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The Description of Platphalonia magdalenae (Tortricidae, Tortricinae, Euliini, Cochylina) Found Nectaring Diurnally on Centromadia pungens (Asteraceae) in the Central Valley of California Along with a List of Species of Platphalonia

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Abstract

Platphalonia Razowski, 2011 (Tortricidae, Tortricinae, Euliini, Cochylina) was proposed for Saphenista mystica Razowski & Becker, 1983 (type species) and several species previously assigned to Platphalonidia Razowski, 1985. However, with the exception of the type species, none of the other purported congeners have been listed. We formally transfer 16 species to Platplialonia, resulting in the following new combinations: P. albertae (Razowski, 1997), P. assector (Razowski, 1967), P. californica (Razowski, 1986), P. campicolana (Walsingham, 1879), P. dangi (Razowski, 1997), P. decrepita (Razowski & Becker, 2002), P. dubia (Razowski & Becker, 1983). P.fusifera (Meyrick, 1912), P. galbanca (Meyrick, 1917), P. lavana (Busck, 1907), Platphalonidia luxata (Razowski & Becker, 1986), P. mendora (Clarke, 1968), P. ochraceana (Razowski, 1967), P. paranae (Razowski & Becker, 1983), P. plicana (Walsingham, 1884), and P. sublimis (Meyrick, 1917). We describe Platplialonia magdalenae Metzler & Albu, new species, from a series of specimens that were discovered nectaring on Centromcdia (=Hemizonia) pungens (Hook. & Arn.) Greene ssp. pungens (Asteraceae) during the day-time on 2 May 2011 in Tulare County, in the Central Valley of California. Adults and male and female genitalia of P. magdalenae are illustrated.
156
156 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
Journal of the Lepidopterists’ Society
67(3), 2013, 156–160
THE DESCRIPTION OF PLATPHALONIA MAGDALENAE (TORTRICIDAE, TORTRICINAE, EULIINI,
COCHYLINA) FOUND NECTARING DIURNALLY ON CENTROMADIA PUNGENS (ASTERACEAE) IN
THE CENTRAL VALLEY OF CALIFORNIA ALONG WITH A LIST OF SPECIES OF PLATPHALONIA
ERIC H. METZLER
Michigan State University, Adjunct Curator of Lepidoptera, Research Collaborator U.S.N.M. Natural History,
P.O. Box 45, Alamogordo, New Mexico 88311-0045, USA; email: metzlere@msu.edu
AND
VALERIU ALBU
23032 Oak Meadow Ln., Friant, California 93626, USA; email: valalbu@netptc.net
ABSTRACT.
Platphalonia Razowski, 2011 (Tortricidae, Tortricinae, Euliini, Cochylina) was proposed for Saphenista mystica
Razowski & Becker, 1983 (type species) and several species previously assigned to Platphalonidia Razowski, 1985. However, with
the exception of the type species, none of the other purported congeners have been listed. We formally transfer 16 species to Plat-
phalonia, resulting in the following new combinations: P. albertae (Razowski, 1997), P. assector (Razowski, 1967), P. californica (Ra-
zowski, 1986), P. campicolana (Walsingham, 1879), P. dangi (Razowski, 1997), P. decrepita (Razowski & Becker, 2002), P. dubia (Ra-
zowski & Becker, 1983). P. fusifera (Meyrick, 1912), P. galbanea (Meyrick, 1917), P. lavana (Busck, 1907), Platphalonidia luxata
(Razowski & Becker, 1986), P. mendora (Clarke, 1968), P. ochraceana (Razowski, 1967), P. paranae (Razowski & Becker, 1983), P.
plicana (Walsingham, 1884), and P. sublimis (Meyrick, 1917). We describe Platphalonia magdalenae Metzler & Albu, new species,
from a series of specimens that were discovered nectaring on Centromedia (=Hemizonia) pungens (Hook. & Arn.) Greene ssp. pun-
gens (Asteraceae) during the day-time on 2 May 2011 in Tulare County, in the Central Valley of California. Adults and male and
female genitalia of P. magdalenae are illustrated.
