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Accepted by R. Anderson: 27 Jun. 2012; published: 8 Aug. 2012
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2012 · Magnolia Press
Zootaxa 3413: 55–63 (2012)
www.mapress.com/zootaxa/Article
55
A remarkable new species of flesh-fly mimicking weevil (Coleoptera:
Curculionidae: Conoderinae) from Southeastern Brazil
SERGIO A. VANIN1,4 & TADEU J. GUERRA2, 3
1Departamento de Zoologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, Travessa 14, 101, 05508-900, São
Paulo, SP, Brazil. E-mail: savanin@ib.usp.br
2Programa de Pós-Graduação em Ecologia, Instituto de Biologia, Universidade Estadual de Campinas (UNICAMP), C.P. 6109,
13083970, Campinas, SP, Brazil
3Actual address: Departamento de Biologia Geral, Instituto de Ciências Biológicas, Universidade Federal de Minas Gerais. C.P. 486,
30161-970, Belo Horizonte, Minas Gerais, Brazil. Email: guerra.tj@gmail.com
4Corresponding author. E-mail: savanin@ib.usp.br
Abstract
Timorus sarcophagoides, new species (type-locality: Brazil, Minas Gerais, Santana do Riacho - Serra do Cipó, 43o35'W
19o17'S, 1200–1300m ASL), is described and illustrated. The new species can be distinguished mainly from the other spe-
cies of the genus by the sexual dimorphism of the male rostrum armed with a hooked tubercle at the base of the dorsal
carina, while the tubercle is absent in the female and the corresponding region of the carina is only tumid. The new species
has a striking pattern of coloration and behavior that mimics flesh-flies in the family Sarcophagidae. Observations on the
natural history of the new species are reported and discussed.
Key words: mimicry, natural history, Sarcophagidae, Serra do Cipó, taxonomy, Timorus, Zygopini
Resumo
Timorus sarcophagoides, nova espécie (Localidade-tipo: Brasil, Minas Gerais, Santana do Riacho - Serra do Cipó,
43o35'W 19o17'S, 1200–1300m), é descrita e ilustrada. A nova espécie pode ser distinguida das outras espécies do gênero
principalmente pelo marcante dimorfismo sexual no rostro dos machos armados com um tubérculo em forma de gancho
na base dorsal da carina, enquanto nas fêmeas o tubérculo é ausente e a região correspondente da carina é apenas intu-
mescida. A nova espécie possui um notável padrão de coloração e comportamento que mimetiza moscas da família Sar-
cophagidae. Observações sobre a história natural da nova espécie são apresentadas e discutidas.
Introduction
The subfamily Conoderinae 1833 (until recently known as Zygopinae Lacordaire, 1866) includes more than 200
genera and approximately 1,500 species worldwide (Alonso-Zarazaga & Lyal, 1999). Most of the genera of
Conoderinae were described in the 19th or early 20th century and have never been revised. The only available keys
for genera of Conoderinae of the world were provided by Heller (1894, 1895). The delimitation of genera is very
difficult because of the lack of a phylogenetic hypothesis of the subfamily and becomes even more difficult with
the discovery of large numbers of undescribed species that link genera or do not fit current concepts (Hespenheide,
2009).
Weevils in the Conoderinae are known often for their distinct color patterns related to predator avoidance
(Hespenheide, 1995). For instance, in Panama and Costa Rica nearly 20% of species are involved in mimicry com-
plexes based on putative models such as flies, ants, beetles and bees (Hespenheide, 1995). However, most mimetic
species in Conoderinae resemble flies (Hespenheide, 1995). Hespenheide (1973) first described a mimicry com-
plex involving Neotropical beetles and flies, and listed nearly 60 species in 21 genera in six families with a conver-
VANIN & GUERRA
56 · Zootaxa 3413 © 2012 Magnolia Press
gent coloration pattern resembling flies, mostly from Central America tropical forests. Since these species are
found in different beetle families and genera in which many species are not mimetic, fly mimicry is probably an
adaptive form of homoplasy (Hespenheide 1973, 1995). Hespenheide (1973) reported 34 fly mimicking species in
nine genera among Neotropical Zygopinae (Conoderinae), but in a subsequent review (Hespenheide, 1995) the
author indicated 75 species in nine genera among Conoderinae as mimics of flies. However, many species of this
group remain undescribed, and descriptions of those fly mimicking weevils available are based in specimens from
museum collections and, therefore, basic information on their biology is still lacking.
