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Reassessment of the enigmatic Lepidopteran family Lypusidae (Lepidoptera: Tineoidea; Gelechioidea)

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  • University of Helsinki, Helsinki, Finland

Abstract and Figures

The genus LypusaZeller, 1852 has been assigned to Tineoidea (Lepidoptera) with varying family positions. The systematic affinities of this genus, currently placed in its own family Lypusidae, were studied using extensive data derived from larval, pupal and adult morphology and certain behavioural traits. In total, 193 characters were considered. On the basis of the results of a parsimony analysis, Lypusa is transferred to the superfamily Gelechioidea, in a monophyletic, exclusively Palaearctic assemblage with the genera Amphisbatis and Pseudatemelia. Several phylogenetically relevant characters support this position, including similarly constructed larval cases, densely porose larval head, and a modification of the pupal abdominal segment 8. The composition of the family Amphisbatidae is discussed and it is proposed that it be delimited as comprising solely the Palaearctic genera Pseudatemelia, Amphisbatis and Lypusa. A detailed diagnosis of the genus Lypusa is given.
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Systematic Entomology (2010), 35, 71 –89 DOI: 10.1111/j.1365-3113.2009.00483.x
Reassessment of the enigmatic Lepidopteran family
Lypusidae (Lepidoptera: Tineoidea; Gelechioidea)
MARIA HEIKKIL ¨
AandLAURI KAILA
Finnish Museum of Natural History, University of Helsinki, Finland
Abstract. The genus Lypusa Zeller, 1852 has been assigned to Tineoidea (Lepi-
doptera) with varying family positions. The systematic affinities of this genus, currently
placed in its own family Lypusidae, were studied using extensive data derived from
larval, pupal and adult morphology and certain behavioural traits. In total, 193 char-
acters were considered. On the basis of the results of a parsimony analysis, Lypusa is
transferred to the superfamily Gelechioidea, in a monophyletic, exclusively Palaearctic
assemblage with the genera Amphisbatis and Pseudatemelia. Several phylogenetically
relevant characters support this position, including similarly constructed larval cases,
densely porose larval head, and a modification of the pupal abdominal segment 8. The
composition of the family Amphisbatidae is discussed and it is proposed that it be
delimited as comprising solely the Palaearctic genera Pseudatemelia, Amphisbatis and
Lypusa. A detailed diagnosis of the genus Lypusa is given.
Introduction
Lypusa (Zeller, 1852) is a Palaearctic moth genus, which, until
recently, was supposed to comprise a single widespread species
(Fig. 1), L. maurella (Denis & Schifferm¨
uller, 1775), found
from Western Europe (Fibiger & Svendsen, 1981; Zagulyaev,
1989; Davis & Robinson, 1998) to the Urals, Siberia and north-
eastern China (unpublished data in the Finnish Museum of
Natural History). However, as has been found with many other
species thought to be widespread, it has become evident that it
actually comprises a species group (Elsner et al., 2008; Kaila
& Heikkil¨
a, personal communication), but in the current article
we refrain from elaborating further on the species taxonomy
of the genus as the number of species in this group remains to
be investigated.
Currently, Lypusa is placed in its own family, Lypusidae,
within the superfamily Tineoidea. Because of its peculiar adult
morphology, combined with poorly known immature stages, its
systematic position has been unstable and remains unresolved
(e.g. Davis & Robinson, 1998). Lypusa has usually been
assigned either to Tineidae, Psychidae or in its own family
Lypusidae within Tineoidea.
Lypusa maurella was described first as Tinea maurella
by Denis & Schifferm¨
uller (1775). Not satisfied with that
Correspondence: Maria Heikkil¨
a, Finnish Museum of Natural
History, P. O. Box 17, FI-00014 University of Helsinki, Finland.
E-mail: maria.heikkila@helsinki.fi
generic placement, subsequent authors included maurella in
various diverse genera, such as Adela (Eversmann, 1844) and
Oecophora (Zeller, 1839; Duponchel, 1844). At that time, these
genera were understood to belong to the large assemblage
Tineina. In 1852, although agreeing that the species shared
a likeness with Oecophora flavifrontella (=Pseudatemelia
flavifrontella), Zeller established a new genus, Lypusa, with
maurella as its only included species. The family status of L.
maurella continued to be disputed, and in 1870 Heinemann
placed the species in its own family, Lypusidae. Subsequently,
several other species were temporarily placed in Lypusidae or
Lypusinae (Staudinger & Wocke, 1871; Mees & Spuler, 1910),
but these were later removed by other authors. Differences in
opinion did not cease, however, and Eyer (1924), for example,
retained L. maurella in Tineidae.
The case-bearing habit of the larva (Eversmann, 1844;
Heinemann, 1870–1877; Frey, 1880; Sorhagen, 1886) prob-
ably influenced the decision of some later authors to place
Lypusa within Psychidae (e.g. Tutt, 1900; Dalla Torre & von
Strand, 1929; Petersen, 1964; Fibiger & Svendsen, 1981; Zag-
ulyaev, 1989). Sauter (1982), however, stated that Lypusiinae
[sic] did not fit within Psychidae and moved them to a place
in the group of lichen-feeding tineids. Sauter & H¨
attenschwiler
(1991) likewise argued that Lypusa was not a psychid on the
basis of the structure of the genitalia and the pupa, and retained
it with Tineidae.
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society 71
72 M. Heikkil¨a and L. Kaila
Fig. 1. Habitus of Lypusa maurella Germany, Potsdam, Museum
f¨
ur Naturkunde der Humboldt-Universit¨
at, Berlin.
In reviewing the literature on L. maurella, Dierl (1996) stud-
ied 21 specimens of the species but could not resolve the
question as to which family Lypusa belonged, and so provi-
sionally retained it in Lypusidae. The most recent comment
on the systematic position of L. maurella is that of Davis &
Robinson (1998), who stated that L. maurella is ‘not a psy-
chid as evidenced by the free arms of the adult metafurca’ and
not a tineid due to the ‘unusual larval case construction’. They
placed the family Lypusidae within Tineoidea, but emphasized
that its affinities were unknown and that study of the immature
stages was needed.
The aim of the current study was to investigate the sys-
tematic position of Lypusa using the extensive morphologi-
cal dataset developed by Kaila (2004) for the study of the
interrelationships within Gelechioidea. We deemed this appro-
priate after noting that although reduced, the base of the
haustellum of Lypusa is scaled. Likewise, the pupal anten-
nae of Lypusa meet each other mesially. These are rare
traits among Lepidoptera, but one of the few universal or
nearly universal synapomorphies of Gelechioidea. As the
data matrix of Kaila (2004) also includes representatives of
11 families placed in nine superfamilies, in addition to the
densely sampled Gelechioidea, we considered this matrix a
suitable background for the study of the systematic position of
Lypusa.
Material and methods
Study material
The morphological analysis and coding of character states
in our study were based on specimens of L. tokari, described
and named by Elsner et al. (2008) (Fig. 2), which were
collected on Mount Olympus in Greece (coll. J. Kaitila and
Finnish Museum of Natural History). For the determination
of larval and pupal characters, only the larval and pupal
exuviae and a larval case from Tyrol in Austria were available
(Tiroler Landesmuseum Ferdinandeum, Innsbruck, Austria).
These were in fairly good condition and most character
states could be coded from them. However, a few ambiguous
characters of the pupal exuvia (characters 140– 142) could
Fig. 2. Habitus of Lypusa tokari Greece, Olympos, Finnish
Museum of Natural History.
not be coded because this region had been lost at adult
eclosion.
Morphological methods
In total, 188 morphological characters and five charac-
ters from larval behaviour were coded. The list of characters
(Appendix 1) and terminology are adapted from those pub-
lished by Kaila (2004), where explanations and illustrations of
several of the characters can be found. The data matrix of that
study was left unchanged (i.e. the other included taxa were not
re-examined or recoded). The characters in question encom-
passed 241 informative character states. Ambiguous, inappli-
cable or missing data are denoted in the matrix as ‘?’.
External characters of the adults and pupa were examined
with a Leica MZ 75 stereomicroscope, magnification up to
400×. For examination of structural characters, adults were
macerated in KOH solution, denuded of scales and hairs, kept
in glycerol or ethanol and examined with a Wild M10 stere-
omicroscope, magnification up to 512×. The larval skin was
also preserved in glycerol and studied with this microscope.
Abdominal skins, genitalia and wings were mounted on per-
manent slides using standard procedures (Robinson, 1976) and
examined with the Leica MZ 75 stereomicroscope and a Leitz
Diaplan phase contrast microscope (maximum magnification
1560×).
The results of the character coding of L.tokari are shown
in Appendix 2, which also includes data on four taxa studied
by Kaila (2004): Scardia boletella (Tineidae), which was
the outgroup taxon used to root the trees, and the three
Amphisbatidae species with which L.tokari groups in the
present analysis (see below). The full matrix for the other taxa
was published by Kaila (2004).
Phylogenetic methods
Phylogenetic analysis was undertaken first using TNT version
1.0 (Goloboff et al., 2000), and complemented with NONA
version 2.0 (Goloboff, 1993) and WINCLADA version 1.00.08
(Nixon, 2002). In TNT, the memory was set to 100 000 trees
and general RAM to 1000 MB. Characters were not weighted
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
Reassessment of Lypusidae 73
Fig. 3. Lypusa maurella. Base of female antenna. Arrow points to
the enlargement due to denser scale coverage.
differentially a priori and multistate characters were treated
as non-additive. From the New Technology Search menu, all
techniques were selected (sectorial search, ratchet, drift and
tree fuse). The program was set to find the minimum-length
tree 20 times. The most-parsimonious trees found by TNT were
then submitted to WINCLADA and NONA, in which the multistate
characters were treated also as non-additive. NONA was set to
store 100 000 trees in memory (hold 100 000) and the shortest
trees searched for with the command max*. The resulting trees
were saved (sv*) and the analysis was continued in WINCLADA.
In WINCLADA, the following commands were applied to all
trees: hard collapse all unsupported nodes in all trees; keep best
trees only. The consensus tree was calculated by the application
of the Nelsen consensus (collapse +consensus). The Bremer
support values (the number of extra steps required before a
clade is lost from the strict consensus tree of near-minimum
length cladograms) for each node were obtained in NONA by
use of the following commands: hold 2000; sub 1; find*; hold
4000; sub 3; find*; hold 8000; sub 5; find*; hold 16 000; sub
7; find*; hold 32 000; sub 10; find*; bsupport;.