Additional key words:
Platphalonia magdalenae, Tulare County, California native plants, Platphalonidia, Phalonidia
Based on the type species of Platphalonidia Razowski,
1985 (i.e., Phalonia felix Walsingham, 1895), Razowski
(2011) synonymized that genus with Phalonidia Le
Marchand, 1933 (Tortricidae, Tortricinae, Euliini,
Cochylina). He concomitantly proposed Platphalonia
Razowski, 2011 for Saphenista mystica Razowski &
Becker, 1983 (type species) and the remaining species of
Platphalonidia (i.e., all species formerly included in
Platphalonida, except its type species). In his description
of Platphalonia, Razowski stated that 29 unnamed
species are transferrable to Platphalonia, but he did not
list them
In 2011 Valeriu Albu collected a series of a previously
unknown species of Cochylina found nectaring during
the day in the Central Valley of California. Razowski (in
litt. 2012) concurred that the new species from
California’s Central Valley belonged in the genus
Platphalonia. Because there was no list of North
American species of Platphalonia, there was no way to
properly diagnose the new taxon.
The purposes of this paper are to transfer 16 species
to Platphalonia and to describe a new species of
Platphalonia from California’s Central Valley.
MATERIALS AND METHODS
The classification Euliini, Cochylina comes from
Regier et al. (2012). Józef Razowski from the Polish
Academy of Sciences, Krakow, (in litt., 2012) provided a
list of species that he referred to Platphalonia. We
carefully compared published illustrations of male and
female genitalia of species from the list to published
illustrations of the type species, Platphalonia mystica.
We examined the male and female genitalia of all other
species listed in Platphalonidia in Brown (2005), and we
made similar examinations of all species described in
Platphalonidia subsequent to Brown (2005).
Valeriu Albu collected a series of 21 specimens of an
undescribed species of Platphalonia nectaring on
Centromadia (=Hemizonia) pungens (Hook & Arn.)
Greene ssp. pungens (common spikeweed) (Asteraceae)
during the day in Tulare County, California. The moths
were common on the flower blossoms. He sent the
specimens to Eric Metzler for study.
Identification was made by comparing the specimens
to published illustrations of adults and genitalia.
Photographs of adults were sent to Kevin Tuck (The
Natural History Museum, London) who opined that the
Tulare County species is different from P. plicana
(Walsingham, 1884), to which the male genitalia
structures bear some resemblance. Photographs of the
adults and male genitalia were sent to Józef Razowski
who confirmed that the species was undescribed.
Genitalia were dissected following procedures
outlined in Hardwick (1950), Lafontaine (2004), and
VOLUME 67, NUMBER 3
157
Pogue (2002). Abdomens were removed from the
moths, wetted in 95% ethanol, and soaked in 10% KOH
for 1 hour at 50°C. Genitalia were dissected in water,
dehydrated in 100% ethanol, cleared in oil of cloves,
rinsed in xylene, and slide mounted in Canada balsam.
The genitalia were stained with Chlorozol Black in water
and/or Safranin O in 95% ethanol. Many authorities (e.g.
Hardwick 1950, Lafontaine 2004, Pogue 2002)
presented techniques for dissection and examination of
genitalia. Not all authors agree on each technique and
each author offered unique ideas. Over time Metzler
assembled a collection of techniques from the several
sources, not all of which can be found in any one source,
yet when taken together the techniques provide an
overview that makes the task of dissection and slide
preparation easier for Metzler to accomplish.
Terminology for elements of wing pattern follows
Horak (1991). Terminology for morphology and genital
structures follows Horak (1991) and Razowski (1970,
2008). Forewing length was measured to the nearest 0.1
mm, from the base to the apex excluding fringe, using a
stereo-microscope with reticle.
Specimens cited in this paper are deposited in the
following collections:
VA Valeriu Albu, Friant, CA
MSU Albert J. Cook Arthropod Research Collection,
Department of Entomology, Michigan State
University, East Lansing, MI
UCB University of California Berkeley, Berkeley, CA
USNM US National Museum of Natural History (Smith-
sonian Institution), Washington, DC
RESULTS
We transfer 16 species to Platphalonia.
Platphalonia Razowski, 2011
Type species: Saphenista mystica Razowski &
Becker, 1983 by original designation.
Platphalonia albertae (Razowski, 1997)
[Platphalonidia] comb. n.
Platphalonia assector (Razowski, 1967) [Cochylis]
comb. n.
Platphalonia californica (Razowski, 1986)
[Platphalonidia] comb. n.
Platphalonia campicolana (Walsingham, 1879)
[Cochylis] comb. n.
Platphalonia dangi (Razowski, 1997)
[Platphalonidia] comb. n.
Platphalonia decrepita (Razowski & Becker, 2002)
[Platphalonidia] comb. n.
Platphalonia dubia (Razowski & Becker, 1983)
[Saphenista] comb. n.
