Evolutionary gain of male secondary sexual structures

Transcription

Insect Systematics & Evolution 46 (2015) 47–78
brill.com/ise
Evolutionary gain of male secondary sexual structures in the
widespread Neotropical montane genus Lathecla (Lepidoptera,
Lycaenidae, Eumaeini)
Robert K. Robbinsa,* and Robert C. Busbyb
Department of Entomology, PO Box 37012, NHB Stop 105, Smithsonian Institution,
Washington, DC 20013-7012, USA
b
7 Countryside Way, Andover, MA 01810-6041 USA
*Corresponding author, e-mail: [email protected]
Published online 20 March 2014; published in print 6 February 2015
a
Abstract
Lathecla is a widespread, primarily montane, Neotropical genus. It consists of seven species that have a
relatively uniform adult wing pattern coupled with a diverse set of male secondary sexual structures.
Taxonomically, we describe five species — Lathecla mitzi Robbins new species, Lathecla fernandezi
Robbins & Busby new species, Lathecla vichai Robbins & Busby new species, Lathecla carolyna
Busby new species and Lathecla winnie Robbins & Busby new species and remove one name from
synonymy — Thecla mimula Draudt, revised status. Evidence is presented for transferring Lathecla from
the Thestius Section of the Eumaeini to the Micandra section, next to Podanotum. Topology of an inferred
phylogenetic tree for Lathecla is stable when male secondary sexual characters are omitted or under a
variety of implied weighting options. A scent patch on the cubital vein of the dorsal surface of the forewing is unique to Lathecla and evolved (and was not lost) in the ancestor of a four-species lineage. Its sister
lineage contains two species. A scent patch on the ventral surface of the forewing evolved (and was not
lost) in the ancestor of a six-species lineage in Lathecla. Its sister lineage contains one species. These results,
along with previous data, show the viability for the Eumaeini of the hypothesis that the evolutionary gain
of a male secondary sexual organ increases the rate of species diversification.
Keywords
Andes; species diversification; biogeography
Introduction
The primarily Neotropical Eumaeini (Lycaenidae) and Riodinidae consist of more
than 1000 species each, but are otherwise a study of contrasts (Robbins 2004a), The
Eumaeini tend to have a basic wing pattern and shape (despite obvious exceptions),
which is one reason why virtually all species were placed in the catchall genus Thecla
Fabricius for 175 years (Swainson 1820-1823; Godman & Salvin 1887-1901; Draudt
© Koninklijke Brill NV, Leiden, 2015
DOI 10.1163/1876312X-45032115
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R.K. Robbins and R.C. Busby / Insect Systematics & Evolution 46 (2015) 47–78
1919-1920; D’Abrera 1995), while the Riodinidae exhibit an amazing array of wing
shapes and patterns that have long been partitioned into a multitude of genera (Seitz
1916-1918). Further, males of more than 95% of the Eumaeini have androconia or
abdominal brush organs in contrast to males of only about 25% of Riodinidae (Hall
& Harvey 2002). These contrasts suggest that diversification in each lineage was substantially different.
Sexual selection acting on male secondary sexual organs can be an important factor
in speciation and is often thought to result in rapid species diversification (e.g., Carson
et al. 1979; Lande 1981, 1982; West-Eberhard 1983; Kopp & True 2002; Steelman &
Danley 2003). This idea is traditionally tested by a phylogenetic contrast of sister lineages (e.g., Barraclough et al. 1995; Møller & Cuervo 1998; Arnqvist et al. 2000),
especially when molecular data are lacking. For example, the number of species is
compared between a lineage with evolutionary diversification of male secondary sexual
structures and a sister lineage with evolutionary stasis of these structures.
The high incidence of male secondary sexual structures in the Eumaeini suggests
that sexual selection was an important factor in the diversification of the Eumaeini, but
ironically, the evolutionary “loss” of such traits in the Eumaeini, often in terminal taxa,
occurs with much greater frequency than their evolutionary “gain” (Quental 2008;
Robbins et al. 2012; cf., Wiens 2001 for a more general discussion of this widespread
pattern). In fact, the only documented case of a gain in a male secondary sexual organ
in the Eumaeini is the evolution of comb-like androconia along the posterior edge of
dorsal hindwing vein Cu2 in five species of Timaeta K. Johnson, Kruse & Kroenlein
(Robbins & Busby 2008).
The hairstreak butterfly genus Lathecla Robbins (Lycaenidae, Eumaeini) consists of
seven primarily montane, Neotropical species that are small (forewing length slightly
more than 1 cm) and possess a similar, relatively non-descript wing pattern (Figs 1–7).
In contrast to this phenotypic similarity in wing pattern, three different kinds of scent
patches (a male secondary sexual structure) occur in the genus (Figs 8–10), one of
which is unique to four Lathecla species. Molecular sequences for Lathecla are not yet
available.
The taxonomic purpose of this paper is to revise the species level taxonomy of
Lathecla and to infer phylogenetic relations among the species. Using these results, the
biological purpose is to document and assess the evolutionary gain of male secondary
sexual structures. We also discuss aspects of the biogeography of Lathecla.
Materials and methods
The species level taxonomy of Lathecla is based on an analysis of variation among 121
pinned specimens (some examined as images) from various museum and private collections, as noted below. Morphological variation in structures of the genitalia, wings,
and androconia within species is noted in the species accounts; otherwise the species
recognized were morphologically homogenous in the sample. Allopatric taxa were
considered to be distinct if they had consistently differentiating traits and they occurred
49
Figs 1–7. Adult Lathecla. Male (left, dorsal wing surface on left) and female (right) except for L. mitzi,
for which these males are the only known specimens . (1) L. mitzi holotype male (PR, Brazil), paratype
male (RJ, Brazil); (2) L. latagus male (Panama), female (Mexican holotype of Thecla epopeoides Schaus);
(30 L. mimula male (Ecuador), female (RJ, Brazil); (4) L. fernandezi holotype male (eastern Ecuador),
female (eastern Ecuador); (5) L. vichai holotype male (Panama), female (Panama); (6) L. carolyna holotype
male (western Ecuador), female (western Ecuador); (7) L. winnie holotype male (Costa Rica), female
(Costa Rica). Scale 1.0 cm. This figure is published in colour in the online edition of this journal, which
can be accessed via http://booksandjournals.brillonline.com/content/1876312x.
50
Figs 8–10. Scent patches, Lathecla fernandezi (left) and L. vichai. (8) Ventral forewing scent patch;
(9) dorsal hindwing scent patch; (10) dorsal forewing scent patch on cubital vein. Inset on left higher
magnification. Inset on right, SEM of cubital vein cross section through the scent patch, showing that it
is not a scent pad (see text). This figure is published in colour in the online edition of this journal, which
can be accessed via http://booksandjournals.brillonline.com/content/1876312x.