The senior author (SAV) was requested by the second author (TJG) to identify a remarkable fly-mimicking
weevil. The species was promptly identified as belonging to the Conoderinae-Zygopini, because the rostral channel
is formed only by the prosternum. A search of Curculionidae in the collection of the "Museu de Zoologia, Univer-
sidade de São Paulo" (MZSP) disclosed some Conoderinae weevils with similar color pattern, but only two uniden-
tified conspecific specimens, one also from the State of Minas Gerais (Lagoa Santa) and another from the State of
Rio de Janeiro (Rio de Janeiro). A closer examination of those specimens showed that they would key to Timorus
Schoenherr, 1838, in Heller's key (1845). We give below a description of the new species and additionally we
report observations on its natural history.
Material and methods
The taxonomic study was done by S. A. Vanin. Adults were dissected in water, under a stereomicroscope (Wild
M5A). Temporary slides were mounted in glycerin and drawings were made using a Leitz SM-Lux compound
microscope or a stereomicroscope Wild M5A, both fitted with a camera lucida. Adult habitus photos were taken
with a stereomicroscope Leica M125 with coupled Magnifier in DV camera Leica DFC.
The field observations and specimens collections were conducted by T. J. Guerra at Reserva Particular do
Patrimônio Natural (RPPN) Vellozia, a private area of “Campos Rupestres” varying from 1100 to 1400 m above
sea level, in the vicinity of Serra do Cipó National Park, Municipality of Santana do Riacho, Minas Gerais,
Southeastern Brazil (43º 35’W, 19º 17’S). The vegetation is typical of the Espinhaço Mountain Range, comprising
mosaics of open fields and rocky outcrops with developed herbaceous strata, many bushes and sparse small trees
(Fig. 16). These ecosystems harbor a highly diversified flora associated with quartzite rock outcrops and with
shallow, acid and nutrient poor soils in areas higher than 1000 m above sea level (Giulietti et al., 1997). Climate is
mesotermic (Cwb of Köpen) with cold dry winters from May to September and hot wet summers from October to
April (Madeira & Fernandes, 1999). In the field observations, nearly 50 host plants were inspected monthly from
October 2007 to March 2009, when some host plants were pruned and placed in rearing containers for inspection
during the following months.
Results
Taxonomy
Timorus sarcophagoides, sp nov.
(Figs. 1 15)
Type material. Brazil. Holotype male (dissected). Minas Gerais, "Santana do Riacho - Serra do Cipó 43o35'W
19o17'S, altitude 1100–1300m, XII.2007 T.J. Guerra col.; campo rupestre, on Psittacanthus robustus, Loran-
thaceae", (MZSP). Paratypes: same data as holotype, 4 (1 dissected), 7 (2 dissected) (MZSP); Minas Gerais,
"Lagoa Santa, 26.XI.1960, Araújo e Martins", 1 (MZSP). Rio de Janeiro, "Rio de Janeiro (Corcovado), X. 1957,
M. Alvarenga", 1 dissected (MZSP).
Description. Length (rostrum excluded), male: 6.3–8.4 mm; female: 8.7–10.0 mm. Body rhomboidal. Integu-
ment black, vestiture formed by colored scales; frons ochreous, yellow above and behind eye; middle longitudinal
carina and lateral triangular-shaped area behind eye glabrous and shiny-black; rostrum with whitish scales, denser
in basal half. Prothorax with one broad patch of dense carmine scales on anterior margin, extending down to flanks;
yellow scales forming irregular fringe almost bordering each carmine patch laterad and posteriad, and less dense
Zootaxa 3413 © 2012 Magnolia Press · 57
NEW SPECIES OF FLESH-FLY MIMICKING WEEVIL FROM BRAZIL
yellow scales forming three pairs of spots, one elongate oval on each side of median longitudinal pronotal carina
and two irregular-shaped behind carmine patch; basal third of elytra with ochreous and yellowish-white scales con-
centrating densely on interstriae 2, 4, 6 and 8, while interstriae 1, 3, 5 and 7 blackish, resulting in contrasting pat-
tern of light and dark irregular stripes; epipleura with two dark spots. Ventral margin and legs covered by elongate
whitish scales; with oval patch of ochreus scales on each side, extending from ventrite II to base of ventrite V, sur-
rounded by yellow scales. Metespisternum and base of ventrites I and II glabrous.