Platphalonia fusifera (Meyrick, 1912) [Phtheochroa]
comb. n.
Platphalonia galbanea (Merick, 1917) [Phalonia]
comb. n.
Platphalonia lavana (Busck, 1907) [Phalonia] comb.
n.
Platphalonia luxata (Razowski & Becker, 1986)
[Platphalonidia] comb. n.
Platphalonia mendora (Clarke, 1968) [Cochylis]
comb. n.
Platphalonia ochraceana (Razowski, 1967)
[Cochylis] comb. n.
Platphalonia paranae (Razowski & Becker, 1983)
[Saphenista] comb. n.
Platphalonia plicana (Walsingham, 1884)
[Conchylis] comb. n.
Platphalonia sublimis (Meyrick, 1917) [Phalonia]
comb. n.
Platphalonia magdalenae Metzler & Albu,
new species
(Figs 1–6)
Diagnosis. Platphalonia magdalenae is a small tan
moth with a pale reddish-brown transverse fascia. The
diagnostic features are the tan ground color, the slightly
reflective nature of the fore wing scales, the curved
reddish-brown postmedial fascia that extends from the
inner margin to the costa, and the brownish-gray hind
wings. As is typical with many species of Cochylina, the
generic assignment of P. magdalenae cannot be
determined from the external appearance of the adult
moth. Superficially, P. magdalenae might be confused
with several species, in several genera, of Cochylina. An
examination of the male genitalia is required for positive
assignment to a genus. Within the genus Platphalonia,
the only other described species from California is P.
campicolana (Walsingham, 1879). The fore wing of P.
campicolana is cream-white, the reddish fascia extends
from the inner margin to mid-wing, the postmedian
fascia does not reach the costa, and the fringe is black.
The fore wing of P. magdalenae is tan and the fringe is
not black. Platphalonia albertae, P. californica, P. dangi,
P. lavana, P. luxata, P. parvimaculana, and P. plicana,
also occur in the Nearctic Ecozone. The width of the
mesal process of the transtilla narrows subapically in P.
albertae; whereas in P. magdalenae it is not narrowed
subapically. The acutely pointed apices of the bifurcate
terminus of the mesal process of the transtilla are
distinctly pointed laterally in P. californica; whereas in P.
magdalenae are not pointed laterally. The fore wing of P.
dangi is dirty-white with an oblique dark colored fascia
extending from just before the apex to outer margin 2/3
from the tornus; the fore wing of P. magdalenae is tan
without an oblique dark fascia. The ground color of the
fore wing of P. lavana is dirty-white, the fore wing has an
excurved subterminal gray fascia extending from just
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FIGS. 1–5. Platphalonia adults and genitalia. 1, P. magdalenae male holotype. 2, P. magdalenae female paratype. 3, P. magdalenae
male paratype genitalia. 4, P. magdalenae male paratype aedeagus. 5, P. magdalenae female paratype genitalia.
VOLUME 67, NUMBER 3
159
before the apex to the tornus, and the fringe is black; in
contrast, the ground color of the fore wing of P.
magdalenae is tan without a subterminal fascia, and the
fringe is concolorous. The ductus bursae of P. luxata is
ringed with sclerotization at the junction with the
corpus bursae; the ductus bursae of P. magdalenae is
encircled with linear rugosities at the junction with the
corpus bursae. The ground color of the fore wing of P.
plicana is dirty-white, the subterminal area contains a
patch of contrasting dark-brown scales, the terminal line
is black, the fringe is contrastingly dark brown, and the
hind wing is gray; the fore wing of P. magdalenae is tan,
there is no patch of dark-brown scales in the
subterminal area, the terminal line and fringe are not
contrastingly dark, and the hind wing is dark fuscous.
Description. Adult male (Fig. 1). Head: Front smooth, tan, with
scattered tan-tipped orange scales; vertex pale-tan, with scattered
dark-tan scales, smooth, posterior margin with scales erect. Labial
palpus porrect, segments 1 and 2 mixed with tan and chamois-colored
scales, lateral and mesal surfaces with scales appressed, dorsal and
ventral surfaces weakly tufted, segment 3 directed anteriorly, with
light tan, appressed scales. Antenna with dorsal surface scaled, tan,
ventral surface naked with setae numerous, length = width of antennal
segment. Thorax: Dorsum tan and chamois-color, smoothly scaled;
underside with scales appressed, shining white and silver. Fore-leg
femur with scales appressed, dorsal surface tan, with scattered brown
and orange scales, ventral surface dirty-white, scales of tibia, femur
and tarsi appressed, dorsal surface tan, scattered brown scales, ventral
surface dirty-white. Mid-leg scales appressed, femur dirty-white, tibia,
femur, and tarsi dorsal surface tan, scattered light-brown scales,
ventral surface dirty-white. Hind-leg scales appressed, dirty-white.
