Phylogenetic Classification of the Atlides Section of the Eumaeini (Lepidoptera, Lycaenidae)
Author
Martins, Ananda Regina P.
Author
Duarte, Marcelo
Author
Robbins, Robert K.
text
Zootaxa
2019
2019-02-28
4563
1
119
134
journal article
28153
10.11646/zootaxa.4563.1.6
0d849a4d-5dd1-4f60-9036-939f4486fe72
1175-5326
2601017
8E2935BD-3608-4BE1-B565-AE50966EE510
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.
FIGURE 20.
Scent patch of
Denivia hemon
scent patch. Note tan colored androconia mixed with iridescent blue wing scales.
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.