FIGURES 1–7. Timorus sarcophagoides, sp. nov., habitus: 1, male holotype (length: 8.4 mm), lateral view; 2, male holotype,
dorsal view; 3, female paratype from Santana do Riacho (length: 8.7 mm), lateral view; 4, female paratype, dorsal view; male
paratype from Rio de Janeiro (length 6.3 mm), detail of head and pronotum, lateral view; 6, same, frontal view of head and ros-
trum; 7, female paratype from Santana do Riacho, frontal view of head and rostrum.
VANIN & GUERRA
58 · Zootaxa 3413 © 2012 Magnolia Press
FIGURES 8 15. Timorus sarcophagoides, sp. nov. FIGURES 8 11, paratype from Santana do Riacho: 8, body, ventral view,
head, median and hind legs omitted; 9, spermatheca, lateral view; 10, coxites and styli, dorsal view; 11, sternite VIII, ventral
view; FIGURES 12 15, holotype, male terminalia: 12, tegmen, lateral view; 13, tegmen, dorsal view; 14, median lobe, lateral
view; 15, median lobe, dorsal view.
Zootaxa 3413 © 2012 Magnolia Press · 59
NEW SPECIES OF FLESH-FLY MIMICKING WEEVIL FROM BRAZIL
Head (Figs. 5 7) Eyes oval, acuminate inferiorly, inner margin sinuous, large and well separated, narrowest
distance in front about 0.9X width of rostrum at base; inferiorly more separated by distance about 1.2X width of
rostrum at base; front shallowly concave between eyes, with middle carina extending from base of head to central
fovea. Rostrum slightly longer than pronotum, moderately stout, weakly curved, feebly depressed in basal half,
with dorsal carina more developed in males, with tuberculiform process at base; antennal insertion behind middle.
Antenna with scape clavate, not reaching base of eye, shorter than funicle; funicle with seven antennomeres; anten-
nomeres 1st and 2nd with similar lengths; 3rd and 4th subequal , about 0.7X as long as 2nd, 5th to 7th subequal,
about as long as broad, slightly shorter than 4th; club oval, about as long as length of three preceding antennomeres
combined, basal antennomere about half as long as club length.
Prothorax (Figs. 1 4) subtrapezoidal, rather convex, transverse, slightly broader than long (1.1–1.2X);
rounded at posterior angles, wider near posterior fifth and there converging gradually towards weakly constricted
apex; anterior margin straight with acuminate angles, posterior margin bisinuous and with basal lobe produced and
emarginate posteriad before scutellum; dorsal middle carina not attaining frontal and hind margins, anterior half of
carina sharp, posterior half wider, flattened and sulcate at middle; pronotal disc with depressed areas smoother, sur-
rounded by densely rugosely punctate and bare areas. Postocular lobes prominent.
Scutellum exposed, free, rounded, tuberculiform, with withish scales.
Elytra (Figs. 1 4) 1.3 1.4 times as long as broad, wider than prothorax, elongate, margins gently converging
posteriorly and abruptly constricted near apices, sharply margined externally by acute carina on intestriae 9; humeri
prominent; apices obliquely truncate, each produced into a minute spine. Elytral striae poorly developed, each
puncture with small scale inside; alternate interstriae 3, 5, 7 and 9 with sharp, bare and shiny carina from base to
elytral declivity; alternate interstriae 2, 4, 6 and 8 with feeble carina; interstriae 1 and 2 adjacent to scutellum
slightly depressed; margin of elytron recovered by slender yellowish scales, outer margin minutely crenulated
Venter (Fig. 8). Rostral canal formed by prosternum and sides of forecoxae extending to mesosternum. Pros-
ternum with deep canal, bordered by sharp carina reaching the anterior region of forecoxae; inner side of each fore-
coxae prolonged posteriad by flattened process forming continuing margin of rostral canal. Mesosternum declivous
posteriad, depressed anteriad at middle accommodatingtip of rostrum. Metaventrite convex.
Abdomen with ventrites 3 and 4 equal in length. Pygidium covered.
Legs (Figs. 1, 3). Femora curved, compressed and weakly clavate, carinate on both inner and outer surfaces
and armed beneath with oblique tooth; hind femora larger than fore and midfemora, their apices just exceeding pos-
terior margin of ventrite 3. Tarsal claws simple.