51
in recognized biogeographic zones for forest butterflies (as outlined in Brown 1982). A
nomenclatural list and male identification key are presented. Species accounts include
notes when relevant on nomenclature, history, morphological variation, elevation, seasonality and behavior. Males and females were associated by similarity of ventral wing
pattern and distribution. Geographic distribution of each species is mapped. Labels on
holotypes are recorded verbatim with brackets used for information not explicitly
noted on the labels and for descriptions of the labels. Otherwise, months are abbreviated by their first three letters in English. We denote Brazilian states by their two letter
postal codes. Paratype labels are light blue. Citations for original descriptions can be
found in Lamas et al. (1995). Biogeographic zones follow Brown (1982), who partitioned the montane forested continental Neotropics into the Transandean Region,
Andean Region and Atlantic Region. Following Elias et al. (2009) for the nymphalid
genus Ithomia, we classify Lathecla species as low-to-mid elevation (0–1500 m) and
mid-to-high elevation (> 1000 m). Many male eumaeines perch in “territories” at certain times of day to wait for receptive females to fly through and “defend” these areas
by flying at other males that enter the territory (e.g., Powell 1968; Alcock & O’Neill
1987; Cordero et al. 2000). Recorded times from our fieldwork for “territorial” behavior are standard time at that locality. Finally, traps baited with decaying fish attract
some lycaenid species and not others. We note species and sex for species collected
using fish-baited traps.
We used adult morphological characters for the phylogenetic analyses because DNA
sequences and immature stages are currently unavailable for any Lathecla species.
Standard references for morphological terminology are Comstock (1918) for insect
wing veins; Klots (1970) and Robbins (1991) for lepidopteran genitalia as modified for
the Eumaeini; Robbins (1991) and Robbins et al. (2012) for male secondary sexual
structures; and Snodgrass (1935) for all other morphological structures. Methods of
preparation for genitalic dissections and for examination of male wing secondary sexual characters with an SEM follow Robbins (1991).
Seven Lathecla species are recognized (Table 1). Six were previously listed (Robbins
2004b), but based on results in this paper, one name is removed from synonymy. The
first pair of outgroups for the phylogenetic analyses are Thestius epopea (Hewitson) and
T. azaria (Hewitson). The male and female genitalia show an overall similarity with
those of Lathecla, for which reason Thestius Hübner was placed next to Lathecla in a
checklist (Robbins 2004b). The second pair of outgroups are Podanotum salaeides
(Draudt) and a species of Podanotum for which we have associated males and females,
but are uncertain of its specific identity. Evidence is given in this paper that Podanotum
Torres & K. Johnson is closely related to Lathecla. Most taxonomic papers on Podanotum
(e.g., Bálint 2001; Bálint & Wojtusiak 2002; Bálint & Attal 2007) have contained
“minimal” species descriptions in which variation — geographical, seasonal and
elevational — was not assessed. As a result, the application of most available names
in Podanotum is yet premature. The third pair of outgroups are Rhamma commodus
(C. Felder & R. Felder) and R. bilix (Draudt). Podanotum and Rhamma K. Johnson
were sister genera in a phylogenetic analysis of DNA sequences (Quental 2008).
No other eumaeine genera have been proposed to be close relatives of Lathecla.
52
Table 1. Character matrix for Lathecla.
Taxa
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Thestius azaria outgroup
Thestius epopea outgroup
Rhamma commodus outgroup
Rhamma bilix outgroup
Podanotum salaeides outgroup
Podanotum C outgroup
Lathecla mitzi
Lathecla latagus
Lathecla mimula
Lathecla fernandezi
Lathecla vichai
Lathecla carolyna
Lathecla winnie
0
0
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
1
2
2
2
2
2
2
0
0
0
0
0
0
0
0
0
1
1
1
1
0
0
0
0
0
0
0
2
2
1
1
1
1
0
0
0
0
0
0
0
1
1
1
1
1
1
0
0
1
1
0
0
0
0
0
0
0
0
0
0 0
0 0
0 0
0 0
0 0
0 0
? 1
0 1
0/11
0 1
0 1
0 1
0 1
–
–
–
–
–
–
0
1
1
2
0
0
3
–
–
–
–
–
–
0
1
1
2
3
3
3
0
0
0
0
0
0
0
1
1
1
2
2
2
0
0
0
0
0
0
1
1
1
2
1
3
3
1
1
1
1
1
1
1
1
1
2
0
2
2
0
0
0
0
0
0
1
1
1
1
1
1
1
0
0
0
0
0
0
1
1
1
1
1
1
1
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
1
1
0
0
0
0
0
0
0
0
0
0
1
1
1
2
2
3
3
3
3
3
3
3
0
0
0
1
1
1
?
1
1
1
1
0
0
0
0
1
1
0
0
?
0
0
0
0
0
0
Character states for each Lathecla and outgroup species were recorded (Table 2), and
we searched exhaustively for shortest trees using the implicit enumeration option of
TNT software (Goloboff et al. 2008) to derive a most parsimonious cladogram with
the collapse trees after the search option. To test the assumption of equally weighted
characters, implied weighting was performed over a range of values for the parameter
K (1, 10, 50, 250 and 1000). Mapping of characters and biogeographic regions on
trees was done with WinClada software (Nixon 2002) with the unambiguous changes
only options. Bootstrap support was determined in WinClada (1000 replications
with mult*10, memory 1000 trees). Because we are interested in the evolution of
scent patches, we repeated the phylogenetic analysis with the scent patch characters
omitted.
Specimens cited in this study are deposited in the following collections (abbreviations where available are from Evenhuis (2013)).
BMNH
CF
CMNH
DZUP
FSMC
MacD
MC
RCB
SMF
The Natural History Museum (formerly British Museum (Natural
History)), London, UK
Private collection of Christophe Faynel, Montpellier, France
Carnegie Museum of Natural History, Pittsburgh, PA, USA
Museu de Entomología Pe. Jesus Santiago Moure, Universidade Federal
do Paraná, Curitiba, Paraná, Brazil
Florida Museum of Natural History, University of Florida, Gainesville,
FL, USA
Private collection of John R. & Steven L. MacDonald, Starkville, MS,
USA
Private collection of Alfred Moser, São Leopoldo, Rio Grande do Sul,
Brazil
Private collection of Robert C. Busby, Andover, MA, USA
Forschungsinstitut und Naturmuseum Senckenberg Frankfurt,
Frankfurt-am-Main, Germany
53
Table 2. Characters used in the phylogenetic analysis.
1. VFW postmedian line terminates anteriorly: (0) at inner margin (1) at vein 2A (Figs 1–7).
2. Basal part of DFW cubital vein: (0) no swelling (1) slight swelling (2) conspicuously swollen (Fig. 10).
3. Androconia surrounding DFW cubital vein near the middle of the cell: (0) absent (1) present
(Fig. 10).
4. Inner margin of male forewing (Figs. 1-7): (0) relatively straight (1) slightly convex (2) strongly
convex.
5. Inner margin of VFW: (0) without scent patch (1) with scent patch (Fig. 8).
6. Dorsal forewing scent pad: (0) absent (1) present (illustrated in Johnson 1992).
7. Female dorsal wing color (Figs 1–7): (0) with blue iridescence (1) brown.
8. Scent patch on costa of DHW: (0) absent (1) present (Fig. 9).
9. Color of DHW androconia (Fig. 9): (0) blue (1) black (2) off-white (3) black intermixed with gray
scales.
