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The Phylogenetic Relationships of Chalcosiinae (Lepidoptera Zygaenoidea Zygaenidae)_SHEN-HORN YEN

The Phylogenetic Relationships of Chalcosiinae (Lepidoptera Zygaenoidea Zygaenidae)_SHEN-HORN YEN

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 Zoological Journal of the Linnean Society
, 2005,
143
, 161–341. With 71 figures © 2005 The Linnean Society of London,
 Zoological Journal of the Linnean Society,
2005,
143
, 161–341
161
Blackwell Science, Ltd
Oxford, UK 
ZOJZoological Journal of the Linnean Society
0024-4082The Lin-nean Society of London, 2005? 2005
143
2
161341Original Article
PHYLOGENY OF CHALCOSIINAE S.-H.YENET AL.
*Corresponding author. Current address: Department of Biological Sciences, National Sun Yat-Sen University, 70Lien-hai Rd, Kaohsiung 804, Taiwan. E-mail:shenhornyen@mail.nsysu.edu.tw
The phylogenetic relationships of Chalcosiinae(Lepidoptera, Zygaenoidea, Zygaenidae)
SHEN-HORN YEN
1
*, GADEN S. ROBINSON
2
and DONALD L. J. QUICKE
1,2
1
 Division of Biological Sciences and Centre for Population Biology, Imperial College London, Silwood Park Campus, Ascot, Berkshire, SL5 7PY, UK 
2
 Department of Entomology, The Natural History Museum, London SW7 5BD, UK 
 Received April 2003; accepted for publication June 2004
The chalcosiine zygaenid moths constitute one of the most striking groups within the lower-ditrysian Lepidoptera,with highly diverse mimetic patterns, chemical defence systems, scent organs, copulatory mechanisms, hostplant uti-lization and diapause biology, plus a very disjunctive biogeographical pattern. In this paper we focus on the genus-level phylogenetics of this subfamily. A cladistic study was performed using 414 morphological and biochemical char-acters obtained from 411 species belonging to 186 species-groups of 73 genera plus 21 outgroups. Phylogenetic anal-ysis using maximum parsimony leads to the following conclusions: (1) neither the current concept of Zygaenidae northat of Chalcosiinae is monophyletic; (2) the previously proposed sister-group relationship of Zygaeninae
+
Chal-cosiinae is rejected in favour of the relationship (Zygaeninae
+
((Callizygaeninae
+
 
Cleoda
)
+
(
 Heteropan
 
+
Chalcosi-inae))); (3) except for the monobasic Aglaopini, none of the tribes
 sensu
Alberti (1954) is monophyletic; (4) chalcosiinesynapomorphies include structures of the chemical defence system, scent organs of adults and of the apodemal systemof the male genitalia. A paired metathoracic androconial organ and a series of abdominal tergal corematal organshave been discovered, both being new to Lepidoptera. Due to highly homoplastic patterns in copulatory structuresand wings that demonstrate significant sexual dimorphism, polymorphism and mimicry, 17 of the 69 ‘true’ chalcosiinegenera (
c
. 25%) are shown to be either paraphyletic or polyphyletic. The present classification is therefore very mis-leading. Reductions of various parts of the male genitalia in some groups are accompanied by morphological and func-tional replacement involving the 8th abdominal segment. A prominent but convergent lock and key mechanism isrevealed.© 2005 The Linnean Society of London,
 Zoological Journal of the Linnean Society
, 2005,
143
, 161–341.
 ADDITIONAL KEYWORDS:
 
chaetosoma – chemical defence – constraint analysis – cyanogenesis – genitalicreduction – mimicry.
INTRODUCTION
H
ISTORICAL
 
REVIEW
 
OF
 
HIGHER
 
CLASSIFICATION
 
OF
Z
 YGAENOIDEA
Like many other superfamilies within the ditrysianLepidoptera, the Zygaenoidea are not characterized byclearly defined autapomorphies (Epstein
 et al
., 1999).The taxon appears at first glance to have become awastebasket, including a number of family groupsthat may individually be well defined, but do littlemore than share a number, albeit numerous, of plesi-omorphic characters. Its composition has been very farfrom stable, including from seven (Scoble, 1992) to 13families (Epstein
 et al
., 1999), and its monophyly isstill highly debatable (Fig. 1).Dyar (1894) brought Limacodidae (as Eucleidae),Megalopygidae, Zygaenidae and Procridinae (asPyromorphidae) into an assemblage called the ‘Anth-rocerina’ based on larval chaetotaxy, which wassubsequently combined with ‘Cossina’ within thesuperfamily ‘Tineides’ (Dyar, 1896). Chapman (1893,1894) lumped the Zygaenidae with Limacodidae (asCochliopodidae) and Micropterigidae into the pupalgroup ‘Incompletae’, although these views were notaccepted by most authors at that time (Dyar & Mor-ton, 1895; Packard, 1895, Hinton, 1955).In 1895, Packard suggested a close relationshipbetween Limacodidae and Megalopygidae, but heplaced them under Tineina, and thus apart from the
 
