Overview

Brief Summary

Introduction

Introduction:

The Epicopeiidae comprise about 25 species in nine genera (Minet and Scoble 1999). They are found only in the Palearctic and Oriental parts of Asia.  The adults are relatively large and colorful moths which are all day-flying, although some may also come to U.V. lights at night (Minet 2002). They rest with the wings spread out and appressed to the substrate. Many appear to mimic other day-flying Lepidoptera. Some species with tailed wings, such as Epicopeia hainesii, resemble swallowtail butterflies. The larvae, still poorly known, appear to be external leaf feeders on woody angiosperms in several distantly related families. They produce waxy secretions which are incorporated into the cocoon and may also coat the pupa (Jaros and Spitzer 1999; Holloway et al. 2001). Pupation occurs in soil or leaf litter. Some excellent images of live epicopeiid adults and larvae can be found at jpmoth.org
 
 Epicopeia hainesii

  Epicopeia hainesii

 
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Diversity

Diversity description:

About 25 species in nine genera (Minet and Scoble 1998).

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Distribution

Geographical Distribution

Geographic Range:

Palearctic, Oriental

Geographic Range description:

Known only from palearctic and tropical Asia (Minet and Scoble 1998).

Geographic Distribution:

Known only from palearctic and tropical Asia (Minet and Scoble 1998).

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Physical Description

Morphology

Egg morphology

Orientation:

upright

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Larvae Morphology

Larval head description:

mandible with large, flat lateral area, the ventral edge of which is more or less projecting (Minet and Scoble 1998)

Larval body description:

mature larva with white waxy secretions, often entirely covering skin (Minet and Scoble 1998)

Thoracic legs:

present

Pairs of thoracic legs:

from 3

Larval Prothoracic L-group setae:

bisetose

Abdominal prolegs:

present

Pairs of abdominal legs:

from 5

Proleg configuration:

normal

Proleg size:

long

Crochets:

multiserial

Crochet arrangement description:

curved, bi- to multiordinal mesoseries (Minet and Scoble 1998)

Anal comb on A10:

absent

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Pupa/Cocoon morphology

Pupa type:

obtect

Pupa description:

often waxy. (Minet and Scoble 1998)

Cremaster:

present

Number of rows of tergal spines:

from 8

Cocoon:

present

Cocoon description:

Waxy, either spun on or between leaves, subterranean, or hidden in leaf litter. (Minet and Scoble 1998)

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Adult Abdomen Morphology

Reproductive system:

Ditrysian

Oviscapt (ovipositor):

non-piercing

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Adult Thorax Morphology

Epiphysis:

present

Forelegs:

normal

Number of tibial spurs foreleg:

from 0

Number of tibial spurs midleg:

from 2

Number of tibial spurs hindleg:

from 4

Leg description:

Arolium present, claws simple (without a medial tooth). Spinarea absent (Minet and Scoble 1998).

Wing venation:

heteroneurous

Wing coupling:

with frenulum, absent

Wing coupling description:

Frenulum and subcostal retinaculum usually present in males; female frenulum absent or strongly reduced.

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Tympanum Morphology

Counter-tympanum:

absent

Abdomen tympanum:

absent

Thorax tympanum:

absent

Palp tympanum:

absent

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Adult Head Morphology

Ocelli:

absent

Eyes:

smooth, hairy

Labial palpus:

porrect, upcurved

Number of labial palp segments:

from 3

Labial palpus modification:

none

Maxillary palpus:

minute

Proboscis:

present

Proboscis description:

"Well developed, with a broad base" (Minet and Scoble 1998)

Mandibles:

absent

Antennal sensillum:

Antennal sensillum present

General antennae description:

"filiform, lamellate, serrate or bipectinate" (Minet and Scoble 1998).

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Diagnostic Description

Synapomorphies

Apomorphies:

Minet 2002: (1) Compound eyes reduced, their posterior margins never really extending posteriorly beyond the posterior parts of the antennal sockets.  (2)Posterior part of the ocular ridge unpigmented (unlike the anterior part).  (3) Head without ocelli.  (4) Proximal outer surface of the labial palpus clothed with yellow scales; inferred ancestral state, replaced by other colors in some lineages.  (5) Ventral process of tegula ending in a fairly sharp point.  (6) In mesothorax, basisternal mesal suture and its ridge not exactly reaching the anterodorsal margin of the basisternum.  (7) In metathorax, furco-epimeral band very broad, primarily much broader than the coxal membrane lying below it.  (8) Metacoxa with merocosta ending, ventrally, close to the sagittal plane and well above the coxotrochanteral articulation.  (9) Pretarsal claw without medial tooth.  (10) Forewing without an areole.  (11) Forewing vein Rs4 stalked with M1 and separated from Rs1 + 2 + 3 (inferred groundplan condition).  (12) spinarea absent

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Life History and Behavior

Behavior

Adult Behavior

Adult behavior:

diurnal

Adult behavior:

At rest, wings are outspread, closely appressed to substrate. Day flying, often mimetic. Minet and Scoble (1998).

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Reproduction

Life History: Immature Stages

Larval food items include:

Clethraceae.  Cornaceae.  Ericaceae.  Theaceae.  Ulmaceae

Larval food habits description:

Known larvae feed externally on plants of the families listed above.