Results
TNT retained 40 trees of length 1512 steps. Subsequent anal-
ysis by NONA yielded 8448 equally most-parsimonious trees
of 1512 steps (consistency index =0.16; retention index =
0.68). Of these, 768 suboptimal trees were removed as non-
parsimonious following hard collapsing, and 7680 trees were
kept. The strict consensus cladogram of these trees (18 col-
lapsed nodes) (Fig. 5) is nearly identical to that of Kaila (2004),
with the added taxon, L. tokari, positioned in the ‘oecophorid
lineage’. Lypusa tokari is grouped with Amphisbatis incon-
gruella, Pseudatemelia josephinae and Pseudatemelia flav-
ifrontella, supported by six homoplastic characters. With the
addition of L. tokari, this assemblage loses the resolution it
had in Kaila’s study and becomes an unresolved polytomy.
In Kaila’s study, Amphisbatis incongruella and Pseudatemelia
flavifrontella together formed the sister clade of Pseudatemelia
josephinae, supported by synapomorphies in genital structure
(90: 0 and 91: 4).
The main result is that Lypusa does not group with
Tineoidea, where it has been placed traditionally. The mor-
phology of Lypusa shows several character states that argue
against a position in that superfamily. Four unique synapo-
morphies (Fig. 6: node 1) separate all the other taxa, including
Fig. 4. Lypusa maurella. Lamellate scale vestiture at haustellum base
(character 4: 1).
Lypusa, from Tineidae. The maxillary palpi of Tineidae have
five segments and those of all the others are shorter (6: 1) (in
one species this is reversed, Oegoconia deauratella, which also
has five). Lypusa and all other moths outside Tineoidea lack
lateral bristles on the labial palpi (8: 0) and have no sternal rod
on the ovipositor (135: 0). In addition, the transverse rows of
spines at the posterior margin of at least some segments of the
pupal abdomen, present in Tineidae, are absent from most other
moths, including Lypusa (152: 0) (this character is reversed in
Synanthedon scoliaeformis, Cossus cossus, Aphelia viburnana,
Orthotaenia undulana, Deoclona yuccasella and some Xylo-
ryctinae). Node 1 is supported by a further six homoplastic
characters. The unambiguously optimized homoplastic charac-
ters on nodes 1– 12 are commented on in the discussion section
below.
Lypusa associates neither with Tineoidea nor Roeslerstam-
miidae as the pupae of these have transverse rows of spines
at the anterior margin of abdominal segments. As a unique
synapomorphy, the absence of at least some of these rows of
spines (151: 0) unites all the other taxa (Fig. 6: node 2). This
character is reversed in this analysis in eight species: Choreutis
pariana, Synanthedon scoliaeformis, Cossus cossus,thetwo
Tortricidae exemplars, Deoclona yuccasella, Oegoconia deau-
ratella and Xylorycta orectis (note, however, that the spine
rows in at least some gelechioids seem to be different struc-
turally from those of other superfamilies, and coding them as
the same character state may be erroneous). Lypusa is also
excluded from the superfamily Gracillarioidea (node 3) as a
whole as its metafurca has a joint as a pair of lobes between
the stem and the secondary arms (32: 0). In Tineoidea and
Gracillarioidea, these lobes are absent. Another trait excludes
Lypusa from Tineoidea and Gracillarioidea. The pupae in these
superfamilies move out of the cocoon, but pupae of most of
the other taxa (including Lypusa) do not (165: 1) (reversed
in Choreutis pariana, Synanthedon scoliaeformis and Cossus
cossus).
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
74 M. Heikkil¨a and L. Kaila
Gelechioidea
Scardia boletella (Tineoidea: Tineidae)
Nemapogon granellus (Tineoidea: Tineidae)
Callisto denticulella (Gracillarioidea: Gracillariidae)
Roeslerstammia erxlebella (Gracillarioidea: Roeslerstammiidae)
Yponomeuta evonymellus (Yponomeutoidea: Yponomeutidae)
Ypsolopha dentella (Yponomeutoidea: Ypsolophidae)
Choreutis pariana (Choreutoidea: Choreutidae)
Aphelia viburnana (Tortricoidea: Tortricidae)
Orthotaenia undulana (Tortricoidea: Tortricidae)
Synanthedon scoliaeformis (Sesioidea: Sesiidae)
Cossus cossus (Cossoidea: Cossidae)
Udea lutealis (Pyraloidea: Pyralidae)
Epermenia illigerella (Epermenioidea: Epermeniidae)
Perittia obscurepunctella
Stephensia abbreviatella
Elachista adscitella
Elachista gerasmia
Urodeta hibernella
Agriophara axesta
Agriophara platyscia
Agriophara cinerosa
Antaeotricha schlaegeri
Scieropepla typhicola
Uzucha humeralis
Tymbophora peltastis
Lichenaula sp.
Xylorycta orectis
Glyphidocera lithodoxa+juniperella
Scythris empetrella
Scythris limbella
Scythris inspersella
Prionocris sp.
Oecophora bractella
Philobota glaucoptera
Phryganeutis cinerea
Palimmeces sp.
Pleurota bicostella
Hofmannophila pseudospretella
Harpella forficella
Denisia similella
Borkhausenia fuscescens
Bisigna procerella
Promalactis venustella
Polix coloradella
Phaeosaces coarctatella
Izatha peroneanella
Tingena hemimochla
Tingena armigerella
Hierodoris atychioides+callispora
Coelopoeta glutinosi
Stathmopoda pedella
Stathmopoda melanochra
Stathmopoda aenea
Diurnea fagella
Dasystoma salicella
Agonoxena phoenicia
Blastodacna atra
Heinemannia laspeyrella
Trachystola macrostola
Microcolona sollennis
Spuleria flavicaput
Agonopterix heracliana
Agonopterix arenella
Exaeretia ciniflonella
Depressaria badiella
Depressaria pastinacella
Depressaria sordidatella
Levipalpus hepatariellus
Semioscopis steinkellneriana
Barantola panarista
Proteodes carnifex
Donacostola notabilis
Enteremna pallida
Pedois lewinella
Hypercallia citrinalis
Anchinia daphnella
Telechrysis tripuncta
Orophia ferrugella
Aeolanthes siphonias
Carcina quercana
Pseudatemelia josephinae
Machimia tentoriferella
Psilocorsis reflexella
Hypertropha tortriciformis
Eupselia carpocapsella
Eupselia satrapella
Thudaca mimodora
Ethmia clytodoxa
Ethmia pusiella
Ethmia heliomela
Deuterogonia pudorina
Syringopais temperatella
Deoclona yuccasella
Pterolonche inspersa
Coleophora serratella
Coleophora expressella
Coleophora brevipalpella
Coleophora caelebipennella
Goniodoma limoniella
Mompha idaei
Mompha conturbatella
Mompha sturnipennella
Mompha epilobiella
Batrachedra praeangusta
Batrachedra pinicolella
Batrachedra eustola
Idioglossa polliacola
Homaledra sabalella
Holcocera gigantella
Hypatopa binotella
Blastobasis phycidella
Blastobasis decorella
Blastobasis yuccaecolella
Pigritia laticapitella
Lecithocera nigrana
Crocanthes prasinopis
Cophomantella lychnocentra
Odites kollarella
Odites leucostola
Odites xenophaea
Autosticha modicella
Cosmopterix lienigiella+scribaiella
Echinoscelis pandani
Anatrachyntis rileyi
Limnaecia phragmitella
Limnaecia cirrhozona
Limnaecia scoliosema
Vulcaniella rosmarinella
Macrobathra leucopeda
Macrobathra chrysotoxa
Macrobathra alternatella
Paratheta calyptra
Anoncia piperata
Pancalia schwarzella
Pancalia nodosella
Sorhagenia janiszewskae
Walshia miscecolorella
Anatiplora basiphaea
Nemotyla oribates
Oegoconia deauratella
Holcopogon bubulcellus
Symmoca signatella
Deroxena venosulella
Neofriseria peliella
Syncopacma sangiella
Anacampsis populella
Helcystogramma rufescens
Dichomeris juniperella
Pexicopia malvella
Hypatima rhomboidella
Filatima incomptella
Elachistidae
Chimabachidae
Stenomati[n/d]ae
Amphisbatidae s. str.
Carcinidae
Oecophoridae s. str.
Elachistinae
Agonoxeninae
Parametriotinae
Aeolanthinae
Ethmiinae
Depressariinae s. str.
Unplaced Elachistidae
Hypertrophinae
Xyloryctid assemblage
Blastobasinae
Deuterogoniinae
Autostichid assemblage
Cosmopterigidae
Cosmopteriginae
Antequerinae
Chrysopeleiinae
Gelechiidae
Scythrididae
Pterolonchidae
Momphidae
Coleophoridae
Syringopai[n/d]ae
Deoclonidae
Holcopogoninae
Symmocinae
Glyphidocerinae
Oditinae
Autostichinae
Lecithocerinae
Oecophorid
lineage
Xyloryctinae. s. str
Gelechiid
lineage
Apoditrysia
9
>10
1
3
2
3
21
58
2
3
2
2
3
72
2
2
9
2
1
2
>10
1
3
8
31
3
7
1
1
2
10
31
1
2
4
3
4
1
2
2
3
2
3
2
1
2
2
1
2
2
2
1
6
2
4
21
1
1
2
2
1
6
7
11
10
3
3
1
1
1
1
1
1
1
1
13
1
4
1
3
1
1
3
5
>10
1
3
1
1
1
1
1
1
7
72
1
1
1
1
1
3
2
1
3
1
3
6
1
5
3
1
3
3
2
3
1
3
2
Pseudatemelia flavifrontella
Amphisbatis incongruella
Lypusa tokari
Fig. 5. Strict consensus tree of 7680 equally most-parsimonious trees based on 189 morphological and four ecological characters. Bremer support
values are shown above the branches. The black arrow indicates placement of the added taxon, Lypusa tokari .
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
Reassessment of Lypusidae 75
Scardia boletella
Nemapogon granellus
Roeslerstammiidae
Gracillariidae
Yponomeutoidea
Epermenioidea
Choreutoidea-Sesioidea-Cossoidea
Pyraloidea-Tortricoidea
Gelechiid lineage
Autostichi dassemblage
Xyloryctid assemblage
Oecophoridae
s.s.
Lypusa
Pseudatemelia flavifrontella
Pseudatemelia josephinae
Amphisbatis incongruella
Carcina quercana
(continues)
0 6 8 17 30 42119135152163
0101010000
291151
030
12732 165
1101
33 7176 85 181
10110
15 91
0
17 46114
124
01
66 71119
145
121
2023 25 105121
11011
171
175186
111
9699141
184
11
2
1
1866 69159
10 1
71
1011
11 0 1
52133169175 191
49 99107163184
002 1
107
161
21
113118185
190
111
71636 45 49 54 55 61 83 95 96 102
111
132
10103222
0111
1
2
Gelechioidea
3
4
5
6
7
8
9
10
11
12
0
186
119143144
0
1
1
0
00
1
10
1
01
Fig. 6. Simplified tree showing apomorphies on nodes as far as placement of Lypusa tokari within Gelechioidea. The numbered arrows indicate
nodes referred to in the text.
