazon, New Combination

laudonia (Hewitson, 1867) ( Thecla ), type locality: Brazil

Distribution and Habitat. The Amazonian Region to southern Brazil (RJ) in wet lowland forest.

Denivia acontius (Goodson, 1945) (Thecla) , type locality: Colombia, New Combination

Distribution and Habitat. Panama south to the Amazonian Region in wet lowland forest.

Denivia chaluma (Schaus, 1902) (Thecla) , type locality: Brazil (SC), New Combination

hamila (Jones, 1912) ( Thecla ), type locality: Brazil (PR)

secunda Bálint, K. Johnson & Kroenlein, 1998, type locality: Brazil (SP)

Distribution and Habitat. Central and southern Brazil from DF and ES to SC in lower montane and montane forest.

Denivia viresco ( H.H. Druce, 1907) (Thecla) , type locality: Brazil (MA), New Combination

photeinos ( H.H. Druce, 1907) ( Thecla ), type locality: Guyana

Distribution and Habitat. Amazonian Region in wet and dry lowland forest.

Denivia silma ( Martins, Faynel, & Robbins, 2016) ( Theritas ), type locality: French Guiana, New Combination Distribution and Habitat. Amazonian Region south to Paraguay and Brazil (MG) in wet low land forest. Denivia espiritosanto Bálint & Moser, 2007 , type locality: Brazil (RJ)

Distribution and Habitat. Eastern Brazil from RJ to SC in wet lowland forest up to 800 m elevation. Denivia lisus (Stoll, 1790) (Papilio) , type locality: Surinam, New Combination

orsina (Hewitson, 1877) ( Thecla ), type locality: Bolivia

hisbon ( Godman & Salvin, 1887) ( Thecla ), type locality: Guatemala; Costa Rica; Panama

Distribution and Habitat. Mexico to southern Brazil in wet and dry lowland forest, occasionally in lower montane forest.

Denivia adamsi (H.H. Druce, 1909) (Thecla) (generic placement uncertain), type locality: Peru

Distribution and Habitat. Presumably lower montane or montane forest.

Generic Placement. We were unable to examine the male holotype of Thecla adamsi Druce for this project. We know of no other male museum specimen. Based on its ventral wing pattern, this species is tentatively placed in Denivia so that it has a generic name.

Excluded from the Atlides Section. Thecla anna Druce “belongs to a group by itself” ( Druce 1907: 39) and has been described as “an entirely isolated animal” ( Draudt 1919 –1920: 750). Robbins (2004b) provisionally placed it in Theritas based on superficial similarities in brush organ structure. Balint et al. (2006) removed it from the Atlides Section because it lacked a hindwing anal cleft. This species did not cluster with the Atlides Section in an exemplar phylogeny based on DNA sequences (Quental, unpublished dissertation). There is no compelling evidence of which we are aware that it belongs to the Atlides Section.

Figures

FIGURE 20. Scent patch of Denivia hemon scent patch. Note tan colored androconia mixed with iridescent blue wing scales.