Forewing length 3.95.5 mm, mean 4.7 mm, n = 18. Upper surface of
forewing reflective white (appears tan), anterior one half of basal 1/3
narrowly salmon, basal 1/3 of costa dark brown, distal 2/3 of costa tan;
posterior one half of basal area chamois color; subbasal fascia angled
outward from inner margin, chamois color; postmedial fascia angled
outward from inner margin, bent at cell towards costa, pale horn color,
at costa mixed with brown and salmon, brown scales at bend;
subterminal fascia angled from cell to just before apex, pale horn
color, subterminal area below apex with small patch of brown scales;
fringe pale horn color mixed with salmon and brown scales; underside
inner margin reflective white scales, area over cell brown-tipped
reflective white scales, basal one half of costa brown, outer one half
salmon; fringe reflective white. Hindwing with tan-gray tipped
reflective white scales, base lighter, inner angle darker, fringe
concolorous, reflective; underside reflective white, tan-gray tipped
scales numerous, inconspicuous; fringe concolorous. Abdomen:
Dorsum scales appressed, reflective silver-tan; underside scales
appressed, mixed tan and pale gray. Genitalia (Fig. 3) with tegumen
wide, prominent, apex flattened, suture mesially; uncus absent; socii
appressed, membranous, broad, lightly sclerotized apically; transtilla
robust, directed ventrally, mesally bent, mesal process drawn out, apex
robust, bifurcate, each arm drawn to a blunt point, apex with tiny
teeth laterally; juxta broad, elliptical; saccus obscure; valva simple,
elongate, curved dorsally, with scattered setae apically, costa
sclerotized; saccular region not produced, weakly sclerotized, with
scattered setae; phallus (Fig. 4) longer than wide, gradually narrowing
apically, bent 30° at 1/3 length; cornuti poorly defined, obscure, lightly
sclerotized.
Adult female (Fig. 2). Head and thorax: Essential as described for
male. Forewing length 5.05.3 mm, mean 5.1 mm, n = 3. Hindwing
with frenulum of one, two, or three bristles, asymmetrical. Abdomen:
Genitalia (Fig. 5) with papilla analis membranous, narrow, setose;
posterior apophysis extending to anterior margin of 8th segment;
anterior apophysis divided basally, extending to anterior end of ductus
bursae; antrum weakly sclerotized; ductus bursae lightly sclerotized
posteriorly, mesally membranous, anteriorly sclerotized, fluted and
widening at junction with corpus bursae; corpus bursae short, round,
membranous, densely covered with spicules in patches; signa absent.
Holotype. Adult male, pinned with labels as follows: “Legit Val
Albu, Tulare Co/California, Rt. 43 @ Allensworth, 2 V 2011"
FIGS. 6–8. Distribution map and habitat of Platphalonia magdalenae. 6, Distribution map for P. magdalenae. 7, Habitat picture
of type locality of P. magdalenae. 8, Centromadia pungens ssp. pungens.
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160 JOURNAL OF THE LEPIDOPTERISTS’ SOCIETY
“HOLOTYPE USNM Platphalonia magdalenae Metzler & Albu” [red
handwritten label] (USNM).
Paratypes. 20 m,3 f: same data as Holotype. Depositories: VA,
MSU, UCB, USNM.
Systematics. This new species is placed in the genus Platphalonia
based on comparison of the female genitalia with the holotype of
Saphenista mystica Razowski & Becker, 1983, which is the type
species of Platphalonia. Our analysis was confirmed by Józef Razowski
(in litt., 2012).
Etymology. The species is named after Magdalena Albu, who, as
the wife of the second author, lovingly and dutifully followed and
helped with his entomological excesses. The name is in the genitive
case.
Distribution and biology. The diurnal nectaring
behavior of adult Cochylina is unusual, and we call
attention to it here with the idea that the behavior could
be present, and as yet unobserved, in other species of
Cochylina.
Platphalonia magdalenae occurs in the Central Valley
of California. Its distribution outside Tulare County is
unknown. Nothing is known about its life history,
however, an association with Centromadia pungens ssp.
pungens is reasonably inferred.