Lypusa is excluded also from Yponomeutoidea (Fig. 6:
node 4) because the stem and secondary arm of its meta-
furca have a joint between them (33: 1, unique character
defining superfamilies that are more advanced than Gracillario-
idea and Yponomeutoidea). In Tineoidea, Gracillarioidea and
Yponomeutoidea, the stem and secondary arm are fused to each
other without a joint (33: 0). A second unique synapomorphy
excluding Lypusa from Yponomeutoidea is the socius reduced
to a small group of setae at the base of the uncus (85: 1).
Yponomeutoidea and Roeslerstammia erxlebella have a socius
developed as a free setose lobe (reversals; Tortricidae, at least
two times in Scythrididae, Pexicopia malvella, Hierodoris,sev-
eral times in Elachistidae).
Two homoplastic characters support node 5 and separate
Epermenia illigerella (Epermenioidea) from the superfamilies
further along the tree. These homoplastic characters are hair-
like scales on lateroposterior area of metascutum (15: 0) (also
the state present in Lypusa) and absence of scobination of
mesal part of gnathos (91: 0) (Lypusa coded with a ? as the
mesal part of gnathos is rudimentary).
Two synapomorphies are shared by most taxa in Pyraloidea,
Tortricoidea and Gelechioidea, including Lypusa (Fig. 6: node
6). The first is the foldable membrane of fultura inferior,
which in these superfamilies is medially overlaid to the phallus
(114: 0). In Tineoidea, Gracillarioidea, Yponomeutoidea, Eper-
menioidea, Choreutoidea, Sesioidea and Cossoidea, the mem-
brane is present posteriorly. The posterior position is also found
in several other taxa, where it is interpreted as a reversal: Deo-
clona yuccasella, several times in Coleophoridae, Pterolonche
inspersa, and several times in the autostichid and xyloryctid
assemblages. The second concerns the structure of the signum.
In many taxa in Pyraloidea, Tortricoidea, Gelechioidea and
Lypusa the teeth of the signum are directed towards the margin
(124: 1), but this is never the case in the more basally originat-
ing superfamilies of the taxon sampling. Again, however, there
are several reversals in the gelechiid lineage and the xyloryctid
assemblage.
The adult character placing Lypusa among Gelechioidea
is the scaled base of its haustellum (4: 1) (Fig. 4). This
is a character state that all Gelechioidea possess, but it
can also be observed in Tischerioidea, Choreutoidea and
Pyraloidea (Kaila, 2004). The second important character
present in Lypusa supporting its position in Gelechioidea
is a character derived from pupal morphology: antennae
touching each other mesially where they lie parallel (145: 1)
(Fig. 7), although it is reversed in some instances (see e.g.
Common, 1990; Kaila, 2004). In the present taxon sampling,
this character state is a unique apomorphy of Gelechioidea.
Outside of this superfamily, antennae touching each other are
also found in some hieroxestine Tineidae (Davis, 1978), some
Roeslerstammiidae (Common, 1990) and Lycaenidae (e.g.
Patoˇ
cka & Turˇ
c´
ani, 2005) that are not included in the present
study. In addition to mesially touching antennae, another three
homoplastic characters support the monophyly of Gelechioidea
(Fig. 6: node 7). These are further discussed below.
Lypusa is placed within the ‘oecophorid lineage’, the mono-
phyly of which (Fig. 6: node 8) is supported by three homo-
plastic characters: ecdysial line of larval adfrontal sclerites does
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
76 M. Heikkil¨a and L. Kaila
Fig. 7. Lypusa tokari. Pupal antennae on exuvia mesially touch each
other (character 145: 1).
not reach the cranial incision (171: 1) (this condition is also
present in Lypusa, but is reversed in several other instances);
horizontal arrangement of the two anterior L group setae on
the larval prothorax (175: 1) (in Lypusa, however, they are
oblique); and hair-like SD1 on larval A9 (186: 1) (in Lypusa
it is as thick as the other setae).
Node 9 is supported by four homoplastic characters: pres-
ence of a pair of sclerotized lobes on the lateral or distolateral
margin of the ventral plate of the juxta (96: 1) (absent in
Lypusa); presence of a sclerotized lobe in the distal area of
the sacculus (99: 1); pupal labial palpus concealed (141: 1)
(Lypusa was coded as ? because the pupal skin available was
broken in this region); and SV group of setae on the prolegs
of A3– A6 on one pinaculum (184: 0) (arranged this way in
Lypusa also).
Node 10 is supported by one homoplastic character: pecten
on the scape of the antenna comprising a row of narrow stiff
scales (2: 1) (this character state is also observed in Lypusa).
Six homoplastic characters from the female and larval mor-
phology support the grouping of Lypusa with Amphisbatidae
(node 12), the first four of which are also found in all three
amphisbatid species, namely: length of S8 on female pro-
longed as an extensible telescope (133: 1); portable sack on
larva present (169: 1); oblique arrangement of the two ante-
rior setae in the L group of the prothorax, with the anterior
seta dorsad of the posterior seta (175: 0); anal comb present
(191: 1). Character state 186: 0 (shape of seta SD1 at A9 does
not differ from other setae) is a character that Lypusa shares
with at least Pseudatemelia flavifrontella. However, for the two
other amphisbatids the state was coded as (?) (missing data).
The sixth synapomorphy (52) concerns the basal forking of
forewing 1A +2A, the costal branch of which is absent, or
present as at most a vestigial fold, in the three Amphisbati-
dae. That of Lypusa, however, is ambiguous and coded as ‘?’
because there were differences in wing venation between the
sexes and among different species of Lypusa.
Discussion
Superfamily placement of Lypusa
The present results support a position of the genus Lypusa
in the superfamily Gelechioidea instead of Tineoidea. Several
unique synapomorphies place Lypusa within the ingroup taxa
and exclude it from Tineoidea (Fig. 6: nodes 1 6), whereas
the mesially touching pupal antennae (node 7, character 145)
is strong evidence for Lypusa being a gelechioid.
In addition to the apomorphies on nodes 1– 4 and 6 7 noted
above, the position of Lypusa in Gelechioidea is supported
by a number of unambiguously optimized, but homoplastic,
characters. The strength of the support afforded by these
characters is discussed in this section. Several of the characters
on nodes 1–12 are known to be widely homoplastic within
Lepidoptera, and in the light of the present knowledge, the
phylogenetic information they offer is highly ambiguous at
best. Thus, the following characters are not discussed further
here: scale vestiture on frons (0); shape of metascutellum
(17); intercoxal lamella (30); vestiture behind antenna (1);
obliquely directed suture directed to median incision of meron
(27); vestiture on lateroposterior area of metascutum (15);
pterostigma (45) [see Kaila (2004) for a critical discussion];
ecdysial line of adfrontal sclerites (171); position of L3
on prothorax (176); pair of sclerotized lobes in lateral or
distolateral margin of ventral plate of juxta (96); sclerotized
lobe in distal area of sacculus (99); pupal labial palpus (141);
setae of SV group (184); and pupal abdominal segment 8 (159).
The other homoplastic characters unambiguously opti-
mized on to nodes 1–12 are here commented on in greater
detail, beginning with those supporting the division between
Tineoidea and the other taxa (node 1). The occurrence of the
dorsoposterior process of the metafurcal apophysis (42: 1) is
poorly known among Lepidoptera, but appears generally rare.
Node 1 is also supported by a character state of female geni-
talia, the membrane between the papillae anales and segment 8
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
Reassessment of Lypusidae 77
being only minimally extensile or not at all telescopic (119: 0).
Telescoping of the female postabdomen seems to be unusual
in Ditrysia. In Gelechioidea (node 7), however, both of these
characters are frequent, including Lypusa and the Amphisbati-
dae. The absence of the lateral spine at the pupal segment
10 (163: 0) unites most of the more ‘advanced’ taxa in the
study. The presence of this lateral spine is common in ‘primi-
tive’ Lepidoptera, rare in ‘advanced’. In our study, the lateral
spine on segment 10 was found to be present on tineid pupae,
Ypsolopha dentella, Choreutis pariana and Cossus cossus,in
the Coleophoridae and three times elsewhere within the Gele-
chioidea.
Nodes 2 and 10 are supported by different states of the
homoplastic character 2, which codes for the presence or
absence of the pecten on the scape of the antenna. We found the
pecten to be absent from Gracillarioidea, Yponomeutoidea and
Epermenioidea. However, outside the present taxon sampling
a pecten or tuft of scales on the scape has been recorded
elsewhere in these superfamilies (Davis & Robinson, 1998;
Dugdale et al., 1998a). Our finding of the presence of a pecten
(2: 1 and 2: 2) supports several monophyletic groupings (in
the half of the gelechiid lineage from Mompha onwards,
Blastobasinae, Oecophoridae s.str. +Amphisbatidae s.str. +
Carcinidae and in Elachistidae). Otherwise, the pecten seems
to be relatively infrequent in Lepidoptera. In addition to the
superfamilies mentioned above it has been observed in some
Tineoidea, Zygaenoidea, Galacticoidea and Noctuoidea (Davis
& Robinson, 1998; Dugdale et al., 1998b; Epstein et al., 1998;
Kitching & Rawlins, 1998).
Nodes 2 and 5 are supported by different states of the
homoplastic character 91, scobination of the mesial part of
the gnathos. The type of gnathos scobination, or its absence,
is variable across the included taxa.
The homoplastic characters supporting node 4 have been
proposed as supporting the monophyly of Apoditrysia (Kyrki,
1983; Minet, 1991), but in the study by Kaila (2004) the char-
acter complex of sternum 2 proved to be quite homoplastic.
The sternal rod in sternum 2 (71) is absent from the included
taxa of Epermenioidea, Choreutoidea, Sesioidea, Cossoidea,
Pyraloidea and Tortricoidea (node 4). In Gelechioidea, both
‘tortricid-type’ broad indistinctly delimited ridges and ‘tineid-
type’ sharply delimited narrow long ridges (71: 2) are present.
Many Gelechioidea have a mixture of tineid-type sternal rods
and tortricid-type lateral corners (see Kaila, 2004). Otherwise,
in Lepidoptera narrow sternal rods are exceptional. The charac-
ter was ambiguous in Lypusa because of its sexual dimorphism,
females having sternal rods whereas some males have indis-
tinctly delimited ridges, others none. The same discussion is
largely applicable to character state 76: 1.
Node 4 is also supported by character state 181: 0, the close
proximity of setae L1 and L2 on larval segments A1– A8.
Minet (1991) proposed this character as a putative apomorphy
for Apoditrysia, with their wide spacing in many advanced
Apoditrysia due to multiple character reversals. Examination
of larvae of groups assigned traditionally to the Apoditrysia
indeed shows that both approximated and widely spaced setae
are found in Apoditrysia. The close approximation of the setae
is, however, the most commonly encountered character state in
Gelechioidea. Lypusa larva has closely approximated setae L1
andL2onsegmentsA1A8.