10. Location and size of DHW scent patch (Fig. 9): (0) large patch from Sc + R1 to Rs (1) large scent
patch from Rs to M1 (2) small patch touching vein Sc + r1 (3) small patch touching vein Rs.
11. Curvature of HW coastal margin (Figs 1–7): (0) slightly convex (1) gradually convex (2) highly
convex.
12. Shape of valvae (Figs 13–19): (0) not basally lobed (1) basally lobed with long needle-like tips
(2) basally lobed with short needle-like tips (3) basally lobed with tip not needle-like.
13. Length of valvae (Figs 13–19): (0) tips extend to base of gnathos (1) tips distal of base of gnathos but
anterior of labides (2) tips extend to the labides
14. Gnathos tips (Figs 13–19): (0) with a short abrupt hook (0.25 mm or less) (1) long apiculus (about
0.5 mm).
15. Dorsal posterior end of penis (Figs 13–19): (0) cylindrical more or less (1) well anterior of the
ventral tip.
16. Posterior penis with a serrate ventral keel: (0) absent (1) present (illustrated in Torres et al. 1996).
17. Process on the ventro-lateral vinculum/tegumen: (0) absent (1) present (illustrated in Johnson 1992).
18. Male genitalia cornutus in vesica of penis: (0) single posteriorly rounded with teeth (1) double
(2) none (3) single posteriorly slim not rounded (Figs 13–19).
19. Sclerotized “cross bar” on the anterior part of the female genitalia signa: (0) absent (1) present
(Figs 20–23).
20. Ductus bursae in dorsal or ventral aspect: (0) evenly sclerotized (Figs 20–25) (1) lightly sclerotized in
the middle so that it appears to be transparent (illustrated in Johnson 1992).
USNM
ZMHB
National Museum of Natural History, Smithsonian Institution,
Washington, DC, USA
Museum für Naturkunde der Humboldt-Universität, Berlin, Germany
Systematics
Nomenclatural checklist
LATHECLA Robbins, 2004, type species: Thecla latagus Godman & Salvin,
designated in the original description (Robbins & Duarte 2004).
Lathecla mitzi Robbins, type locality: Brazil (Parana) new species
Lathecla latagus (Godman & Salvin, 1887) (Thecla), type locality: Panama
epopeoides (Schaus, 1902) (Thecla), type locality: Mexico (Veracruz)
Lathecla mimula (Draudt, 1920) (Thecla), type locality: Peru revised status
Lathecla fernandezi Robbins & Busby, type locality: Ecuador new species
54
Fig. 11. Most parsimonious cladogram for Lathecla with unambiguous character state changes (34 steps,
CI = 0.91, RI = 0.94). Hollow circles are homoplastic changes. Numbers to right of nodes are bootstrap
values. Nodes A, B, and C denote where scent pads evolved on the dorsal hindwing, ventral forewing, and
dorsal forewing, respectively. See text for further explanation.
Lathecla vichai Robbins & Busby, type locality: Panama new species
Lathecla carolyna Busby, type locality: Ecuador new species
Lathecla winnie Robbins & Busby, type locality: Costa Rica new species
Key to male Lathecla based on wing characters (Figs 1–10)
1. Anterior portion of dorsal hindwing with black androconial patch�� 2
Anterior portion of dorsal hindwing without black androconial patch �� 3
2. Bright blue dorsal color with black FW outer margin �� L. mimula
Deep blue/purple dorsal color with black FW outer margin �� L. latagus
3. Anterior portion of dorsal hindwing with small patch of
grey/white androconia �� L. fernandezi
Anterior portion of dorsal hindwing without small patch of
grey/white androconia�� 4
4. Dark androconial patch on the basal side of the VFW inner
margin �� 5
Without dark androconial patch on the basal side of the VFW inner
border �� L. mitzi
55
Fig. 12. Três Picos, Itatiaia National Park, RJ, Brazil. Arrow points to the mountain summit (1650 m)
where the paratype male of L. mitzi was found. This figure is published in colour in the online edition
of this journal, which can be accessed via http://booksandjournals.brillonline.com/content/1876312x.
5. Dorsal hindwing with a thick black border �� 6
Dorsal hindwing with a very thin black border �� L. vichai
6. Blue scales restricted to the posterior half of the dorsal forewing �� L. winnie
Blue scales extend well into the anterior portion of the dorsal
forewing �� L. carolyna
Lathecla Robbins 2004
Diagnosis
Lathecla is a monophyletic genus in the phylogenetic results with a bootstrap support
of 99%. The synapomorphies that characterize Lathecla in the phylogenetic analysis
(unambiguous changes, Fig. 11) are a scent patch on the dorsal surface of the hindwing
near the costa (character 8), male genitalia valvae lobed basally (character 12), male
genitalia gnathos with a long apiculus about 0. 5 mm in length (character 14), dorsal
tip of the penis well anterior of the ventral tip (character 15), and male genitalia penis
with a single, posteriorly blunt cornutus. Characters 14 (Figs 13–19) and 15
(Figs 13–19) are either unique within the Eumaeini or highly unusual, and will be the
56
Figs 13–16. Lathecla male genitalia. Ventral and lateral view of the genital capsule (top, middle) and
lateral view of the penis (bottom). Posterior of insect to the right. (13) L. mitzi; (14) L. latagus; (15) L.
mimula; (16) L. vichai. Scale 1 mm.
57
Figs 17–19. Lathecla male genitalia. Ventral and lateral view of the genital capsule (top, middle) and
lateral view of the penis (bottom). Posterior of insect to the right. (17) L. fernandezi; (18) L. Winnie; (19)
L. carolyna. Scale 1 mm.
58
Figs 20–25. Lathecla female genitalia. Dorsal (top) and lateral view of the ductus copulatrix. Posterior of
insect to the right. (20) L. latagus; (21) L. mimula; (22) L. fernandezi; (23) L. vichai; (24) L. carolyna; (25)
L. winnie. Scale 1 mm.
most useful in determining whether species discovered in the future belong to Lathecla.
Although the gnathos of Lathecla resemble those of Arumecla Robbins & Duarte in the
Calycopidina, the apiculate gnathos tips are 2–3-times longer in Arumecla than in
Lathecla (Duarte & Robbins 2010). A potential distinguishing trait mentioned in the
original description of the genus (Robbins & Duarte 2004) is the convex shape of the
male forewing inner margin (character 4), but according to the phylogenetic analysis,
this trait evolved within the genus. Shape of the posterior vinculum was also used to
distinguish Lathecla (Robbins & Duarte 2004), but this trait appears to be too variable
to be useful for identification.
59
Superficial recognition that a hairstreak butterfly belongs to Lathecla can be
difficult because adults are relatively small (forewing length typically about 1.1 cm),
non-descript montane species that resemble many other small montane forest
eumaeines. The inability to recognize individuals of Lathecla is undoubtedly one reason why the genus has been poorly known. An excellent external distinguishing trait
(character 1) is the point where the postmedian line on the ventral surface of the forewing terminates posteriorly (Robbins & Duarte 2004). It terminates at vein Cu2 in
Lathecla, Rhamma and Podanotum, and at vein 2A or at the inner margin in many
other small primarily montane eumaeine genera, such as Hypostrymon Clench, Dicya
K. Johnson, Nesiostrymon Clench, Timaeta, Temecla Robbins, and Thestius Hübner. All
male Lathecla have a scent patch at the costa on the dorsal surface of the hindwing
(lacking in the outgroups except for some species of Rhamma) and lack a scent pad on
the dorsal surface of the forewing (always present in Rhamma). These external characters distinguish male Lathecla from virtually all other small male eumaeines except for
a few species of Thereus Hübner (distinguishing traits for Thereus in Robbins 1991).