Male terminalia. Aedeagus (Figs. 14, 15): median lobe slender, feebly curved, 3.3 X as long as wide (median
struts excluded), lateral margins converging gradually from base to near apex, then more strongly convergent and
ending in triangular point; median lobe about 1.3 X as long as median struts; endophallus with a pair of uncinate
sclerites. Tegmen as in Figs. 12, 13.
Female terminalia. Coxites (Fig. 10) elongate, weakly sclerotized; stylus apical, elongate, cylindrical, slightly
curved outwards, apex rounded and setulose; sternite VIII elongate (Fig. 11), setulose, apex truncate. Spermatheca
(Fig. 9) U-shaped, ramus and collum closely approximate.
Etymology: Greek, sarcophagoides, like sarcophagid flies.
Type-locality: Brazil, Minas Gerais, Santana do Riacho - Serra do Cipó 43o35'W 19o17'S, altitude 1100–1300m
ASL.
Geographic distribution: the new species is represented in Southeastern Brazil, in the states of Minas Gerais
and Rio de Janeiro.
Biology. Timorus sarcophagoides was found exclusively on live, adult and reproductive woody mistletoes,
Psittacanthus robustus Mart. (Loranthaceae) (Fig. 16). The new species is a specialized phytophage feeder on P.
robustus in both adult and larval stages. During two consecutive years of observation adult weevils were found
exclusively from November to February, overlapping the flowering period of the host plant species. Adult weevils
fed mostly on flower buds (Fig. 17). They usually pierced receptacles chewing soft tissues of the ovaries, but also
on the tips of closed buds feeding on pollen grains and stamens. Weevils also fed on soft tissues in leaf axils (Fig.
18). In January and February females were observed chewing tiny roles in the haustorial root of the host plant (Fig.
19) where they oviposited (Fig. 20). The larva is rhizophagous, developing as a borer inside the haustorium of the
host plants (Fig.21). Larvae were found from March to September, during the dry season. Pupae were found exclu-
sively inside haustorial roots in October, suggesting that the metamorphosis ends just before the beginning of the
VANIN & GUERRA
60 · Zootaxa 3413 © 2012 Magnolia Press
rainy season. Adults emerged from mistletoes in November 2009, coinciding with the beginning of the host plant
blooming period. While moving, weevils perform unusual jerking movements of the legs and a stereotypical leg
scrubbing behavior mimicking the behavior of flesh-flies. Disturbed weevils usually hide on the underside of
leaves or shoots moving in the opposite direction of the threat stimuli. More effective capture attempts usually
induced weevils to drop off of the host plants. Hand-captured individuals performed thanatosis (Fig. 22). The new
species is diurnal and at night individuals were observed resting completely motionless on leaves. Adult weevils
walk throughout the whole area of the host plant, rarely leaving mistletoes by flight. While chewing, weevils
inserted their long rostrum in host tissues, and this feeding behavior made it difficult for them to drop off or move
away quickly, probably making them more susceptible to attacks of natural enemies.
FIGURES 16 23. FIGURE 16, host plant Psittacanthus robustus Mart. (Loranthaceae); 17–22, Timorus sarcophagoides, sp.
nov; 17, weevil feeding on flower receptacle and detail of pierced flower bud; 18, weevil on leaf axils; 19, female chewing host
plant; 20, oviposition placed in the haustorium; 21, weevil larva developing inside host plant; note the excavated tunnel inside
haustorium; 22, thanatosis behavior of a captured weevil; 23 a flesh-fly model (Sarcophagidae) perched on host plant.