The adults of P. magdalenae were collected in early
May. The habitat is in the intensively farmed Central
Valley, along California State Route 43, in the vicinity of
Allensworth, CA at 60 m elevation (Fig. 7). The plant
upon which the adults were found, Centromadia
pungens ssp. pungens (Fig. 8), is found in the U.S. from
Washington south through Oregon, California, Nevada,
and southern Arizona (USDA 2012). The plant is not
recorded from Canada, and the Mexican distribution is
unknown. The plant’s distribution in California
(Baldwin et al. 2012) is extensive at lower elevations,
from the Sierra Nevada foothills to the coast. It is not
recorded from the eastern deserts of the state. In Tulare
County it occurs abundantly in disturbed areas, along
roads and rail road tracks. It has a long blooming period,
from April to September.
Remarks. Some specimens are slightly darker. They
have scattered dark-brown scales on the head, the palpi,
and an increased number of dark-brown scales on the
legs.
DISCUSSION
Our study examined the published illustrations of the
genitalia of the 24 species included in Platphalonidia
(Brown 2005), the list of species provided by Józef
Razowski (in litt. 2012), and all species described in the
genus Platphalonidia subsequent to Brown (2005) We
were able to confirm the congeneric status of 16 species
which we transferred to Platphalonia. We transferred
species where the genitalia were in agreement with the
type species Platphalonia mystica. Razowski (2011)
indicated that 29 species are referable to Platphalonia.
This paper is not a revision of the genus Platphalonia,
and we do not resolve the correct generic assignments
of the remaining 13 species because such an endeavor
falls outside the scope of this paper.
The many species of Cochylina, even within a single
genus, e.g. Aethes Billberg, 1820, often have forewing
patterns that are incongruous. Several references (e.g.
Horak 1991, Razowski 1970, 2008) illustrate the variety
of forewing patterns of Cochylina. We reference Horak
(1991) in this paper because her illustration on page 6
has wing pattern elements, sub-basal and postmedial
fascia, represented in P. magdalenae.
ACKNOWLEDGMENTS
We thank Joanna Clines, Forest Botanist at the Sierra
National Forest in North Fork, CA for identifying the plant
upon which the moths were found nectaring and for providing
useful information about it. We especially thank Kevin Tuck of
the Natural History Museum in London and Józef Razowski of
the Polish Academy of Sciences in Kraków for their rapid and
detailed replies to our requests for information. We appreciate
the assistance of Jerry A. Powell and John W. Brown with advice
on how to broach the taxonomy of Platphalonia. We thank Jason
J. Dombroskie and two anonymous reviewers for reading the
paper and offering constructive suggestions.
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BROWN, J. W. 2005. World Catalogue of Insects Volume 5 Tortricidae
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HARDWICK, D. F. 1950. Preparation of slide mounts of lepidopterous
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HORAK, M. 1991. Chapter 1 Morphology, phylogeny and systematics,
1.1 Morphology. pp. 122. In Van Der Geest, L. P. S. & H. H.
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LAFONTAINE, J. D. 2004. The Moths of North America Including
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POGUE, M. G. 2002. A World revision of the genus Spodoptera
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RAZOWSKI, J. 1970. Microlepidoptera Palaearctica dritter Band
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REGIER, J. C., J. W. BROWN, C. MITTER, J. BAIXERAS, S. CHO, M. P.
CUMMINGS, & A. ZWICK. 2012. A molecular phylogeny for the
leaf-roller moths (Lepidoptera: Tortricidae) and its implications
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USDA. 2012. United States Department of Agriculture Natural Re-
sources Conservation Service Plants Profile. Available from:
http://plants.usda.gov/java/profile?symbol=CEPUP6 (Accessed
19 April 2012).
Submitted for publication on 11 June 2012; revised and ac-
cepted 13 September 2012.
... The new genera recognized by were identified as "Cochylini new genus 1," "Cochylini new genus 2," and so forth in order to provide a more meaningful taxonomic framework for the fauna, i.e., this action seemed a better alternative to leaving the associated species "unplaced." Taxonomic changes proposed by Razowski (2009Razowski ( , 2011, Metzler & Albu (2013), and others over the last decade resulted in the generic reassignments of many species. Many of the changes proposed by were formalized through various publications (e.g., Pogue 2001, Brown 2005, and new species were added to the faunal list through introduction (e.g., Agapeta zoegana) (Powell et al. 2000) or description of the native fauna (e.g., Metzler & Forbes 2012, Brown 2013, Metzler & Albu 2013. ...