Distribution and habitat (Figs 26–28)
Lathecla inhabits forest from Mexico (Veracruz) to southern Brazil (Paraná). Five species are primarily montane (above 1000 m elevation, infrequent records below this
elevation) while L. latagus and L. mimula occur from sea level to about 1500 m elevation. A majority of Lathecla individuals were found in wet forest.
Scent patches (Figs 8–10)
Scent patches are located on three different parts of the wings in Lathecla. The first
kind of scent patch in Lathecla is located on the ventral surface of the forewing between
the cubital vein and vein 2A (Fig. 8). It occurs in all Lathecla species except L. mitzi
(character 5), and the androconia may be black or off-white. The costa of the hindwing
is enlarged so that most of the scent patch is covered by the hindwings. A superficially
similar scent patch occurs with enlargement of the hindwing costa in a variety of other
eumaeine genera, such as Strephonota K. Johnson, Austin, Le Crom & Salazar and
Janthecla Robbins & Venables, but not in the outgroups. Some Rhamma species on the
ventral surface of the forewing have iridescent blue scales, but such scales do not occur
in Lathecla and do not appear to be androconia.
The second kind of scent patch is located on the dorsal surface of the hindwing,
bordered anteriorly by vein Sc + R1 and posteriorly by vein Rs or M1 (Fig. 9, character
10). It occurs in all Lathecla species (character 8), and the androconia may be blue,
black, or off-white (character 9). The inner margin of the forewing is enlarged (except
in L. mitzi) so that the scent patch is partially covered by the forewings. A similar scent
patch occurs in some Rhamma species, perhaps being most conspicuous in R. comstocki
K. Johnson, and in a variety of other eumaeine genera, such as Thereus Hübner and
Erora Scudder.
The third kind of scent patch in Lathecla is located on the dorsal surface of the forewing on a swollen part of the basal cubital vein (Fig. 10). It occurs in four Lathecla
60
Fig. 26. Geographic distribution of L. mitzi (hearts), L. fernandezi (stars) and L. vichai (diamonds).
species (L. fernandezi, L. vichai, L. carolyna, L. winnie), and in no other eumaeines.
Although it superficially resembles a scent pad (cf., Thomas 1893 for scent pad histology, Robbins 1991; Robbins et al. 2012 for scent pad morphology), it lacks supporting
columns and invaginations containing secretory cells, structures that characterize scent
pads (Fig. 10, insert on right).
61
Fig. 27. Geographic distribution of L. mimula (squares), L. latagus (circles).
Biology
There is a paucity of recorded biological data on Lathecla. Beebe (1949, 1951) documented insects “migrating” in a southward direction against the wind through
62
Fig. 28. Geographic distribution of L. carolyna (hearts), L. winnie (flowers).
Portachuelo Pass (Rancho Grande, 1100 m) in Aragua, northern Venezuela, primarily
at the beginning of the wet season. Two males of L. fernandezi were found in 1985 flying through Portachuelo Pass during the wet season “migration” (vouchers in USNM).
However, L. fernandezi typically occurs in montane wet forest, so the status of this
species as a migrant is unclear. Four Lathecla species have been recorded setting up
63
mating territories. The montane species (L. mitzi, L. fernandezi, L. carolyna) were territorial during the morning while a lowland species (L. mimula) was recorded at 14:00
hours. The known males of L. mitzi were found on mountain tops (Fig. 12). Males and
females of L. mimula were attracted to decaying fish. No larval food plants have been
recorded for Lathecla.
Lathecla mitzi Robbins, new species
Type material
Holotype ♂: (Fig. 1, left). [red printed label] Holotype/Lathecla mitzi/Robbins. [white
printed label] Morro do Anhangava/Quatro Barras/Paraná/Brasil/20-1-2010 Dolibaina
leg [white printed label] DZ 23. 217. Deposited DZUP.
Paratype 1♂: (Fig. 1, right). Brazil, RJ, P. N. de Itatiaia, Três Picos, 22°27’S, 44°36’W,
27 May 1994, 1650m, Leg. Robbins & Caldas. Territorial Behavior Time: 09:46
hours. Deposited USNM.
Etymology
It is with great pleasure that we name this unique and lovely species after Mitzi Caldas,
the mother/mother-in-law of Caldas and Robbins, who discovered this species in
1994. She is a beautiful and wise Carioca who has resided her whole life in southern
Brazil, where this species occurs. The name is a noun in apposition.
Type locality
The holotype was collected by Diego Dolibaina on the summit of Morro do Anhangava
at 1430 m elevation with coordinates 25°23´ S, 49°00´ W. The vegetation at the type
locality was described as “campos rupestres” by Koehler et al. (2002) and Simão (2008).
Morro do Anhangava is part of Parque Estadual Serra da Baitaca.
Diagnosis and description
The male is immediately distinguished from all other male Lathecla by the straight
inner margin of the forewing and the lack of a scent patch on the ventral surface of the
forewing. The female is unknown. According to the phylogenetic results, L. mitzi is the
sister lineage of the remainder of the genus. The wing pattern of the two known males
(Fig. 1) and the male genitalia (Fig. 13) are illustrated. Mean forewing length is 1.2 cm,
SD = 0.14, N = 2.
Variation
The primary difference between the two known males is that the holotype has
fewer blue scales on the dorsal wings, especially the hindwing, than the paratype
(Fig. 1). The significance, if any, of this difference is difficult to assess with only two
known individuals.
64
Habitat, elevation and seasonality
The known habitat of L. mitzi is mountaintops in the midst of relatively wet forest at
1430–1650 m elevation (Fig. 12). An image of the spot where the holotype was collected (figure 13 in Simão 2008) looks very much like the spot where the paratype was
found. Dates of capture are January and the end of May (towards the end of autumn
in southern Brazil), so the flight season of this species encompasses those months of the
year when it is sufficiently warm for activity at these elevations.
Behavior
The holotype, according to its collector, Dolibaina, was flying with males of other
lycaenid species (an indication of territorial behavior) at about 10:00 h. The paratype
male was recorded setting up a mating territory at 09:46 h.
Distribution (Fig. 26)
Lathecla mitzi is known only from the mountaintops in the southern Brazilian states
of Rio de Janeiro and Paraná. It is allopatric with all other Lathecla species. Although
L. mitzi and L. mimula both occur in the states of Rio de Janeiro and Paraná — and
potentially could be sympatric at some localities — the former is unknown under 1430
m elevation while the latter is unknown above 1000 m elevation.
Remarks
This species is rare in collections because, in part, it occurs in high elevation forest that
is frequently not easily accessible. Additionally, males of L. mitzi do not occur consistently at the localities where they are found. For example, Dolibaina found only one
male of L. mitzi in about 10 collecting trips to the type locality. Similarly, after the
paratype male was discovered almost 20 years ago, the senior author returned several
times to the same mountaintop in the morning, but did not find additional males.