Zootaxa 3413 © 2012 Magnolia Press · 61
NEW SPECIES OF FLESH-FLY MIMICKING WEEVIL FROM BRAZIL
Discussion
The tribe Zygopini were first recognized as "Zygopides vrais" by Lacordaire (1866) and defined by him as
"Zygopides with the rostral channel formed only by the prosternum". Zygopini Lacordaire, 1866, as recognized by
Alonzso-Zarazaga and Lyal (1999) is a predominantly Neotropical tribe with 33 genera, of which 31 are
represented in the Neotropics and only two are exclusive to Africa (Cameroon and Tanzania). Heller (1894, 1895)
provided the only available keys to the genera of Conoderinae, where 23 of the genera of Zygopini considered valid
by Alonso-Zarazaga & Lyal (1999) are included, 18 of which are represented in Brazil according to Wibmer &
O'Brien (1986). Heller's key (1895) has many generic names considered synonyms by Alonso-Zarazaga & Lyal
(1999) and is outdated because 12 new genera of Zygopini have been described after 1895 (10 Neotropical and two
Ethiopian). Nevertheless, it is still useful for helping in identification of the Brazilian Zygopini since only two of
the genera represented in Brazil are not included in the key, Lissoderes Champion, 1906 (monobasic) and
Copturomorpha Champion, 1906. Timorus sarcophagoides differs from these two genera mainly by the following:
in Copturomorpha the metasternum is more or less hollowed between the middle coxae and the femora are not or
are only obsoletely dentate beneath, while in Timorus the metasternum is not hollowed and the femora are
distinctly dentate beneath; in Lissoderes the femora are linear and unarmed whilst in Timorus the femora are
weakly clavate and distinctly dentate beneath.
The new species cannot be assigned to any of the other eight Neotropical genera of Zygopini described by
Champion (1906) and Hustache (1932) after the publication of Heller's key (1895). Considering the seven genera
described by Champion (1906), Arachnomorpha, Microzygops and Philenis can be easily distinguished by having
the eyes subcontiguous or narrowly separated in front (eyes are widely separated in Timorus). Helleriella, Larides,
Phileas and Zygopsella have eyes well separated as in Timorus, however in Helleriella, Phileas and Zygopsella the
ocular lobes are very weak or absent. Moreover, in Heleriella the prothorax is cylindrical and much more elongate,
in Phileas and Larides the rostrum is shorter, more curved and robust, and in Zygopsella the prothorax is very gib-
bous and each elytron is armed with a stout dentiform process. Paramnemynellus Hustache, 1932 differs by the
femora sulcate beneath, the posterior femora not carinate and reaching the elytral apices, while in Timorus the fem-
ora are not sulcate or carinate on the outersurface, and the apex of the hind femora just surpasses the posterior mar-
gin of ventrite III.
The new species is tentatively assigned to the genus Timorus Schoenherr, 1838. Timorus has never been
revised and shares with other Zygopini genera many of the characters presented in the original description
(Schoennher 1838), such as the prominent postocular lobes, the femora unidentate beneath and carinate on the
outer edge, the hind femora slightly more elongate than the fore- and midfemora, and the hind femora with their
apices barely exceeding the posterior margin of ventrite II, the antennomeres 1st and 2nd with about equal length,
the rostrum dorsally carinate, moderately slender and weakly curved, the pygidium concealed by elytra, the
inferiorly acuminate eyes, and ventrite 2 as long as the length of 3 and 4 combined. Timorus includes seven
described species, five from Brazil and two from French Guyana (Wibmer & O'Brien, 1986). None of the seven
species described in Timorus by Chevrolat (1879), Heller (1904), Hustache (1938) and Rosenschoeld (1838) has
the sexual dimorphism or the coloration pattern similar to those reported above for T. sarcophagoides.
Sexual dimorphism involving modifications of the rostrum have been reported for other Neotropical
Conoderinae, including Lissoderes spp. (Hustache, 1938; Hespenheide, 1987) and Pseudolechriops spp.
(Hespenheide & Lapierre, 2006). Other outstanding sexual dimorphism was reported for some males of Old World
Zygopini, which have a pair of long thoracic spines similar to those occurring in some Baridinae weevils and used
for intra-specific contests (Kojima & Lyal, 2002). These sexually dimorphic weapon-like structures are expected to
evolve as a result of intra-sexual selection, when males interact agonistically to compete for females (Emlen,
2008). However, adaptive significance of sexual dimorphism of weevils` rostrum remains controversial (Wilhelm
et al., 2011). In effect, the natural history of the group is still poorly known (Souza et al. 1998; Weng et al., 2007)
and contests between males have been reported for only two Conoderinae (Lyal, 1986). The rostrum armed with a
hooked tubercle could be related to fighting ability of male weevils, although function of this structure in sexual
selection needs further investigation.