Of the characters supporting the monophyly of Gelechioidea
(node 7), the extensible membrane between the papillae anales
(119) was discussed above. Node 7 is supported by two
other homoplasies. Scales transformed into setae on the terga
(66: 1) is a character typical of only Yponomeutoidea and
Gelechioidea, but the reversed condition is also common
(node 11). Amphisbatidae and Lypusa all lack these setae,
and they are also lacking from their sister clade (except for
Chimabachidae).
Systematic position of Lypusa within Gelechioidea
Discussion of nodes 8– 10 has been elaborated by Kaila
(2004) and is not repeated here.
Node 11 is supported by, among others, the homoplastic
presence of apodemes on male sternum 2 (69). The presence
of apodemes is variable in Gelechioidea and the kinds of
apodeme thought of as uniting Apoditrysia were also observed
in some Gelechioidea. Sexual dimorphism in the presence
of apodemes was found frequently. Apodemes may serve as
supporting structures for the heavier female abdomen (Kaila,
2004). There is sexual dimorphism in Lypusa in the presence
of these apodemes as they are absent in males and present
in females. The oecophorid lineage (node 8) is supported
by the shape of the larval seta SD1 on A9. The hair-like
condition of this seta (186: 1) is typical, although variable, in
Gelechioidea. Some ypsolophids (Yponomeutoidea), noctuids
and pyralids also exhibit a similar hair-like seta. Node 11 is
also supported by character state 18: 1, the mesially interrupted
internal folding of the metascutellum. It is not found outside
Gelechioidea, but within it is quite variable. However, this
character has not yet been well studied throughout Lepidoptera.
Five of the six characters placing Lypusa in a mono-
phyletic group (Fig. 6: node 12) with Pseudatemelia josephi-
nae, Pseudatemelia flavifrontella and Amphisbatis incongruella
are derived from the female, larval and pupal morphology.
The portable sack of the larva (Figs 8, 9) and the anal comb
(Fig. 10) are encountered in only a few species outside this
grouping. The portable sack (169: 1, two steps) is found only
on Goniodoma limoniella and Coleophora, but there are quite
a number of question marks due to unavailable material. The
anal comb (191: 1, three steps) otherwise is present only in
Fig. 8.Lateral view of larval case of Lypusa tokari. Austria, Tiroler
Landesmuseum Ferdinandeum, Innsbruck.
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
78 M. Heikkil¨a and L. Kaila
Fig. 9. Distal opening of larval case of Lypusa tokari. Austria, Tiroler
Landesmuseum Ferdinandeum, Innsbruck.
Fig. 10. Lypusa tokari. Larval anal comb (character 191: 1).
the larvae of the tortricids, Aphelia viburnana and Orthotae-
nia undulana, and the Gelechiidae grouping except Pexicopia
malvella (the presence or absence of the anal comb in Hypa-
tima rhomboidella is unknown). An anal comb is also present
in Eudarcia (Tineidae) larvae (D. R. Davis, personal commu-
nication).
The oblique arrangement of two anterior setae in the L
group of the larval prothorax, with the anterior seta dorsad
of posterior seta (175: 0, steps 15), and seta SD1 on A9
of the same thickness as the other setae (186: 0) are the
most commonly encountered states in the outgroup taxa, the
Fig. 11. Lypusa tokari. Lateral view of distal end of pupal abdomen
with small group of setae (character 164: 0). The arrow points to the
transverse fold and protuberances on A8, a possible synapomorphy of
Amphisbatidae s.str.
‘gelechiid lineage’, and are found in the oecophorid lineage on
several occasions in quite a scattered manner, suggesting that
both have undergone multiple reversals.
In the Lypusa –Amphisbatidae group, sternum 8 of the
female abdomen is prolonged as an extensible telescope
(133: 1). Outside of this grouping, this character state is
present in only seven species included in the study, belong-
ing to many different lineages, pointing to extensive homo-
plasy. These species are Syringopais temperatella, Sorhagenia
janiszewskae, Holcopogon bubulcellus, Deroxena venosulella,
Hierodoris atychioides +callispora, Philobota glaucoptera
and Polix coloradella.
The homoplastic characters uniting Lypusa with the amphis-
batids are complemented by one significant larval character and
one pupal character, not included in the character coding of the
present study, but under study by the same research team (L.
Kaila, personal communication). The larvae of all four species
have ‘porous’ head capsules, that is, the head is densely cov-
ered by distinctive pits. A pitted head of this form is otherwise
rare in Lepidoptera, only to be found in Xyloryctidae (Gele-
chioidea), Thyrididae and some Papilionoidea. The pupae of
all Amphisbatidae s.str. species studied have a transverse fold-
ing on the eighth abdominal segment, usually with a pair of
spines or protuberances (Figs 11, 12). This fold is confused
easily with the somewhat fused margin between segments 8
and 9.
Intriguing support for the close affinity of Lypusa and
Amphibatidae position is provided by preliminary results from
an extensive molecular study of the phylogeny of ditrysian
Lepidoptera (M. Mutanen, personal communication). There, L.
maurella appears with convincing support as the sister taxon
of Pseudatemelia josephinae, among over 200 species across
those Lepidoptera families for which eight gene sequences have
been examined.
The question whether Lypusa could be placed in an existing
amphisbatid genus also needs to be addressed. The male
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
Reassessment of Lypusidae 79
Fig. 12. Lypusa tokari. Ventral view of distal end of pupal abdomen.
The arrow points to the transverse fold and protuberances on A8, a
possible apomorphy of Amphisbatidae s.str. Scars of larval legs are
also visible.
Fig. 13. Lypusa tokari. Male genitalia.
genitalia of Lypusa (Fig. 13), however, differ from those
of Amphisbatidae (see also Tok´
ar et al., 2005) to such an
extent that Lypusa cannot be assigned to either of the genera.
The male genitalia of Amphisbatidae are often characterized
by a gnathos as an articulated band with a mesial spined
bulb (Hodges, 1998). The mesial part of the gnathos of
Pseudatemelia flavifrontella and Amphisbatis incongruella has
a bulbous shape and is fused to the basal arms of the gnathos
with a membranous connection and is covered dorsally with
transverse rows of elongate spines. The mesial lobe of the
gnathos of Pseudatemelia josephinae is scobinate with small
diamond-shaped spines dorsally. The gnathos of Lypusa is
somewhat reduced and lacks this mesial lobe and scobination
(90 and 91).
In the cladogram presented by Kaila (2004), the equivalent
node to node 10 (Fig. 6) in our study was supported by
two homoplastic characters: pecten as a row of narrow stiff
scales (2: 1) and mesial part of gnathos scobinate with small
diamond-shaped spines (91: 1). With L. tokari added, node
10 is supported only by character state (2: 1) as the gnathos
of Lypusa lacks scobination. Another difference in male
genital structure between Lypusa and the amphisbatids is
that the characteristic transtilla (95) of Lypusa with the
paired differentiated structures is absent from Amphisbatis
and Pseudatemelia. The male genitalia of Amphisbatis and
the Pseudatemelia species have a weakly sclerotized pair
of lobes in the (disto)lateral margin of the ventral plate of
juxta. Lypusa lacks these (96). In Lypusa, the sacculus is
separate from the costa, in the other amphisbatids it is not
(102). Lypusa has a paired uncus, in the amphisbatids it is
fused as a single projection (83). In the Amphisbatidae, the
coecum penis is absent; in Lypusa it is present as a sac
ventrad of the basal opening of the aedeagus (111). There
are also genital characters that Lypusa shares with some, but
not all, amphisbatid species: 99, 107, 113 and 118. Although
Amphisbatidae share some genitalia structure, much variation
exists within the Pseudatemelia. The female genitalia differ
only by the sclerotization of S8 (132), medially incised in
Lypusa, entirely sclerotized in the other species.
Other morphological differences between Amphisbatidae
and Lypusa include the shape of the second segment of
the labial palpi (7); scale vestiture on the thorax (16); the
posterior joint in the stem of the metafurca (present in the three
amphisbatids, absent from Lypusa) (36); pterostigma (45);
three characters of wing venation (49, 54, 55) and one from
the scale vestiture of the wing (61); and the sternal rod (present
in the Amphisbatidae as a sharply delimited ridge, but coded
as ambiguous in Lypusa due to some differences between the
sexes and individuals) (71). Lypusa shares larval and pupal
characters with some amphisbatids, but due to incomplete data
we were unable to code characters that would be unique to it
and differentiate it with certainty from the others (Appendix 2).
The family status of Amphisbatidae is not settled and
its species composition has been delimited in various ways.
Becker (1984) treated Amphisbatini as a very speciose tribe (48
genera) within the subfamily Depressarinae of the Oecophori-
dae. Of the species included in this concept of Amphis-
batini, many were grouped differently by Kaila (2004). In
Hodges (1974) and Passoa (1995), the tribe Amphisbatini
comprised the genera Machimia, Eupragia and Psilocorsis.
In Hodges (1998), the family Amphisbatidae included 11
genera (65 species). Yet another delimitation is presented
on the web pages of Fauna Europaea (Lvovsky, 2004):
Amphisbatidae (Amphisbatinae (Amphisbatini (Amphisbatis,
Pseudatemelia) Fuchsiini (Fuchsia ) Hypercalliini (Anchinia,
Hypercallia) Telechrysidini (Telechrysis ))). These taxa, exclud-
ing Amphisbatis and Pseudatemelia, are, however, probably
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
80 M. Heikkil¨a and L. Kaila
representatives of Elachistidae s.l. (e.g. Hypercalliinae) as the
male genitalia alone do not justify this systematic position-
ing. On the basis of the male labial palpus structure, Lvovsky
(2005) erected a new tribe Periacmini in the subfamily Amphis-
batinae (Amphisbatidae) for the genera Periacma, Irepacma
and Ripeacma. Recently, Lvovsky (2007) described a fur-
ther amphisbatid genus, Paratemelia, with two species from
Namibia. In the same article, he stated, ‘Anchinia and Hyper-
callia should be transferred to Amphisbatidae’ (on the basis
of male genitalia structure). Kaila (2004) found that the New
World genera, Machimia and Psilocorsis, did not group with
Amphisbatis and the two Pseudatemelia species included in
that study. The obtained grouping, consisting of only the two
last mentioned Palaearctic genera, is thus in agreement with
that of Minet (1990).