... Taxonomic changes proposed by Razowski (2009Razowski ( , 2011, Metzler & Albu (2013), and others over the last decade resulted in the generic reassignments of many species. Many of the changes proposed by were formalized through various publications (e.g., Pogue 2001, Brown 2005, and new species were added to the faunal list through introduction (e.g., Agapeta zoegana) (Powell et al. 2000) or description of the native fauna (e.g., Metzler & Forbes 2012, Brown 2013, Metzler & Albu 2013. In a phylogenetic analysis of Tortricidae based entirely on molecular data, Regier et al. (2012) revealed that Cochylini are a monophyletic lineage embedded within the larger Euliini and should be treated as the subtribe Cochylina until further evidence contradicts this proposed change in classification. ...
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Background: Tortricidae, one of the largest families of microlepidopterans, comprise about 10,000 described species worldwide, including important pests, biological control agents and experimental models. Understanding of tortricid phylogeny, the basis for a predictive classification, is currently provisional. We present the first detailed molecular estimate of relationships across the tribes and subfamilies of Tortricidae, assess its concordance with previous morphological evidence, and re-examine postulated evolutionary trends in host plant use and biogeography. Methodology/principal findings: We sequenced up to five nuclear genes (6,633 bp) in each of 52 tortricids spanning all three subfamilies and 19 of the 22 tribes, plus up to 14 additional genes, for a total of 14,826 bp, in 29 of those taxa plus all 14 outgroup taxa. Maximum likelihood analyses yield trees that, within Tortricidae, differ little among data sets and character treatments and are nearly always strongly supported at all levels of divergence. Support for several nodes was greatly increased by the additional 14 genes sequenced in just 29 of 52 tortricids, with no evidence of phylogenetic artifacts from deliberately incomplete gene sampling. There is strong support for the monophyly of Tortricinae and of Olethreutinae, and for grouping of these to the exclusion of Chlidanotinae. Relationships among tribes are robustly resolved in Tortricinae and mostly so in Olethreutinae. Feeding habit (internal versus external) is strongly conserved on the phylogeny. Within Tortricinae, a clade characterized by eggs being deposited in large clusters, in contrast to singly or in small batches, has markedly elevated incidence of polyphagous species. The five earliest-branching tortricid lineages are all species-poor tribes with mainly southern/tropical distributions, consistent with a hypothesized Gondwanan origin for the family. Conclusions/significance: We present the first robustly supported phylogeny for Tortricidae, and a revised classification in which all of the sampled tribes are now monophyletic.
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Diagnoses of the 68 described genera of Cochylini are provided along with redescriptions and comments where necessary. Platphalonidia is synonymized with Phalonidia, and Platphalonia Razowski, gen. n. is described.
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One of the most essential procedures in the identification of many Lepidoptera is an examination of the genitalia, and for a most critical examination of these organs slide mounts must be prepared. In many cases, the preparation of slides is one of the most neglected arts in lepidopterology. This is to be deplored, for a well-prepared slide is not only handsome but also much more valuable taxonomically than a messily prepared one. For large and medium-sized specimens, I have found the following technique very gratifying in producing slides of uniformly high quality. Certain phases of the technique are not particularly well adapted for working with smaller moths.
World Catalogue of Insects Volume 5 Tortricidae (Lepidoptera) Apollo Books. Stenstrup, Denmark. 741 pp. HARDWICK, D. F. 1950. Preparation of slide mounts of lepidopterous genitalia
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Chapter 1 Morphology, phylogeny and systematics, 1.1 Morphology
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LAFONTAINE, J. D. 2004. The Moths of North America Including Greenland, Fascicle 27.1, Noctuoidea Noctuidae (part) Noctuinae (part -Agrotini). The Wedge Entomological Research Foundation, Washington, DC. 385 pp.
Microlepidoptera Palaearctica dritter Band Cochylidae
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RAZOWSKI, J. 1970. Microlepidoptera Palaearctica dritter Band Cochylidae. Verlag Georg Fromme & Co. Wien, Germany. 2 volumes. 528 pp. ----. 2008. Tortricidae of the Palaearctic Region Volume 1. Frantisek Slamka, Bratislava, Slovakia. 152 pp. ----. 2011. Diagnoses and remarks on genera of Tortricidae, 2: Cochylini (Lepidoptera: Tortricidae). SHILAP Revta. Lepid. 39:397-414.