162
S.-H. YEN
 ET AL.
 © 2005 The Linnean Society of London,
 Zoological Journal of the Linnean Society,
2005,
143
, 161–341
Zygaenidae. The term ‘Zygaenoidea’ was first intro-duced by Fracker (1915), who included within itChalcosiinae (as Chalcodidae [sic]), Procridinae (asPyromorphidae), Epipyropidae, Dalceridae, Megalopy-gidae, and Limacodidae. This classification has beenexpanded by modern workers to include all of the cur-rent family groups of Zygaenoidea (Common, 1975,1990; Kuznetzov & Stekolnikov, 1981; Minet, 1986;Scoble, 1992).Except for Zygaenidae, Brock (1971) placed mostfamilies previously assigned to Zygaenoidea in theCossoidea, based primarily on characters of the adultthorax and forewings. He also transferred Epipyropi-dae and Heterogynidae to the Tineoidea. Heppner
Figure 1.
Schematic classifications and phylogenetic concepts of Zygaenoidea and Zygaenidae proposed by differ-ent authors. In each of the hypotheses, members of Zygaenidae are in bold and non-zygaenoid groups are initalic. A, Alberti (1954). B, Minet (1986, 1991, 1994). C, Common (1970) and Nielsen & Common (1991). D, Scoble(1992). E, Epstein (1996). F, Heppner (1998). G, Fänger
 et al
.
1999). H, Epstein
 et al
. (1999). I, Holloway
 et al
.
2001).
ZygaeninaePhaudinaeCharideinaeCallizygaeniniProcridiniHimantopterinaeAnomoeotinae
A
Chalcosiinae
MegalopygidaeLimacodidaeChrysopolomidaeEpipyropidaeCyclotornidaeHeterogynidae
Lactura 
-groupPhaudinaeAnomoeotinaeHimantopterinaeCharideinaeProcridinaeZygaeninae
B
Chalcosiinae
Charideinae (Thyrididae) 
EpipyropidaeCyclotornidaeDalceridaeLimacodidaeChrysopolomidaeMegalopygidaeAnomoeotidaeHimantopteridaeHeterogynidae
ProcridinaeZygaeninae
C
Chalcosiinae
DalceridaeEpipyropidaeCyclotornidaeMegalopygidaeLimacodidaeChrysopolomidae
ProcridinaeZygaeninaeCharideinaePhaudinaeHimantopterinaeAnomoeotinae
D
Chalcosiinae
SomabrachyidaeCyclotornidaeEpipyropidaeMegalopyginaeTrosiinaeAididaeDalceridaePant/CrothChrysopolominaeLimacodinae
Zygaenidae
E
Bombycina Tortricoidea Castnioidea Other Cossoids Cyclotornidae Epipyropidae Dalceridae Chrysopolomidae Limacodidae Tineina Sesioidea 
SomabrachyidaeMegalopygidaeHeterogynidaeLacturidaeHimantopteridae
AnomoeotinaePhaudinaeProcridinaeZygaeninae
F
ChalcosiinaePhaudinae
MegalopygidaeSomabrachyidaeAnomoeotidaeHimantopteridaeEpipyropidaeCyclotornidaeAididaeLimacodidaeDalceridaeLacturidaeHeterogynidae
CallizygaeninaeProcridinaeZygaeninae
G
Chalcosiinae
MegalopygidaeSomabrachyidaeAnomoeotidaeHimantopteridaeEpipyropidaeCyclotornidaeAididaeLimacodidaeDalceridaeLacturidaeHeterogynidae
PhaudinaeCallizygaeninaeProcridinaeZygaeninae
H
ChalcosiinaeMegalopygidaeAididaeLimacodidaeDalceridaeEpipyropidaeHimantopteridaeLacturidaePhaudinaeProcridinaeCallizygaeninaeZygaenidae
I
Chalcosiinae
 