Hodges (1998) listed character states defining the Amphis-
batidae. Three acanthi on the female frenulum was one of these,
but Kaila (2004) excluded this character because he observed
that the number of acanthi was closely linked to body size. With
regard to the hair-like SD1 seta on A9 of the larval abdomen, in
Pseudatemelia flavifrontella and Lypusa the seta was not hair-
like, but as thick as the other setae. In Amphisbatis incongruella
and Pseudatemelia josephinae, this character state could not,
unfortunately, be studied. The female retinaculum of Amphis-
batidae was described by Hodges (1998) as diffuse and to have
anteriorly directed scales on CuA or between CuA and R. In the
three amphisbatid species and Lypusa, the anteriorly directed
scales at these locations were absent. The female of Lypusa
has anteriorly directed scales on R.
We concur that for the time being, Amphisbatidae merits
family status and comprises the exclusively Palaearctic genera,
Lypusa, Pseudatemelia and Amphisbatis, i.e. the sack-bearing,
presumably detritivore, members of the oecophorid lineage
sensu Kaila (2004). Restricting the family to these genera
is justified further by the presence of a ventrally located
transverse folding on the pupal abdominal segment 8, which
these genera share (Figs 11, 12). This character could represent
a unique synapomorphy not found elsewhere in Gelechioidea.
Lypusa and the Pseudatemelia species also possess a pair of
protuberances on the distal margin of A8, which is absent in
other gelechioids.
The interrelationships of the genera Pseudatemelia and
Amphisbatis should be scrutinized further. Kaila (2004) found
that Amphisbatis incongruella grouped with Pseudatemelia
flavifrontella and these two form the sister group of Pseu-
datemelia josephinae. In the present study, this resolution is
lost in the consensus tree and the three species form a tetra-
chotomy with L.tokari. Amphisbatis incongruella shows only
four morphological characters by which it differs from the
Pseudatemelia species [one from the wing venation (49: 0),
one from the male genitalia (99: 0), and two from the pupae
(163: 1 and 184: 1)]. The question thus arises, on what basis
are Amphisbatis and Pseudatemelia considered as separate
genera or should they be synonymized? We propose they are
kept separate pending further research. We consider the char-
acters on which the resolution of the Amphisbatidae in Kaila
(2004) was based of uncertain value at this systematic level
and therefore refrain from guessing their interrelationships (cf.
Tok ´
ar et al., 2005).
Notes on the biology of Lypusa
Information on the biology of Lypusa is scarce and has been
thought to relate to L. maurella. The morphology and biology
of the larval stage have been treated even less in the literature.
Now that there is evidence for the existence of several species
in the genus, one has to be cautious because it is uncertain as
to which species the published observations apply.
Lypusa maurella, in the general older concept, is associated
with forests and moorlands supporting heathers and other small
shrubs (Palm, 1976–1977; Dierl, 1996). In Central Europe,
Lypusa has been reported in coniferous forests and even in
maple ‘bushes’ in Tuscany (Zeller, 1852). Lypusa also occurs
in mountainous habitats, such as the Alps and on Mount
Olympus in Greece. Frey (1880) wrote that Zeller met the
species several times in Berg¨
un, Switzerland, flying around
flowers of Pulsatilla alpina. In Finland, L. maurella lives in
forests and marshy peat bogs with heather and other low
vegetation (M. Mutanen, personal communication). In the
southern Urals, it inhabits forest steppe (K. Nupponen, personal
communication). The adult is active diurnally and flies from
May to July (Zeller, 1852; Dierl, 1996). In Denmark, according
to Palm (19761977), the males fly on open terrain between
10 and 11 a.m. looking for females. Sorhagen (1886) described
the flight style as ‘ponderous’.
The case-bearing larvae can be found in March and April
(Sch¨
utze, 1931; Palm, 1976 1977). Palm (19761977) sup-
posed that the larva overwinters and speculated whether the
larval stage takes 1 or 2 years. In the Catalogue of the Lep-
idoptera of Belgium (De Prins & Steeman, 2007), the larval
stage of Pseudatemelia josephinae is said to take 2 years, dur-
ing which period the larva can construct as many as eight cases.
The larva is reported to be able to turn around in its case and
to move in both directions.
We coded the larval diet (character 168) of Lypusa with
a question mark (?). The exact food sources are unknown
and there are no detailed reports on sites where larval or
pupal cases have been found. In some literature sources the
larva of L.maurella is said to feed on lichen, and, for
example, Sch¨
utze (1931) and Sauter (1982) placed the species
in the category of lichenivorous Microlepidoptera. Zagulyaev
(1989) reported that larvae were found ‘on scaly lichens on
stone in April’. Sorhagen (1886) mentioned that the larva
attaches its case to lichens and according to the nature of its
case, also lives on heather. However, G. Palmqvist (personal
communication) finds lichenivory unlikely and sees Calluna,
Erica or Empetrum as more probable food sources. J. Jalava
(personal communication) has indeed found larvae climbing on
Empetrum (Fibiger & Svendsen, 1981) and P. Falck (personal
communication) reported having seen Lypusa larvae feeding
on the fruit of this plant. Palm (1976– 1977) also listed some
other food plants mentioned in different sources, such as plants
belonging to the genus Vaccinium and heather.
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Reassessment of Lypusidae 81
The portable sack of L. tokari (Figs 8, 9) is a piece of
leaf wrapped into a tubular sack, but we have not been able
to identify the plant species. As Sch ¨
utze (1931) has already
pointed out, the piece of leaf the larva wraps around itself is not
cut from heather plants [as Sorhagen (1886) wrote], which have
tiny leaves. The larvae of Amphisbatis are also case-bearers and
feed on Ericaceae (Hodges, 1998), eating the dead leaves of
Calluna at the base of the plant. Pseudatemelia flavifrontella
was coded with a question mark (?) in this study because there
are no published sources as to what kind of food on which
it feeds. However, there is a website report of Pseudatemelia
flavifrontella feeding on withered leaves (Wall, 2008).
In Lepidoptera of Europe (Karsholt & Razowski, 1996:
302), in a personal communication, P. Falck doubted that the
larval case described by Dierl (1991) [sic] is L. maurella, but
could more probably be a Pseudatemelia species. He based
his argument on his personal observation that the first larval
stage of L.maurella did not make a case and the second instar
larva spun a case similar to that of the larva of Psyche betulina.
Unfortunately, he did not manage to follow larval development
further (O. Karsholt, personal communication). In an ex-ovo
rearing of L. maurella performed by M. Mutanen in the summer
of 2008, the female moth laid whitish eggs at the base of leaves
of Empetrum. After eclosion, the first instar larvae each cut a
leaf of the plant and tied it into a portable sack. The larvae fed
on both fresh and dried Empetrum leaves. Unfortunately, the
reared L. maurella larvae died in the autumn and the material
from which the next case would have been made remains
unknown. The larvae had retained the same cases all this time,
which suggests that larval development takes 2 years as with
Pseudatemelia (M. Mutanen, personal communication).
It should be considered a possibility that the first larval stages
of Lypusa construct a different type of case than later instars, or
that the case is simply made from whatever material is available
and therefore may differ depending on rearing conditions or
habitat. The Lypusa larvae observed by Falck had Empetrum
as material from which to construct the case. Dierl (1996)
proposed that the first instars may not build a case, but that
it would be made later, used mainly for pupation.
The specimen we studied is not the same as Dierl (1996)
studied, which was from a Bavarian forest in Germany; ours
was from Tyrol in Austria. The portable case of Lypusa we
studied does not differ from that of the larvae of Pseudatemelia.
However, on the basis of these observations, several reports of
a case-bearing larva (Eversmann, 1844; Sorhagen, 1886; Dierl,
1996) and the specimen we studied (the adult and the case
containing larval and pupal exuviae are preserved on the same
pin), it seems obvious that the larva of Lypusa makes a case
at some stage.
Diagnosis of
Lypusa
Specimens of both known species of Lypusa are shown in
Fig. 1 (L. maurella ) and Fig. 2 (L. tokari ).
Adult
The male and female of Lypusa differ little in exter-
nal appearance, but, as already pointed out by Heinemann
(1870–1877), the base of the antenna of the female appears
thicker due to a denser coverage of scales (Fig. 3).
Head. Scale vestiture on frons smooth, rough on vertex. A
pecten formed as a row of narrow stiff scales present on anten-
nal scape. Antenna nearly touching eye, filiform, ciliate-setose
in male. Lamellate scales on dorsal side of antenna. Some-
what reduced proboscis present with lamellate scales at its base
(Fig. 4). Second segment of labial palpus straight. No bristles
on labial palpus. Maxillary palpus vestigial. Chaetosemata and
external lateral ocelli absent.
Thorax. Both male and female dark brown, scales on
thorax arranged at regular intervals. Wingspan from 13 to
20 mm. Colour of wings of different species of Lypusa in
the grey–brown range, with differences in darkness, lustre
or dullness of the bronzy or brassy sheen. In the literature,
there is disagreement as to the colour of the sheen on the
wings (Eversmann, 1844; Zeller, 1852; Zagulyaev, 1989), but
this may reflect true distinction between the different species.
Fringe concolorous with wings.
Forewing. Retinaculum of male forewing arises from
spur of Sc. Only female with row of narrow anteriorly
directed scales ventrally on R of forewing. Pterostigma absent.
Forewing chorda absent, M1 +2 present at least as a fold
(vestigial, more visible), CuA1 and CuA2 separate, CuP
present at least distally as a well-developed vein, basal forking
of 1A +2A ambiguous.
Hindwing. Male hindwing with one frenulum, female with
varying number of acanthi. Hindwing R1 runs into Sc, Rs being
laterally free, M1 +2 absent, Rs and M1 entirely separate,
M3 arising apical to M2, M3 and CuA1 arising from the same
point.
Metendosternum. Secondary arm and stem of metafurca
present. For further details see the character descriptions
(Appendix 1) and matrix (Appendix 2).
Abdomen. Integument between terga smooth. Sexual dimor-
phism in presence of apodemes on sternum 2; absent in male
(Fig. 15), present in female (Fig. 16).
Male genitalia. (Fig. 13). See also comparison with geni-
talia of Amphisbatidae above. Uncus with paired projection.
Socius present as small group of setae at base of uncus.
Gnathos present, mesially fused, articulated with tegumen
and uncus. Tuba analis projected ventrad of uncus. Transtilla
present as band with paired differentiated structures [‘penincu-
lus’ of Davis & Robinson (1998)]. Sclerotized lobe in distal
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
82 M. Heikkil¨a and L. Kaila
Fig. 14. Lypusa tokari. Female genitalia.
area of sacculus. Sacculus separate. Valvae symmetrical and
mobile. Ventral shield of juxta present as a free sclerotization
ventrad of aedeagus. Juxta not connected to valvae. Aedea-
gus entirely sclerotized. Coecum penis present as sac ventrad
of basal opening of aedeagus. Foldable membrane of fultura
inferior present on aedeagus medially.
Female genitalia. (Fig. 14). Membrane between papillae
anales and segment 8 telescopic. Papillae anales connected
by dorsoposterior sclerotization. Signum with teeth directed
towards margin present in corpus bursae.