Lathecla latagus (Godman & Salvin, 1887)
Nomenclature
Thecla latagus (Godman & Salvin) — the Panamanian male holotype from the
Staudinger collection is in ZMHB — and Thecla epopeoides (Schaus) — the Mexican
female holotype from the Schaus collection is in USNM (Fig. 2) — were synonymized
(Robbins 2004b) because they are the only two lowland species in Central America
with the same ventral wing pattern. The holotype is illustrated online (http://butterfliesofamerica.com/, accessed 25 September 2013).
Lathecla latagus and L. mimula share a sharply convex inner margin of the male forewing and black androconia in the scent patch on the dorsal surface of the hindwing.
65
They are sisters in the phylogenetic results. Although these species were treated as synonyms (Robbins 2004b), three distinguishing traits suggest that they are distinct species. (1) In both sexes of L. latagus, the postmedian line segment in cell M3-Cu1
on the ventral surface of the hindwing is displaced basally (compared to the segment
in cell M2- M3) while these two segments are almost smoothly co-linear in L. mimula.
(2) In males of L. latagus, the distal edge of the postmedian line on the ventral surface
of the forewing is narrow and interrupted while the white is broader and continuous
in L. mimula. (3) In females of L. latagus, the dorsal surface of the wings is bright blue
while it is brown in L. mimula (Figs 2, 3). These characters consistently differentiate
the species except that two (both from southern Brazil) of the 14 females of L. mimula
examined have some blue scaling on the dorsal surface of the wings. The wing pattern (Fig. 2), male genitalia (Fig. 14), and female genitalia (Fig. 20) are illustrated.
Mean male forewing length is 1.1 cm, SD = 0.14, N = 3. Mean female forewing length
is 1.3 cm, SD = 0.07, N = 2.
Habitat and elevation
Lathecla latagus occurs in very wet forest from sea level to 1100 m elevation in Central
America and from 850 to 1500 m elevation in western Ecuador. The three Ecuadorian
specimens from the Nanegal-Garcia Moreno Road (Pichincha Province) were collected
under 1,500 m elevation, even though the data labels were less precise.
Lathecla latagus is a Transandean lowland species (Brown 1982, Robbins et al. 2010)
occurring from Mexico to western Ecuador. It is allopatric with its sister species,
L. mimula. Because of elevational overlap, it is possible that L. latagus will be found to
be sympatric with L. winnie and L. fernandezi.
Material examined (8♂ and 5♀)
Mexico, Veracruz: 3♂ Veracruz, Catemaco, 26 Jul 1986, 27 Jul 1986, 27 Jul 1986
(RCB). 1♀ Coatepec, no date, holotype of Thecla epopeoides Schaus (USNM, Fig. 2,
right). Panama, Chiriquí: 1♂, no date, holotype of Thecla latagus (ZMHB); 1♂
Potrerillos, 3000’, 29 Jan 1966 (USNM, Fig. 2, left); 1♀ Potrerillos, 3600’, 8 Aug
1965 (USNM). 1♂ Canal Area: Chagres Road, Ft. Sherman, May 1976 (MacD).
Ecuador, Pichincha: 2♂&1♀ 5 km Nanegal-García Moreno Rd., 0°09.2’S, 78°39.4’W,
1375-1700m, 15 Jan 2007, 15 Jan 2007, 3 Jun 2011 (RCB). 1♀ 20 km PactoGuayabillas Road,0°11.6’N, 78°51.5’W, 900 m, May 2010 (RCB). Esmeraldas: 1♀
Alto Tambo, 850m, Aug 2000 (RCB).
Misidentifications
The male of Thecla latagus illustrated in D’Abrera (1995: 1259) is a male of Lathecla
mimula.
66
Remarks
Schaus considered the female of L. winnie to be the female of L. latagus (see below),
which is undoubtedly why he described L. epopeoides as a distinct species.
Lathecla mimula (Draudt, 1920), Revised Status
Nomenclature
The identity of this name has long been in doubt because the original description and
illustration were poor, but after an extant type was found in SMF (Frankfurt) (Lamas
& Robbins, data not shown) (illustrated in http://butterfliesofamerica.com/, accessed
25 September 2013), Robbins (2004b) synonymized this name with L. latagus. With
the additional evidence gathered for this paper, we remove the name L. mimula from
synonymy.
The recognition of L. mimula as a distinct species was detailed in the account for
L. latagus. The wing pattern (Fig. 3), male genitalia (Fig. 15), and female genitalia
(Fig. 21) are illustrated. Mean male forewing length = 1.0 cm, SD = 0.07, N = 5. Mean
female forewing length = 1.0 cm, SD = 0.07, N = 3.
Variation
Females from French Guiana, Trinidad, Ecuador, Peru, and Brazil (RJ) are brown on
their dorsal wings, lacking any blue scaling. However, two of nine females from southern Brazil (MG, SC) have suffused light blue scaling on the dorsal wing surface, which
we attribute to intraspecific variation because males from the same localities are typical
of L. mimula.
Lathecla mimula occurs in forest from sea level to 1000 m elevation.
Behavior
Males and females of this species are attracted to traps baited with rotting fish. A male
in French Guiana was observed setting up a mating territory at 14:00 h (Faynel 2003).
Lathecla mimula inhabits lowland forest in the Amazonian and Atlantic biogeographic
zones of Brown (1982). Most records are from wet forest. As noted, it is allopatric or
parapatric with L. latagus. It occurs at lower elevations than L. fernandezi, L. carolyna
and L. mitzi (so far as is known).
67
Material examined (22♂ and 14♀)
French Guiana, 1♂ Saül, Boeuf Mort, 28 Oct 2001, hilltopping 14:00 hours (CF).
2♀ Saül, 3°37’N, 53°12’W, 20 Jan 2013 on a hill-top at 12:40 hours (CF), on a hilltop at 13:50 hours (USNM). Trinidad, 1♂ St. Ann’s, 1300 ft (listed as Thecla odinus
in Kaye 1940, Barcant 1970) (M. Cock compared this specimen in the Barcant
Collection with a specimen identified by us). 1♂ (papered specimen from F.C. Urich
Collection sent for identification). 1♂ and 1♀Lalaja Ridge, Aug 1989, May 1993 in
the Alston-Smith Collection (images sent for identification). Ecuador,1♂ Sucumbíos,
5 km. Puerto Libre-La Bonita Road, 0°13.0’N, 77°29.3’W,700m, 9 Sep 2005 (RCB).