The new species has a striking pattern of coloration and behavior which resembles flesh-flies from Sarcophagi-
dae. In the weevil collection of the "Museu de Zoologia, Universidade de São Paulo", we found unidentified spe-
cies belonging to other genera of Conoderinae which are similarly colored and also resemble flesh-flies (Copturus,
VANIN & GUERRA
62 · Zootaxa 3413 © 2012 Magnolia Press
2 spp., Mnemynurus, 1 sp. and Zygops, 1 sp.). It should be noted that T. sarcophagoides has a pattern of coloration
extremely similar to a species of another genus listed by Hespenheide as a fly mimic (l.c.), the monobasic Panama-
nian Lechriopini Euzurus ornativentris Champion, 1906 (Champion, 1906, vol. 4, tab. 3, figs 21, and 21a). The dis-
tribution of the colored patches is very similar in both species, even on the abdominal ventrites.
This is the first case of fly mimicry reported for a South American weevil species and also the first case of such
a mimicry system in a savanna ecosystem. Red headed flies have been record as models for at least 58 species in
eight genera among Conoderinae (Hespenheide, 1995) in Central America tropical forests. However, Hespenheide
(1973) observed flies in families Tachinidae, Muscidae and Tabanidae as the models of mimicking weevils from
Central America. Flesh-flies were commonly observed on rock outcrops through the whole year, but they seem to
be especially abundant from November to March, during the wet and hot season. These flies frequently perch on
the vegetation, including flowers and leaves of P. robustus (Fig. 23). However, resemblance with flesh-flies is
probably not species specific. Considering size, presence of bright red eyes, variegated coloration of the dorsum’s
midsection and transparent wings, this weevil presents external convergence with flesh-fly species in the genera
Ravinia, Peckia (Euboettcheria) and Oxysarcodexia (C.A. Mello-Patiu, pers. com.). More studies are required to
assess model species involved in this mimicry system, their micro-habitat use and abundance through the year.
Hespenheide (1995) concluded that fly mimicry among weevils is a tropical phenomenon, occurring more fre-
quently among 4–8 mm phytophagous weevils that use branches and trunks as micro-habitat during their daily
activities. The new species is a tropical phytophagous weevil with life cycle associated with one plant species, as
most Curculionidae (Anderson, 1993; McKenna et al., 2009), matching size and micro-habitat use of other mimic
species (Hespenheide, 1995). Although weevils associated to mistletoes have been reported by Anderson (1994),
the author found no fly mimic species. According to Hespenheide (1973), adaptive significance of fly mimicry by
beetles is related to predator avoidance. Nevertheless, flies are not known to be chemically protected and distaste-
ful to predators. Hespenheide (1973) argued that external convergence with fly models confers protection to beetles
because flies are agile and are a hard to catch prey group. Since these mimicry systems involve bright coloration,
Hespenheide (1973) realized that the evolution of these adaptations must be related to selective pressure exerted by
visually oriented predators, particularly flycatching and foliage-gleaning birds. Because beetles are also quick and
difficult to capture, Hespenheide (1973) argued that this mimicry system can be classified as Mullerian, although
color convergence is probably not mutual.
Our observations on the natural history of T. sarcophagoides led us to agree with Hespenheide's hypothesis.
However, the ecological background of this mimicry system is still unexplored. Hespenheide's predictions regard-
ing adaptive significance of fly mimicry by Neotropical beetles remain untested in the field or under laboratory
conditions. Indeed, the evolution of mimicry systems based on palatable but difficult to capture models remains
controversial, and up to now, this phenomenon has not been demonstrated unequivocally (Ruxton et al., 2004). The
new species described here could be used as model for future studies addressing the occurrence of evasive mimicry
in nature.
Acknowledgments
To Dr. Cátia Antunes de Mello Patiu (Museu Nacional do Rio de Janeiro) for identification of the genera of the sar-
cophagid flies; to Dr. Ricardo Pinto da Rocha (Instituto de Biociências, Universidade de São Paulo) for providing
access to his photo equipament (FAPESP 2008/06604-7); to Bruno Medeiros for the stereomicroscope photos; to
Ricardo Pires Vanin for the electronic treatment of the habitus photographs; to Juares Fuhrmann, Cassio Coletinha
and Fabiano Albertoni for assistance with the electronic editing of figures 1 to 15 and 16 to 23, respectively; to
Daniela de Cassia Ben for inking the line drawings. Tadeu J. Guerra was supported by a doctoral fellowship from
Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) and a reseach grant (no. 07/59444-4)
from Fundação e Amparo a Pesquisa do Estado de São Paulo (FAPESP).
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