Fig. 15. Denuded adult sternum 2 of male Lypusa. Apodemes
absent (69: 0). The arrow points to the obliquely directed lateroanterior
sclerotization; character 76: 1.
Larva
Cuticle densely covered with minute lens-shaped sculpture;
no integumental trichiae present. Secondary setae not present,
except for a group of short triangular spinules around anal
orifice; pinacula of primary setae not erected.
Head. Primary setae only; densely porose, i.e. sculpted
with pits. Ecdysial line does not reach cranial incision. Six
stemmata present; stemmata 5 and 6 positioned more ventrally
than other stemmata; submental pit not present; fusulus of
spinneret tubular, straight; stipular setae relatively long, in front
of sclerotized prementum.
Thorax. Spiracle of T1 rounded; prothorax without sclero-
tization ventrally; all thoracic segments with dorsal sclerotiza-
tion, not extended to spiracle or L group of setae. L group in all
thoracic segments trisetose. Dorsal setae D1 and D2 of meso-
and metathorax approximately on same level, not approximate,
on their own pinacula; D1 shorter than D2. SV group bise-
tose in prothorax, unisetose in meso- and metathorax. V setae
mediocaudad from midpoint of coxae of thoracic legs. Tarsi
bare, tarsal setae not differentiated; axial seta not present in
claw.
Abdomen. Spiracles rounded, spiracle of A8 larger than
those of A1–A7, approximately at same level. Setae L1 and
L2 approximate in A1–A8. Three SV setae in A3 A6. Prolegs
present in A3–A6 and A10; crochets homoideus, arranged as
uniserial ring in A3–A6, as somewhat curved transverse row
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
Reassessment of Lypusidae 83
Fig. 16. Denuded adult sternum 2 of female Lypusa. Apodemes
present (69: 1). The arrow points to the obliquely directed lateroante-
rior sclerotization; character 76: 1.
in A10. No setiform secondary setae in or near prolegs. SV1
of A9 setiform. Anal comb present (Fig. 10).
Pupa
Appendages firmly fused to body, not loosened during
eclosion. Pupa sparsely covered by secondary setae. Flagellae
ventromesially parallel, touching each other in median part,
distally diverging (Fig. 7). Metathoracic legs distally shortly
free. Spiracle of A1 not visible, spiracle of A2 dorsally
surrounded by suture. Spiracles of A2– A7 somewhat erected.
Scars of larval legs present. A8 ventrally with transverse
folding, and distal margin with pair of protuberances (Figs 11,
12). Segments A9 and A10 fused with no visible suture present,
ventrally swollen. Cremaster absent, dense group of hook-
tipped setae at apex of A10.
Conclusions
The results of this study offer a plausible solution to the
systematic position of the genus Lypusa within Gelechioidea.
This is so far the most thorough analysis on the morphology
of Lypusa. Previously, there was little information on the
morphology of the larva and pupa of Lypusa,whichmay
partly explain the difficulty in finding a convincing systematic
position, although the scaled haustellum base of the adult is
quite a straightforward character. Examination of both sexes
and of the immature stages has proved fruitful for yielding
congruent evidence. Surely, however, intact larva and pupal
exuviae would yield more information.
The study of Kaila (2004) resulted in a hypothesis of the
interrelationships within Gelechioidea. As Kaila stated, the
resultant groupings would have been better tested had more
of the immature stages of the basally originating species of the
oecophorid lineage been available and a more comprehensive
taxon sampling undertaken. That the tree resulting from that
study was not altered by the addition of a new species in the
present study is, however, a sign of its stability.
No new characters or character states were sought in this
study. Examination of the morphology of Lypusa did serve,
however, to rephrase and thus clarify some of the character
descriptions (characters 19, 114 and 186) in Kaila (2004).
These did not alter the coding in any way.
No unique synapomorphies for the Lypusa amphisbatid
grouping were found in this study, but together with the infor-
mation from molecular data (M. Mutanen, personal communi-
cation) and unpublished data derived from the immature stages
(see above), this position seems probable. Further research on
the systematics of Amphisbatidae is needed before more can
be said about the relationships among the genera in this family,
and more research on the taxonomy, adult and larval biology,
and geographical range of the different species of Lypusa is
also highly desirable.
Acknowledgements
We express our thanks to the following persons who pro-
vided us with material: Jari Kaitila, Kari Nupponen, Wol-
fram Mey (Museum f¨
ur Naturkunde der Humboldt-Universit¨
at
zu Berlin) and Peter Huemer (Tiroler Landesmuseum Ferdi-
nandeum, Innsbruck), Sabine Gaal-Haszler (Naturhistorisches
Museum Wien), and Bernard Landry (Mus´
eum d’histoire
naturelle Gen`
eve). Marko Mutanen provided us with valuable
information on the biology of adult L. maurella and performed
an ex-ovo rearing of this species. We are grateful to Bengt
˚
A. Bengtsson and G¨
oran Palmqvist for their valuable com-
ments and help with searching for literature. We would also
like to thank one anonymous referee and Don R. Davis for
their helpful comments and especially Ian J. Kitching for the
outstanding in-depth review of our manuscript, which helped
to significantly improve the quality of this work. We also
wish to thank Kevin Tuck (The Natural History Museum, Lon-
don), for searching (albeit unsuccessfully) for L. fulvipennella,
Roy Danielsson (Lund Zoological Museum) for loaning ‘Adela
aethiopella’, and Ole Karsholt (Zoological Museum, Univer-
sity of Copenhagen) and Peter Huemer for their information.
M. Heikkil¨
a is grateful for the help of Marjatta Mikkonen,
Ursina Jud and Sanna Saari. This study was funded by the
Academy of Finland, project 1110906 ‘Phylogeny of Ditrysian
Lepidoptera’.
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Accepted 14 March 2009
First published online 23 August 2009
Appendix 1. Character states coded in the study.
0. Scale vestiture on frons: smooth (0); rough, lateral scales
prolonged, directed mesially or upward forming a ridge in
the meson (1); rough without any direction (2).
1. Vestiture behind antennae: cylindrical hair-like (0); only
long and elongate scales that are parallel-sided but flat and
with longitudinal striae (1); lamellate, spatulate scales that
are basally abruptly widened and with longitudinal striae,
present solely or among narrower scales (2).
2. Pecten in scape of antenna: absent (0); formed as row of
narrow stiff scales (1); formed as tuft of lamellate scales (2).
3. First flagellar article of male antenna: as other articles of
flagellum (0); basally dilated and medially notched (1).
4. Lamellate scale vestiture on haustellum: absent (0); present
(1).
5. External lateral ocellus: absent (0); present (1).
6. Maxillary palpus: five-segmented (0); with four or less
segments (1).
7. Shape of second segment of labial palpus: Sickle-shaped,
ascending (0); straight (1); downward bent (2).
8. Lateral bristles on labial palpus: absent (0); present (1).
9. Inception of antenna: incised in eye (0); nearly touching
eye (1); at a distance from eye so that scales are found at
integument between antenna and eye (2).
10. Pilifer: present with at least few setae (0); absent (1).
11. Mid-cranial sulcus: present (0); absent (1).
12. Parapatagium: present as distinct sclerotized lobe or as
row of scales (0); present as tuft of coremata (1).
13. Lateral extension of pre-episternum: medially with window
(0); fused (1). See Landry (1991: fig. 122) for explanation.
14. Lower sector of precoxal suture: short, not extended past
middle of precoxa (0); long, nearly or quite fused to
anapleural cleft (1). See Landry (1991: fig. 127) for
explanation.
15. Vestiture on lateroposterior area of metascutum: hair-like
scales (0); lamellate spatulate scales (1).
16. Arrangement of vestiture on thorax dorsally: scales evenly
distributed (0); scales distributed to form bands (1).
17. Shape of metascutellum: without lateral corners thus being
evenly rounded (0); with lateral corners with a trace of
lateroanteriorly directed suture (1) (see Kaila, 2004: fig. 6).
18. Internal folding of metascutellum: lining metascutellum lat-
erally and anteriorly (0) (see Kaila, 2004: fig. 6); mesially
interrupted (1).
19. Anterior margin of metathoracic laterophragmata anteri-
orly bent and sclerotized laterad of epinotum: not (0) (see
Kaila, 2004: fig. 7A, B); yes (1) (see Kaila, 2004: fig. 7C).
20. Median ridge in epinotum: absent (0) (see Kaila, 2004:
fig. 7C); present (1) (see Kaila, 2004: fig. 7A, B).
21. Mesial fusion of epinotum: absent (0) (see Kaila, 2004:
fig. 7A, C); present (1) (see Kaila, 2004: fig. 7B).
22. Additional phragmata in epinotum: absent (0); present (1).
23. Posterior suture of inner sclerite of metacoxa dorsally:
present (0) (see Kaila, 2004: figs 6, 13); absent (1).
24. Mesial suture of metacoxa: present (0) (see Kaila, 2004:
fig. 13); absent (1).
25. Anterior suture in the anterior lobe of metacoxal meron :
a suture along anterior margin through to inner sclerite
present (0) (see Kaila, 2004: figs 6, 8); no distinct suture
present except often laterally (1).
26. Anterior lobe of metacoxal meron: present (0) (see Kaila,
2004: fig. 8); absent (1).
27. Obliquely directed suture directed to median incision of
meron: present (0) (see Kaila, 2004: fig. 8); absent (1).
28. Metacoxa ventrally: with oblique incision (0) (see Kaila,
2004: fig. 8); without median incision or the incision
rudimentary (1).
29. Tongue-shaped lobe on posterior part of metacoxa:absent
(0); present (1).
30. Intercoxal lamella between infraepisterna: present as keel
(0) (see Kaila, 2004: fig. 9); present, T-shaped (1); absent (2).
31. Stalk of infraepisternum: free nearly to the median fusion
of infraepisterna (0) (see Kaila, 2004: fig. 9C); fused
forming a stalk-like suture (1) (see Kaila, 2004: fig. 9B);
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
86 M. Heikkil¨a and L. Kaila
free, the anterior margin forming a ‘hook’ towards inter-
coxal lamella, and with a suture from posterior to anterior
margin (2) (see Kaila, 2004: fig. 9A).
32. Joint of metafurca: present as ‘pair of lobes’ fused to
secondary arms of furca or joint between secondary arms
and stem of metafurca (0) (see Kaila, 2004: figs 10, 13,
14); present as point without lateral lobes (1); absent (2).
33. Secondary arm and stem of metafurca: fused to each other
without joint (0); between them a joint (1) (see Kaila,
2004: figs 6, 10, 11, 13).
34. Chiasma between secondary arm of furcasternum and
furcal apophysis: present (0) (see Kaila, 2004: fig. 13);
absent (1) (see Kaila, 2004: fig. 12); secondary arm of
furcasternum and furcal apophysis entirely fused with each
other (2).