2♂ and 1♀ Napo, 14 km Tena-Puyo Road, 0°06.7’S, 77°46.9’W, 600m, 17 Oct 2011,
2-3 Oct 1997, 11 Oct 2011 (RCB). 1♂ Morona-Santiago, Santiago (hill north of
town), 3°02.3’S, 78°00.3’W, 300 m, 24 Sep 2012 (RCB). Peru, 2♂ Loreto, Rio
Ucayali, Contamana, Jul 2007 (identified by C. Faynel) (CF), 1♀ Pachitea, [Huánuco,
08º45’S, 74º32’W, 150 m in Lamas 1976], a syntype of Thecla mimula (SMF). Bolivia,
1♂ Las Juntas, Nov 1913 (CMNH). Brazil, 1♀ MA, Imperatriz (DZUP). 1♂ GO,
Vianópolis, illustrated in D’Abrera (1995: 1258-1259) from the Spitz collection
(BMNH) under the name, Thecla latagus. 1♂ and 1♀ MG, Conceição dos Ouros, 900
m (examined images, MC). 5♂&1♀ RJ, Petrópolis, Independência, 1,000 m 1 May
1969 (USNM), 900m 16 Jan 1972 (USNM), 900m 3 Nov 1972 (FSMC), 25 Mar
1937 (DZUP), 12 Nov 1938 (DZUP), 2 Jul 1939 (DZUP). 1♂ (DZUP) and 3♀ RJ,
Guapimirim, 22°32’S, 42°59’W, 9 May 1969 (USNM), 7 Jul 1974 (USNM), 7 Jul
1974 (USNM). 1♂ RJ, Rio de Janeiro, Morro Dona Marta (DZUP). 1♂ and 1♀ SC,
São Bento do Sul 500-800 m (examined images, MC). 1♂ PR, P[ar]q[ue] Mun[icipal]
Três Morrinhos, Terra Rica, 600 m, 14 Dec 2009. 1♀ PR, Est[ação] Ecol[ógica do]
Caiuá, Diamante do Norte, 6 Oct 2009. 1♀ PR, RPPN Faz[enda]. Matão, Loanda, 20
Apr 2008.
Misidentifications
Kaye (1940) and Barcant (1970) misidentified this species as Thecla odinus G. & S., a
name that is now placed in the synonymy of Ocaria aholiba (Hewitson) (Robbins
2004b). D’Abrera (1995: 1212) illustrated a male and female that he identified as
Thecla mimula. However, the male does not belong to Lathecla, and the generic placement of the female cannot be determined from the image. D’Abrera also illustrated a
male of L. mimula under the name L. latagus, as noted in the material examined
section.
Remarks
This species is widespread in South American lowland forest east of the Andes,
but does not seem to be especially common or regular in its occurrence at any
locality.
68
Lathecla fernandezi Robbins & Busby, new species
Type material
Holotype (Fig. 4, left). [red printed label]Holotype/Lathecla fernandezi/Robbins &
Busby. [white printed label]ECUADOR: Napo/10 km El Chaco-El Reventador
Rd/0°16.5’S, 77°45.6’W/11 February 2008 1800-1950m/Robert C. Busby leg.
Deposited USNM.
Paratypes (9♂ and 3♀). Venezuela: Aragua, Rancho Grande, 1100 m: 2♂ 26 May,
19 Jun 1985 (USNM). Ecuador: Pichincha, 4 km E Tandapi, 2050 m, 0°26.6’S,
78°46.7’W, 1♂ 23 Sep 2002 (USNM); Loja, 25 km Catamayo-Piñas Road, Las
Chinchas, 3°57.6’S, 79°29.2’W, 2100-2250 m, 1♂ 12 May 2004 (USNM), 1♂ 16
May 2010 (RCB), 4♂ 22 May 2010 (RCB); 2♀ Napo, 10 km El Chaco-El Reventador
Rd., 0°16.5’S, 77°45.6’W, 1800-1950 m, February 2010 (Fig. 4, right, RCB).
Etymology
It gives us great pleasure to name this species in honor of the late Francisco Fernández
Yépez. He was the leading lepidopterist in Venezuela for decades and facilitated the
collecting trip in 1985 when the first specimen of this species was collected (it is not
the holotype because it was a worn specimen). The masculine name is in the genitive
case, as requested by Francisco’s students.
Type locality
The holotype was collected along a ridge that is fairly steep and heavily forested on one
side and gently sloped and partially deforested on the other.
Males of L. fernandezi are immediately identifiable by the off-white color of the dorsal
hindwing androconia (Fig. 9). This androconial color is unique in Lathecla and very
unusual for any eumaeine. Wing pattern (Fig. 4), androconia (Figs 8–10. left), male
genitalia (Fig. 17) and female genitalia (Fig. 22) are illustrated. Mean male forewing
length = 1.1 cm, SD = 0.10, N = 5. Mean female forewing length = 1.2 cm, SD = 0.08,
N = 2.
This species occurs in wet montane forest at a wide range of elevations from 1100 m to
2200 m.
Behavior
Males at Las Chinchas (Loja Province in southwest Ecuador) exhibited territorial
behavior in the morning on tall vegetation along a ridge top. Specific records for the
individuals from 16–22 May 2010 are 09:40, 10:06, 10:20, 11:19 and 11:25 h. Four
of the five males were landed 7 to 9 m above the ground while the fifth was at 2 m. One
69
of the males from eastern Ecuador (Napo Province) was found on a flowering bush at
11:15 h on a ridge top. The Venezuelan paratypes were found flying south against the
wind through Portachuelo Pass, which is a renowned site for insect migration, including Lycaenidae (Beebe 1949, 1951).
Lathecla fernandezi occurs widely in the Andes from western Ecuador to northern
Venezuela to eastern Ecuador. It may yet be found in eastern Peru. Like L. carolyna, it
occurs on the western and eastern slopes of the Andes at the same elevations and has
been found at the same locality in western Ecuador as L. carolyna.
Lathecla vichai Robbins & Busby, new species
Type material
Holotype ♂ (Fig. 5, left). [red printed label] Holotype/Lathecla vichai/Robbins &
Busby. [white printed label, date handwritten in India ink] PANAMA: Chiriqui:/
Volcan Baru: 1,800 m/13 March 1984//Leg. G. B. Small. Deposited USNM.
Paratypes (20♂ and18♀). Panama, Chiriquí, Volcán Barú, 8°48’N, 82°29’W (same
as the type locality, all collected by G. B. Small), listed by elevation. 5,600’: 1♂ 6 Mar
1976, 1♀ 12 Mar 1976, 1♂ 14 Mar 1976, 1♂ 21 Mar 1976; 1,700 m: 1♀ 28 Feb
1976, 2♂ 5 Mar 1976, 1♀ 14 Mar 1976, 1♀ 19 Mar 1976, 1♀ 19 Feb 1977, 1♂&1♀
22 Feb 1977, 1♂ 23 Feb 1977, 2♂ 19 Feb 1979; 1,750 m: 4♂&3♀ 14 Feb 1981, 1♀
14 Feb 1985, 1♂&1♀ 15 Feb 1985, 1♀ 16 Feb 1985; 1,800 m, 2♂ 22 Feb 1976, 1♂
27 Dec 1976, 2♂ 28 Dec 1976, 3♀ 13 Mar 1984, 1♀ 4 Mar 1985, 1♀ no date; 1,300
m, 1♂ 19 Mar 1976 (a female was collected at 1,700 m on the same day with the same
locality, for which reason we consider the 1,300 m elevation on this handwritten label
an error); no elevation, 1♀ 23 Mar 1985.
Other specimens examined (excluded from the type series)
Guatemala, 1♀ Volcán Sta. María, Jul (USNM). Nicaragua, 2♂ Matagalpa, 10 km.