PHYLOGENY OF CHALCOSIINAE
163
 © 2005 The Linnean Society of London,
 Zoological Journal of the Linnean Society,
2005,
143
, 161–341
(1984, 1992, 1998) retained Megalopygidae, Som-abrachyidae, and Heterogynidae in Zygaenoidea, butplaced Epipyropidae, Cyclotornidae, and the remain-der of the limacodid group (see below) in the Cos-soidea. Brock’s Cossoidea, except for the position of Epipyropidae and Heterogynidae, has been followedby Fletcher & Nye (1982). Brock did not provide jus-tification for this arrangement. This classification hasbeen criticized, however, both for the weakness of theadult characters involved and because of the strengthof the immature stage characters as evidence for themonophyly of Zygaenoidea in its earlier sense (Com-mon, 1975; Kuznetzov & Stekolnikov, 1981).Minet (1986, following Common, 1975), proposedtwo potential autapomorphies of Zygaenoidea: theretractile head of the larva (at least in the laterinstars); and the position of the second abdominal spi-racle of the pupa, which is covered by the wings. Thelatter character has been verified for Epipyropidae,Megalopygidae (including Somabrachyidae), Limaco-didae, Heterogynidae, Zygaenidae (Epstein
 et al
.,1999), Cyclotornidae, Himantopteridae and Zyga-enidae (Fänger, Yen & Naumann, 1999). Although ithas been argued that both characters may easily havedeveloped independently (Heppner, 1998), Epstein
 et al
. (1999) argued that they can be used as charac-ters to define the superfamily.The first cladistic study of the superfamily was car-ried out by Epstein (1996) who defined the monophylyof the limacodid group, viz. Limacodidae, Megalopy-gidae, Somabrachyidae, Aididae and Dalceridae,based on a number of apparently synapomorphic char-acters of both adult and immature stages, includingthe presence of additional prolegs or crochets on A2and A7, a sculptured eye flange in pupae, dense sen-silla trichodea on all legs of adult females and theabsence of ocelli. As a result of this study, two majorlineages, the limacodid families, and Zygaenidae
 sensu lato
, are tentatively recognized within the Zyg-aenoidea
.
Fänger
 et al
. (1999) attempted a morpholog-ical survey of several family groups which had beengiven very little attention in previous studies, and pro-vided a preliminary phylogenetic structure of thesuperfamily. However, the inter-relationships of thebasal zygaenoids, ant-associated Cyclotornidae andHomoptera–parasitic Epipyropidae are still unclear,due to insufficient information on immature stagesand uninformative adult characters.
T
HE
 
FAMILY
Z
 YGAENIDAE
 
IN
 
FLUX
The Zygaenidae, colloquially Burnet moths (Zygaeni-nae) and Forester moths (European Procridinae), isone of the largest families within the Zygaenoidea,estimated to include
c.
1200 species worldwide (Bryk,1936, Epstein
 et al
., 1999). According to NaumannTarmann & Tremewan (1999) and Epstein
 et al
.(1999), the current concept of zygaenids is not sup-ported by general morphological features but ratherby their ability to synthesize two cyanogenic com-pounds from the amino acids valine and isoleucine(Witthohn & Naumann, 1984a, b, 1987a, b; Epstein
 et al
., 1999; Naumann
 et al
., 1999).The chemical defence ecology of Zygaenidae wasfirst reported by Jones
 et al
. (1962) who demonstratedthat HCN is released from crushed tissues of allinstars of 
 Zygaena filipendulae
, and that the highestconcentrations of HCN precursors are found in theeggs. Subsequently, various studies on the biochemicalmechanisms, interactions amongst hostplants, mothsand their parasitoids, and the relevant morphologicalstructures of several representative genera of Zyga-enidae were conducted by a number of research teams.The chemical source of HCN in Burnet mothsremained unknown until Davis & Nahrstedt (1979)demonstrated that it is derived from two cyanogluco-sides, linamarin and lotaustralin. Linamarin andlotaustralin have long been known to occur in a num-ber of plant families, e.g. Fabaceae, widely utilized asa host plant by the species of 
 Zygaena
(see Figs 5, 6).However, the fact that some Zygaeninae live on acya-nogenic host plants, but remain cyanogenic, suggeststhat these insects are capable of synthesizing thesecompounds
de novo
. Although cyanogenesis in the surveyed zygaenidspecies has been considered to be autapomorphic forthis family (e.g. Naumann
 et al
., 1999), a similar chem-ical defence mechanism is known to exist in its poten-tial sister group, the Heterogynidae (Zilli, 1987; Zilli &Racheli, 1989; Epstein
 et al
., 1999; Fänger & Nau-mann, 2001), and the current taxonomic compositionof the Zygaenidae remains problematic (Yen, 2003c).The consistency of apomorphies recognized in previousstudies has yet to be tested in a comprehensive study.The chaotic taxonomic history of Zygaenidae hasinvolved most family groups within the superfamily,as well as various non-zygaenoid groups. When estab-lished by Latreille (1809) as Zygaenides, only the well-known western Palaearctic genus
 Zygaena
Fabricius,1775 was included in Zygaenidae. Subsequently, thefollowing groups were included in the family, and itsconcept varied between different authors:
 Heterogynis
(in Zygaenides by Walker, 1854), Syntominae and Cte-nuchini of Arctiidae (by Walker, 1854), ProcridinaeBoisduval, 1828 (as Procridae
=
Pyromorphina Her-rich-Shaffer, 1855), Chalcosiinae Walker, 1864 [1865](as Chalcosiidae), Charideinae Butler, 1876 (as Chari-deinae in Arctiidae
=
Glaucopidae Harris, 1839,Pompostolinae Jordan, 1907), HimantopteridaeRogenhofer, 1884, Phaudinae Kirby, 1892 and Ano-moeotidae Hering, 1937. There were, in addition, twolittle-known groups,
 Lactura
-group (
 sensu
Kyrki,

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