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Evolution and Systematics

Evolution

Phylogenetic Relationships

Discussion of Phylogenetic Relationships:

Monophyly for Epicopeiidae seems secure. Minet (2002) cited 12 synapomorphies of the adult moth in support of the family as currently constituted:

 

(1) Compound eyes reduced, their posterior margins never really extending posteriorly beyond the posterior parts of the antennal sockets. (2)Posterior part of the ocular ridge unpigmented (unlike the anterior part). (3) Head without ocelli. (4) Proximal outer surface of the labial palpus clothed with yellow scales; inferred ancestral state, replaced by other colors in some lineages. (5) Ventral process of tegula ending in a fairly sharp point. (6) In mesothorax, basisternal mesal suture and its ridge not exactly reaching the anterodorsal margin of the basisternum. (7) In metathorax, furco-epimeral band very broad, primarily much broader than the coxal membrane lying below it. (8) Metacoxa with merocosta ending, ventrally, close to the sagittal plane and well above the coxotrochanteral articulation. (9) Pretarsal claw without medial tooth. (10) Forewing without an areole. (11) Forewing vein Rs4 stalked with M1 and separated from Rs1 + 2 + 3 (inferred groundplan condition). (12) spinarea absent.

 

Minet (1991) regarded Epicopeiidae as sister group to the Drepanidae (see also Minet and Scoble 1999, Minet 2002). Minet (2002) proposed the following synapomorphies for Drepanoidea comprising these two families: (a)imaginal abdomen base with complete or nearly complete tergosternal sclerites;  (b) imaginal abdomen base with neotergite distinctly fused with marginotergites; (c) larval mandible with a large, flat, lateral area, which is delimited ventrally by a well defined carina;  (d) most larval abdominal segments with at least one secondary seta cephalad of L3; (e) pupa never exhibiting conspicuous portion of the fore femur.  Several of these apomorphies were regarded as tentative.

  

Epicopeiidae and Drepanidae have recently been included in two large molecular studies of ditrysian phylogeny. In neither study do these two families ever group together. Regier, Zwick et al. (2009) found Epicopeiidae to group invariably with or within Geometroidea, although bootstrap support for Geometroidea including Epicopeiidae was weak (<50%). The most strongly supported specific position for Epicopeiidae was as sister group to Sematuridae (67% bootstrap). The Approximately Unbiased test of Shimodaira showed near-significant rejection (P=0.08) of monophyly for Drepanoidea as currently defined. Mutanen et al. (2010)found Epicopeiidae to group with Lasiocampidae, but with bootstrap <50%. While additional evidence will be needed to determine the exact phylogenetic position of Epicopeiidae, it now seems very unlikely that they are the sister group to Drepanidae.

  Minet (2002) provided an estimate of phylogenetic relationships among the genera of Epicopeiidae based on an analysis of morphological characters, as shown in this figure.

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Systematic and taxonomic history

Systematic and taxonomic history:

Account taken from Minet 2002: For most of its history, this family was restricted to the genus Epicopeia. Several authors had however noted similarities of this genus to others. In a series of papers beginning in 1983, Minet formally moved additional genera into Epicopeidae which had previous been placed in Epiplemidae, now regarded as a subfamily of Uraniidae (Geometroidea).  
 
 Minet (1991) regarded Epicopeiidae as sister group to the Drepanidae (see also Minet and Scoble 1998, Minet 2002), although Scoble (1982) had allied them tentatively with Uraniidae. Minet (2002) proposed the following synapomorphies for Drepanoidea comprising these two families: (a)imaginal abdomen base with complete or nearly complete tergosternal sclerites;  (b) imaginal abdomen base with neotergite distinctly fused with marginotergites; (c) larval mandible with a large, flat, lateral area, which is delimited ventrally by a well defined carina;  (d) most larval abdominal segments with at least one secondary seta cephalad of L3; (e) pupa never exhibiting conspicuous portion of the fore femur.  Several of these apomorphies were regarded as tentative. 
 Epicopeiidae and Drepanidae have recently been included in two large molecular studies of ditrysian phylogeny. In neither study do these two families ever group together. Regier, Zwick et al. (2009) found Epicopeiidae to group invariably with or within Geometroidea, although bootstrap support for Geometroidea including Epicopeiidae was weak (<50%). The most strongly supported specific position for Epicopeiidae was as sister group to Sematuridae (67% bootstrap). The Approximately Unbiased test of Shimodaira showed near-significant rejection (P=0.08) of monophyly for Drepanoidea as currently defined. While additional evidence will be needed to determine the exact phylogenetic position of Epicopeiidae, it now seems very unlikely that they are the sister group to Drepanidae.

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Fossil Record

Fossil record:

No fossils are known

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Molecular Biology and Genetics

Molecular Biology

Statistics of barcoding coverage

Barcode of Life Data Systems (BOLD) Stats
                                        
Specimen Records:17Public Records:6
Specimens with Sequences:17Public Species:2
Specimens with Barcodes:16Public BINs:2
Species:4         
Species With Barcodes:4         
          
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Barcode data

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Locations of barcode samples

Collection Sites: world map showing specimen collection locations for Epicopeiidae

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