N., [1,000-1,400 m, cited in Robbins et al. 2012], 14 Sep. 1976, 11 Oct 1976 (USNM).
Costa Rica, Puntarenas, Monteverde, 1♂14 Sep 1996, 1♀ 14 Sep 1971)(USNM).
Etymology
It gives us great pleasure to name this species in appreciation of Vichai Malikul. With
a trained eye and enormous native talent, Vichai has illustrated genitalia and other
eumaeine structures for decades. The name is a noun in apposition.
Type locality
The type locality in 1984 was forest at 1700 m elevation on the eastern slopes of Volcán
Barú along a dirt road that went north and then west from the town of Boquete,
70
Chiriquí. According to Gordon Small, who collected the holotype, the type locality on
the eastern slope of the volcano had less annual precipitation than the western and
northern slopes.
The dark gray scaling on basal edge of the postmedian line is wide and gives the impression of a band (Fig. 5). This trait distinguishes L. vichai from all other eumaeines. The
wing pattern (Fig. 5), androconia (Figs 8–10, right), male genitalia (Fig. 16) and female
genitalia (Fig. 23) are illustrated. Mean male forewing length is 1.0 cm, SD = 0.11,
N = 10. Mean female forewing length is 1.0 cm, SD = 0.10, N = 10.
Variation
The number of submarginal orange lunules on the ventral surface of the hindwing
ranges from two to seven.
Habitat, elevation and seasonality
The habitat of L. vichai is montane forest from 1400 to 1800 m elevation. Dates of capture
are November to March at the type locality and July to September at other localities.
Lathecla vichai occurs in montane forest from Guatemala to Panama and overlaps the
range of L. winnie in Costa Rica at the same elevations, but appears to occur primarily
in drier forest than L. winnie.
Remarks
The first female of L. vichai was collected over a century ago in Guatemala (in the
Schaus collection, USNM). It probably was not described because it is in poor condition and because no other specimens were known.
Lathecla carolyna Busby, new species
Type material
Holotype (Fig. 6, left). [red printed label]Holotype/Lathecla carolyna/Busby. [white
printed label]ECUADOR/Pichincha Province/Tandapi (1800m)/4. X. [Oct] 1988/
leg. Robert C. Busby. Deposited USNM.
Paratypes (6♂ and 3♀). Ecuador:Pichincha 1♂ Tandapi, 1,800 m, 4 Oct 1998;
1♂&1♀ 4 km E Tandapi, 0°26. 6’S, 78°46. 7’W, 2,050 m, 23 Sep 2002; 30 km W of
Aloag, 2,200 m, 1♂ 12 Oct 1995, 1♀ 19-20 Oct 1991, 1♀ 16-20 Oct1989. MoronaSantiago 2♂ 18 km Macas-9 de Octubre Road (Río Abanico), 2°15. 6’S, 78°12. 9’W,
2,050 m, 2 Oct 2004.
71
Etymology
This beautiful species is named in honor of Carolyn Weigandt, sister of Robert Busby.
She is an accomplished nature photographer, and butterflies in the wild have always
been among her favorite subjects. The name is a non-Latinized noun in apposition.
Lathecla carolyna has a wing shape and pattern that is similar to that of L. vichai and
L. winnie (Figs 5–7). It is immediately distinguished from L. vichai by the lack of dark
gray banding on the ventral wings. It differs from L. winnie, its phylogenetic sister, by
the more extensive blue on the dorsal surface of the forewings. Wing pattern (Fig. 6),
male genitalia (Fig. 19) and female genitalia (Fig. 24) are illustrated. Mean male forewing length is 1.1 cm, SD = 0.08, N = 4. Mean female forewing length is 1.1 cm,
SD = 0.11, N = 2.
Type locality
In the late 1990s, the type locality was accessed from the main road from Aloag to
Santo Domingo just before the town of Tandapi. A wide dirt path/road “climbed”
200–300 m along a ridge before leveling out. One side of the ridge was steep and well
forested, while the other side was more level with less vegetation. Trees and shrubs
(generally between 5–10 m in height) lined the road along the steep side of the ridge
and were readily used by perching butterflies. By the early 2000s, there was less forest
along this ridge because of increased agricultural activity, and fewer butterflies.
Lathecla carolyna occurs in wet montane forest from 1800 to 2200 m elevation.
Behavior
Males in the western mountains of Ecuador (Pichincha) exhibited territorial behavior
on vegetation along prominent ridges and hilltops in the mid-to-late morning.
Lathecla carolyna has been found on both the eastern and western slopes of the Andes
in Ecuador. It is sympatric with L. fernandezi in the Tandapi area of western Ecuador.
Lathecla winnie Robbins & Busby, new species
Type material
Holotype (Fig. 7). [red printed label]Holotype/Lathecla winnie/Robbins & Busby.
[white label handwritten in India ink]Costa Rica/San Jose Prov. /N. of San Isidro/5600’
– Hwy 2/Talamanca Mts. /7 – vi[Jun] – [19]88– [Brian] Harris. Deposited USNM.
72
Paratypes (6♂ and 3♀). Costa Rica, Heredia, Rio Sarapiquí: 1,300 m, 1♂ 23 Jun
1976, 3♂ 25 Jun 1976, 1♂ 29 Jun 1976, 1♂ 10° 43’N83° 56’W, 700 m, 26 Jun 1976,
1♀ 1,150 m 6 Sep 1977. 1♂ Costa Rica, San José, Parque Braulio Carillo, La Montura,
10°05’N 83°30’W, 1,000 m. 1♀ Costa Rica, [Alajuela], Mount Poás, May.
Etymology
It gives us great pleasure to name this beautiful species, which is endemic to Costa
Rica, to honor biologist Winnie Hallwachs. She and her husband Daniel Janzen have
made prodigious strides in documenting the Costa Rican lepidopteran fauna, their
immature stages, and their caterpillar food plants. The name is a noun in apposition.
The blue scaling on the dorsal surface of the forewing in males is restricted to the inner
margin area, which is a unique trait in Lathecla. This species is the phylogenetic sister
of L. carolyna. Wing pattern (Fig. 7), male genitalia (Fig. 18), and female genitalia
(Fig. 25) are illustrated. Mean male forewing length is 1.0 cm, SD = 0.13, N = 7. Mean
female forewing length is 1.0 cm, SD = 0.02, N = 3.
Type locality
In 1988 the type locality was lightly disturbed montane forest bordering paved
Highway 2.
Lathecla winnie is recorded from montane wet forest from 700 to 1700 m elevation.
Lathecla winnie is endemic to Costa Rica, but it should also occur on the wet Atlantic
slopes of Panama. Both L. winnie and L. vichai have been found in the Talamanca
Mountains at similar elevations, but it appears that L. winnie occurs in wetter habitats
than L. vichai.
Remarks
As with L. vichai, a female of this species was deposited in the Schaus Collection
(Mount Poás, USNM). It bears a label in Schaus’s handwriting identifying it as Thecla
latagus, which is probably the reason that L. winnie was not described long ago.
Phylogenetic analyses
There was one most parsimonious 34-step equal weight tree (CI = 0. 91 and RI =
0. 94) with bootstrap values as noted (Fig. 11). When the data were analyzed with the
73
androconial characters omitted (characters 3, 5, 8), the tree topology was the same.