35. Stem of metafurca: anteriorly broadened as cusp-like lobes
(0) (see Kaila, 2004: fig. 10); not broadened (1) (see Kaila,
2004: fig. 13).
36. Posterior joint in stem of metafurca: present (0) (see Kaila,
2004: fig. 13); absent (1); posteriorly fused as single
widened plate (2) (see Kaila, 2004: fig. 14).
37. Anteromedian process of metafurca formed as pair of bent
shields: not (0); yes (1) (see Kaila, 2004: fig. 12).
38. Separate distal shields (lamellae) on anteromedian process
of furca: absent (0); present (1) (see Kaila, 2004: figs
10, 11).
39. Distal shield of furcal apophysis: present (0) (see Kaila,
2004: figs 10, 11); absent (1).
40. Ventrodistal hook in metafurca: absent (0); present (1).
41. Distal lobe of lamina of secondary arm of furca: expanded
as cusp-like (0); vestigial or absent (1). See Kaila (1999a,
figs 8–12) for explanation.
42. Dorsoposterior process of metafurcal apophysis:absent
(0); present (1) (see Kaila, 2004: figs 10 12).
43. Shield-like lamella on dorsoposterior process of metafurcal
apophysis or at its place if furcal apophysis is absent:
absent (0); present (1) (see Kaila, 2004: figs 10, 11).
44. Tibia and tarsal articles of metathoracic leg with tufts of
stiff scales: absent (0); present (1).
45. Granulose reinforcement along forewing costa (‘pterostigma’):
absent (0); present (1) (see Kaila, 2004: fig. 15).
46. Retinaculum of male forewing: arising from spur of Sc (0);
arising from Sc (1).
47. Groups of erect scales arising from modified pinacula
that are detectable on upper surface of denuded forewing :
absent (0); present (1).
48. Forewing chorda: present at least vestigially at base and
apex (0); absent (1).
49. Forewing M1 +2: present at least as fold (0); absent or at
most present as small swelling distally at the end of cell
(1).
50. Forewing CuA1 and CuA2: separate (0); stalked (1).
51. Forewing CuP: present at least distally as well-developed
vein, otherwise as fold (0); present at most as fold (1).
52. Basal forking of forewing 1A +2A—costal branch:present
(0); absent or nearly absent (1); inapplicable (?).
53. Basal forking of forewing 1A +2A—dorsal branch:present
(0); absent or nearly absent (1).
54. Hindwing Rs: separate from Sc +R1 (0); basally parallel,
touching but not fused to Sc +R1 (1); basally fused to Sc
+R1 (2); anastomosed to Sc +R1 (3) (see Hodges, 1998:
fig. 9.6P).
55. Hindwing M1 +2: present as fold from base to end of cell
(0); present as fold from base beyond cell (1); absent (2).
56. Hindwing veins Rs and M1 : entirely separate (0); at least
narrowly fused or coalescent (1).
57. Hindwing crossvein M2 – M3 : present, M3 arising apical to
M2 (0); present, M3 arising basal to M2 (1); reduced as
point-like (2); absent as M2 and M3 coalescent (3); absent
as cell open (4).
58. Hindwing M3 and CuA1 : separate (0); coalescent or arising
from same point (1); only one vein present (one of them
absent or veins fully fused) (2); if the crossvein M2– M3
is absent, the inception point of M3 is impossible to detect
(?).
59. Termen of hindwing: straight or convex (0); concave
between M1 and M2 (1).
60. Row of narrow anteriorly directed scales ventrally between
Sc and R of forewing of both male and female (in male
sometimes broader than in female): absent (0); present (1).
61. Row of narrow anteriorly directed scales ventrally on R of
forewing: absent (0); present on both sexes (1); present on
female only (2). See note on character 60.
62. Row of narrow anteriorly directed scales ventrally between
CuA and R of female forewing: absent (0); present on both
male and female (1). See note on character 60.
63. Row of narrow anteriorly directed scales ventrally on CuA
of female forewing: absent (0); present on both male
and female (1); present on female only (2). See note on
character 60.
64. Row of narrow anteriorly directed scales ventrally between
CuA and dorsum of female forewing: absent (0); present on
both male and female (1). See note on character 60.
65. A tuft of stout scales in costal margin of hindwing:absent
(0); present (1).
66. Scales transformed as variably stout spurs in terga:absent
(0); present in single area in each tergum (1); present as
divided areas in each tergum (2).
67. Spines arising without pinacula in anterior margin of terga:
absent (0); present (1).
68. Integument between terga: smooth (0) (see Kaila, 2004:
fig. 16A); microgranulate at least between some terga
(1) (see Kaila, 2004: fig. 16B).
69. Apodeme in sternum 2 of male: absent (0) (see Kaila, 2004:
fig. 17E, F); present (1) (see Kaila, 2004: fig. 17A– D).
70. Apodeme in sternum 2 of female: absent (0); present (1).
71. Sternal rod in sternum 2: absent (0); present as broad
not distinctly sclerotized longitudinal ridge (1) (see Kaila,
2004: fig. 17E); present as sharply delimited narrow long
ridge (2) (see Kaila, 2004: fig. 17A D, F); ambiguous (?).
72. Triangular keel in anterior end of sternal rod of male :
absent (0), present (1) (see Kaila, 2004: fig. 17F).
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
Reassessment of Lypusidae 87
73. Sternal rod anteriorly modified as a sharp-tipped scleroti-
zation fused to skin: no (0), yes (1).
74. T-shaped sclerotization mesially in S2: absent (0); present
(1).
75. Lateroposteriorly directed hook from sternal rod of S2:
absent (0); present (1) (see Kaila, 2004: fig. 17B).
76. Obliquely directed lateroanterior sclerotization in sternum
II : absent (0) (see Kaila, 2004: fig. 17B, F); present
(1) (see Kaila, 2004: fig. 17A, C– E).
77. A group of minute spines without pinacula at sternum 2 :
absent (0); present (1) (see Kaila, 2004: fig. 17E).
78. Posteromedially directed sclerotized ridge from lateral rod
of tergum 1 : absent (0); present as short ridge (1);
present, extended to posterior margin of tergum 1 leaving
less sclerotized windows to the lateroposterior corners of
tergum 1 (2) (see Kaila, 2004: fig. 18).
79. Coremata: absent (0); present as pair of lobes in sternum
2 (1); present as one lobe in sternum 2 or in integument
between sternum and tergum 2 (2).
80. Pleural lobes arising from integument between S8 and T8:
absent (0); present (1).
81. Sternum 8 formed as a lobe covering the basis of valvae,
often to varying extent fused to vinculum: absent (0);
present (1).
82. Presence of uncus: uncus present (0); uncus absent (1).
83. Fusion of uncus: uncus fused as single projection (0);
uncus paired (1).
84. Asymmetry of uncus: uncus symmetric (0); uncus asym-
metric (1).
85. Socius: present as tongue-shaped or variably shaped
setose-free lobe (0); present at most as small group of setae
at ventral margin of tegumen or base of uncus (1).
86. Asymmetry of socii : socii symmetric (0); socii asymmetric
(1).
87. Presence of gnathos: gnathos present (0); gnathos absent
(1).
88. Articulation of basal arms of gnathos with tegumen: artic-
ulated from tegumen (0), fused to tegumen (1), connected
to tegumen with membranous connection (2); inapplicable
or homology unclear (?).
89. Articulation of basal arms of gnathos with uncus: articu-
lated from uncus (0); basally fused to uncus (1); inappli-
cable or homology unclear (?).
90. Shape of mesial part of gnathos: bulbous, fused to basal
arms with membranous connection (0); laterally com-
pressed, downwards directed, fused to basal arms without
distinct limit (1); variably differentiated from basal arms
of gnathos, distally variously pointed-rounded lobe, fused
to basal arms without distinct limit (2); articulated from
basal arms, with sickle-shaped, ascending hook (3); sickle-
shaped, downwards directed, fused to basal arms without
distinct limit (4); rudimentary or inapplicable (?).
91. Scobination of mesial part of gnathos: no scobination
present (0); scobinate with small diamond-shaped spines
dorsally (1); scobinate with large thorns (2); irregularly
covered by elongate spines dorsally (3); covered with trans-
verse rows of elongate spines dorsally (4); inapplicable.
92. Asymmetry of gnathos: gnathos symmetric (0); gnathos
asymmetric (1); inapplicable (?).
93. Division of gnathos: gnathos mesially at least partially
fused (0); gnathos totally divided (1).
94. Tuba analis: projected ventrad of uncus (0); projected
dorsad of uncus (1).
95. Transtilla: absent, or present as variably developed paired
appendix from valval costa (0); present as mesially differ-
entiated band often resembling a gnathos (1); present as
band with paired differentiated structures (2).
96. Pair of sclerotized lobes in lateral or distolateral margin
of ventral plate of juxta: absent (0); present as setose
weakly sclerotized lobe (1); present as expanded strongly
sclerotized lobe with stout spurs; this lobe is sometimes to
varying extent fused to valva (2).
97. Pair of lobes in distal margin of ventral plate of juxta:
absent (0); present (1).
98. Sclerotized lobe in basal or medial area of sacculus:absent
(0); present, projected towards costa (1).
99. Sclerotized lobe in distal area of sacculus:absent(0);
present (1).
100. Free sclerotized lobe from sacculus distally: absent (0);
present (1).
101. Costa: not separate from rest of valva (0); costa separate
(1).
102. Sacculus: not separate (0); separate (1). See comment of
character 101.
103. Proximal flange in valva: absent (0); present (1). See
Adamski & Brown (1989) for definition.
104. Stout distally multilobed setae in valva: absent (0); present
(1).
105. Mobility of valvae: valvae mobile (0); movement restricted
(1).
106. Asymmetry of valvae: valvae symmetric (0); valvae asym-
metric (1).
107. Ventral shield of juxta : absent (0); present as free scle-
rotization ventrad to aedeagus (1); present, surrounding
aedeagus (2); present, connected to vinculum (3).
108. Differentiated dorsal shield of juxta: absent (0); present as
lobe or pair of lobes connected to ventral shield of juxta
(1); present, surrounding aedeagus (2). See Kaila (1999a,
b) for definition.
109. Connection between juxta and valva : juxta not connected
to valva (0); narrow sclerotized valval process connecting
valval costa and juxta present (1); valva broadly connected
to juxta (2).
110. Sclerotization of aedeagus: aedeagus entirely sclerotized
(0); aedeagus at least partly membranous (1).
111. Coecum penis: absent (0); present as sac ventrad to basal
opening of aedeagus (1); present as sclerotized usually
spirally coiled lobe (2).
112. Sclerotized manica: absent (0); present (1).
113. Sclerotized anellus: absent (0); present (1).
114. Foldable membrane of fultura inferior overlain to aedea-
gus: present on aedeagus medially (0); very extensible,
present on aedeagus posteriorly (1).