Further, each implied weight most parsimonious tree with different values for the
parameter K resulted in one tree, also with the same topology as the most parsimonious
equal weight tree.
The three kinds of scent patches in Lathecla evolved at different points on the cladogram. The ventral forewing patch (Fig. 8, character 5) evolved once at node B and was
not lost (Fig. 11). The dorsal hindwing patch (Fig. 9, character 8) evolved at node A
and was not lost (Fig. 11), but this result depends, in part, on the outgroups species
chosen because a dorsal hindwing patch also occurs in some species of outgroup genus
Rhamma. The dorsal forewing patch on the cubital vein (Fig. 10, character 3) evolved
once at node C and was not lost (Fig. 11).
Discussion
Generic taxonomy
Lathecla was tentatively placed next to Thestius in a checklist (Robbins 2004b) because
Lathecla and Thestius share superficially similar genitalic structures. We transfer Lathecla
to the Micandra Section of the Eumaeini (Robbins 2004b) next to Podanotum on the
basis of two characters. First, most Lathecla species possess a female genitalia “fanshaped signa” (lacking in Thestius). This structure is best developed in L. fernandezi and
L. vichai (Figs 22, 23), but was previously overlooked because the signa are small in
other Lathecla species (Figs. 20, 21), perhaps best described as vestigial in L. carolyna
and L. winnie (Figs 24, 25). Fan-shaped signa occur only in the Calycopidina and the
Micandra Section of the Eumaeini (Robbins & Busby 2008, Duarte & Robbins 2010,
including a morphological description). Further, the relatively straight anterior edge of
the signa on the corpus bursae occurs only in Lathecla and Podanotum. Second, the
postmedian line on the ventral surface of the forewing terminates posteriorly at vein
Cu2 in Lathecla and Podanotum. It terminates at the inner margin in Thestius, at least
in those species in which the wing pattern has not been greatly modified. The genus
Rhamma was closely related to Podanotum in a phylogenetic analysis of DNA sequences
(Quental 2008), for which reason it was also used as an outgroup.
Male secondary sexual structures
The evolution of male secondary sexual structures has long been implicated in speciation because these structures are postulated to promote species recognition among
females and to contribute to reproductive isolation among sympatric populations
(Phelan & Baker 1987; Löfstedt et al. 1991; Symonds & Elgar 2008). More generally,
male secondary sexual character diversification is hypothesized to be correlated with
species diversification (e.g., Streelman & Danley 2003; Polak & Simmons 2009 and
included references).
Lathecla is a highly unusual eumaeine genus in that two male secondary sexual structures were gained evolutionarily. The scent patch on the dorsal forewing cubital vein
74
occurs in no other lycaenid species and evolved at node C (Fig. 11). The scent patch on
the ventral surface of the forewing evolved at node B (Fig. 11) and occurs in none of
the outgroups. The scent patch on the dorsal surface of the hindwing ostensibly evolved
at node A (Fig. 11), but this result is uncertain because the scent patch also occurs in
some of the outgroups.
The purpose of this paragraph is to assess preliminarily the consequences of a gain in
male secondary sexual structures in eumaeines. In Timaeta, the lineage with the comblike androconia diversified into five species while in the same period of time its sister
lineage did not diversify (one extant species) (figure 75 in Robbins & Busby 2008). In
Lathecla, the lineage possessing a ventral forewing scent pad diversified into six species
while its sister lineage did not diversify (one extant species) (Fig. 11, node B). Further
in Lathecla, the lineage with a dorsal forewing cubital scent patch diversified into four
species while its sister lineage diversified into two species (Fig. 11, node C). The probability that in all three cases the lineage gaining a male secondary sexual structure
would have more species at random is 1/8 (p = 0.125). Even though this result is statistically insignificant because of small numbers, it shows the viability of the hypothesis
that a gain in a male secondary sexual structure in eumaeines leads to an increased
diversification of species. We note that statistical methods have been developed for
DNA sequence data that incorporate information on timing of branching events (e.g.,
Maddison et al. 2007, O’Meara 2012). These methods are more powerful when DNA
sequence data are available and when the assumptions on which they are based are met.
The scent patch on the ventral surface of the forewing (Fig. 9) is the only known case
so far in the Eumaeini in which a male secondary sexual structure was re-gained evolutionarily. The cubital vein scent patch (Fig. 10) in Lathecla and the comb-like androconia in Timaeta are structures that evolved once and were not lost (Robbins & Busby
2008). The significance of the scent patch on the ventral surface of the forewing is that
other cases where secondary sexual structures re-evolved are likely to be discovered by
phylogenetic analyses. If so, the hypothesis in the previous paragraph will be able to be
tested.
Biogeography
Willmott et al. (2001) proposed a four-factor model for diversification of montane
Andean genera. Species might speciate between the Andes and other montane regions,
speciate across elevational gradients, radiate within the Andes, or colonize the Andes
from temperate latitudes (not relevant for strictly Neotropical taxa, such as Lathecla).
We briefly assess the apparent importance of the first three factors in the biogeographic
diversification of Lathecla and contrast it with that of Timaeta, the only other primarily
montane eumaeine genus to be revised phylogenetically (Robbins and Busby 2008).
The most frequent biogeographic mechanism of speciation in Lathecla would appear
to be allopatric speciation in isolated mountain ranges (4 of 6 nodes). Lathecla species
appear to have been “spun off” from the Andes to the southern Brazil mountains (M.
mitzi), to the lowlands (M. latagus + M. mimula), and twice to the Central American
mountains (M. vichai, M. winnie). This pattern is arguably the one predicted by
75
William Brown’s (1957) model of centrifugal speciation. In contrast, this mechanism
was unimportant in Timaeta (0 of 14 nodes), which is an Andean endemic. The second
mechanism is change in elevation, in which M. latagus + M. mimula colonized the
lowlands (1 of 6 nodes), which then led to another speciation outside of montane
areas. This is only the second time that a montane eumaeine lineage has been shown to
have colonized the lowlands (Robbins et al. 2010). This mechanism was similarly infrequent in Timaeta (2 of 14 nodes). The third mechanism of radiation within the Andes
apparently played no role in the diversification of Lathecla (0 of 6 nodes). In contrast,
it was predominant in Timaeta (12 of 14 nodes). We suspect that the contrasting biogeographic patterns of Lathecla and Timaeta represent extremes along a continuum,
and as more primarily montane eumaeine genera are revised, the biographic patterns
of these two genera may be useful reference points.
Acknowledgements
We are grateful to Matthew Cock, Christophe Faynel, Dan Janzen, Gerardo Lamas,
John MacDonald and Alfred Moser for kindly sharing information and/or specimens
and for commenting on the manuscript. We are especially grateful to Diego Dolibaina
for making substantive contributions to the paper, including obscure information on
the type locality of L. mitzi. In addition, two anonymous reviewers made insightful
comments that improved the manuscript. We thank Karie Darrow, Vichai Malikul and
Brian Harris for help with imaging, illustration, and a variety of technical aspects.
Thanks to the Hennig Society for making TNT software freely available.
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Evolutionary gain of male secondary sexual structures

Transcription

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