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
88 M. Heikkil¨a and L. Kaila
115. Aedeagus ankylosed by median plate of juxta :no(0);yes
(1).
116. Aedeagus ankylosed by juxta lobes: no (0); yes (1).
117. Aedeagus ankylosed by dorsal shield of juxta:no(0);yes
(1).
118. Aedeagus ankylosed by anellus: no (0); yes (1).
119. Membrane between papillae anales and segment 8 in
female: short and stiff, not or minimally extensible (0);
extensible (telescopic) (1).
120. Ductus bursae and corpus bursae: fully developed (0);
ductus bursae and corpus bursae reduced to minute ducti
(1).
121. Internal granulae in ductus bursae: absent (0); present (1).
122. Signum in corpus bursae: present (0); absent (1).
123. Number of signa: one (0); two, symmetric (1); two,
asymmetric, indicating a possibility for their independent
origin (2).
124. Structure of signum: simple sclerotized plate (0); with teeth
directed to margin of signum (1); with numerous teeth
directed towards centre of signum (2).
125. Curved large hook in signum: absent (0); present (1).
126. Signum folded mesially:no(0);yes(1).
127. Signum with sclerotized tooth outside corpus bursae:no
(0); yes (1).
128. Orientation of basal rim of papillae anales, relative to tran-
section of abdomen, in lateral view : straight or diagonal to
anterodorsal direction (0); diagonal to anteroventral direc-
tion (1); diagonal to anteroventral direction, and ventrally
extended by anterior lobes (2).
129. Connection of papillae anales: papillae anales separated
by membrane dorsally (0); papillae anales connected by
dorsoposterior sclerotization (1).
130. Lateral fusion of S8 and T8 : S8 and T8 separate sclerotized
plates (0); S8 and T8 laterally fused and together form solid
ring (1).
131. Sclerotization of T8 : T8 entirely sclerotized (0); with
median incision or entirely membranous (1).
132. Sclerotization of S8 : S8 entirely sclerotized (0); medially
incised or entirely membranous (1).
133. Integument between S8 and S9 : not extensible (0); pro-
longed as extensible telescope (1).
134. Shape of ductus seminalis: tubular (0); basally dilated as
sack (1).
135. Sternal rod on ovipositor: absent (0); present (1).
136. Cylindrical often protruded tube in ostium bursae:absent
(0); present (1).
137. Lobe in front of ostium bursae: absent (0); present (1).
138. Lobe behind ostium bursae: absent (0); present (1).
139. Pubescence on pupa:absent(0);present(1).
140. Shape of pupal clypeus and frons: unmodified (0); beak-
shaped (1).
141. Pupal labial palpus: exposed (0); concealed (1).
142. Transverse ridges in scape of pupal antenna : absent (0);
present (1).
143. Pupal metathoracic leg: distally exposed (0); distally
concealed (1).
144. Pupal fore femur: exposed (0), concealed (1).
145. Pupal antennae: mesially separate (0); mesially touching
each other (1).
146. Mesial swelling on pupal mesothorax : absent (0); present
(1).
147. Tubercles in mesothorax : absent (0); present (1).
148. Mesothoracic spiracle: as other spiracles (0); with protu-
berance (1).
149. Pupal hindwing: exposed to abdominal segment 3 (0);
exposed at most to abdominal segment 2 (1).
150. Pupal abdominal intersegment 3 –4 : movable (0); immo-
bile (1).
151. Transverse rows of spines in anterior margin of at least
some segments of pupal abdomen: absent (0); present (1).
152. Transverse rows of spines in posterior margin of at least
some segments of pupal abdomen: absent (0); present (1).
153. Lateral condyles on pupal abdominal segments preventing
their lateral movement : absent (0); present (1).
154. Dorsal and ventral condyles on pupal abdominal seg-
ments preventing their dorsoventral movement: absent (0);
present (1).
155. Dorsal ridge on pupal abdomen: absent (0), present (1).
156. Lateral ridge on pupal abdomen: absent (0); present (1).
157. Dense row of setae at posterior margin of abdominal
segment 7 : absent (0); present (1).
158. Concave emargination dorsolaterally at posterior margin
of abdominal segment 7 : absent (0); present (1).
159. Pupal abdominal segment 8 : parallel-sided (0); ventrally
narrowed (1).
160. Pupal segments 9 +10 : cylindrical as other segments (0);
ventrally flattened as plate (1).
161. Pair of spur groups in pupal segment 9 ventrally:absent
(0); present (1).
162. Pupal legs ventrally at abdominal segment 9 ventrally;
sometimes only present as pair of swellings: absent (0);
present (1).
163. Lateral spine at pupal segment 10 : absent (0); present (1).
164. Pupal cremaster: absent or at most present as small group
of setae (0); present as setose appendix (1).
165. Behaviour of pupa prior to eclosion: pupa protrudes from
cocoon by active movement and with aid of transverse
spine band of terga (0); pupa does not move out of cocoon
(1).
166. Pupation: concealed, in cocoon, soil, litter, foliage or
within stem (0); exposed (1).
167. Silken girdle around abdomen to hold pupa attached to
pupation substrate: absent (0); present (1).
168. Larval diet: on green plants (0); on dead material or fungi
(1); predator on other insects (2).
169. Portable sack on larva: absent (0); present (1).
170. Submental pit at ventral side of larval head:absent(0);
present as rounded or oval pit (1); present as sclerotized
pair of grooves (2).
171. Ecdysial line of adfrontal sclerites: reaches cranial incision
(0); does not reach cranial incision (1).
172. Stipular setae of spinneret: minute (0); long but thin (1);
long and stout (2).
173. P setae of head: two setae present (0); one seta present (1).
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
Reassessment of Lypusidae 89
174. L group of prothorax: with three setae (0); with two setae
(1); with one seta (2).
175. Arrangement of two anterior setae in L group of prothorax:
oblique, anterior seta dorsad of posterior seta (0); horizon-
tally (1).
176. Position of L3 of prothorax: horizontally or dorsolaterad
of L2 (0); ventrolaterad of L2 (1).
177. Secondary setae in SV group of meso- and metathorax:
absent (0), present (1).
178. Sclerotization ventrally in T1 :absent(0);present,paired
(1); present, single (2).
179. Shape of metathoracic leg: not different from other thoracic
legs (0); pistol-shaped (1).
180. Ring-shaped pinaculum ring around SD1 leaving non-
sclerotized area around seta at A1 –A8 : absent (0); present (1).
181. Setae L1 and L2 of A1– A8 : setae approximate, in same
pinaculum (0); setae distant, in separate pinacula (1).
182. Arrangement of crochets on prolegs of A3– A6 : multiserial
(0); uniserial (1).
183. Arrangement of crochets on prolegs of A3–A6 :asfullor
nearly full circle (0); as lateral penellipse (1); as mesial
penellipse (2); as pair of transverse rows (3).
184. Setae of SV group of prolegs of A3 A6 : at one pinaculum
(0); at separate pinacula (1); SV2 in its own pinaculum (2);
SV setae absent (3).
185. Secondary setae on prolegs of A3– A6 : absent (0); present
at least on some prolegs (1).
186. Shape of SD1 at A9: as thick as the other setae (0); hair-like
(1); inapplicable (?).
187. Arrangement of D1, D2 and SD at A9 :D1anteriortoD2
(0); in vertical row (1).
188. Size of D1 at A9 : as D2 (0); smaller than D2 (1).
189. Arrangement of D1 and D2 at A9 : D1 ventrad of D2 (0);
D1 dorsad of D2 (1).
190. Secondary setae in SV group of anal legs:absent(0);
present (1).
191. Anal comb: absent (0); present (1).
192. Larva with swellings (verrucae) on which numerous sec-
ondary setae: absent (0); present (1).
Appendix 2. Data matrix of character states for Lypusa
tokari, outgroup taxon Scardia boletella and the amphisbatids
Amphisbatis incongruella, Pseudatemelia josephinae and Pseu-
datemelia flavifrontella. See Kaila (2004) for character states
of the other 149 taxa. The full matrix may also be obtained
from the authors.
1111111111 2222222222 3333333333 4444444444
0123456789 0123456789 0123456789 0123456789 0123456789
Scardia boletella 2010000111 0000001000 0000010000 1120010001 0100001000
Amphisbatis incongruella 0110101001 0101001010 0000010100 0101010001 0110010010
Pseudatemelia josephinae 0110101001 0101001010 0000010100 0101010001 0110010011
Pseudatemelia flavifrontella 0110101001 0101001010 0000010100 0101010001 0110010011
Lypusa tokari 0110101101 0101000010 0000010100 0101011001 0110000010
5555555555 6666666666 7777777777 8888888888 9999999999
0123456789 0123456789 0123456789 0123456789 0123456789
Scardia boletella 0000000100 0000000001 1100000000 00???????? ?0?0?00000
Amphisbatis incongruella 0010000010 0000000000 1200001000 0000010000 0400001000
Pseudatemelia josephinae 0010000010 0000000000 1200001000 0000010000 2100001001
Pseudatemelia flavifrontella 0010000010 0000000000 1200001000 0000010000 0400001001
Lypusa tokari 00??320010 0200000000 1?00001000 0001010000 ??00020001
1111111111 1111111111 1111111111 1111111111 1111111111
0000000000 1111111111 2222222222 3333333333 4444444444
0123456789 0123456789 0123456789 0123456789 0123456789
Scardia boletella 0000000000 0000100001 001?????0? 0000010000 0000000000
Amphisbatis incongruella 0000000200 0000000001 0000100001 0001000000 0100010000
Pseudatemelia josephinae 0000000200 0000000001 0000100001 0001000000 0100010000
Pseudatemelia flavifrontella 0000000100 0001000011 0000100001 0001000000 0100010000
Lypusa tokari 0010000100 0100000001 0000100001 0011000000 ???0010000
1111111111 1111111111 1111111111 1111111111 111
5555555555 6666666666 7777777777 8888888888 999
0123456789 0123456789 0123456789 0123456789 012
Scardia boletella 0110000000 0001000010 000010?000 0110000010 000
Amphisbatis incongruella 1000000001 0001010011 0100000000 001010???? ?10
Pseudatemelia josephinae 1000000001 0100010011 0100000000 001000???? ?10
Pseudatemelia flavifrontella 1000000001 00000100?1 0100000000 0010010110 110
Lypusa tokari 1000000001 00000100?1 0100000000 0010000??0 010
©2009 The Authors
Journal compilation ©2009 The Royal Entomological Society, Systematic Entomology,35, 71–89
... Recently the relationships of the genus Lypusa with the genera Amphisbatis and Pseudatemelia from the family Amphisbatidae were revealed being based on a similar construction of larval case (made from a whole piece of leaf), densely porose larval head, and modified pupal abdominal segment VIII having a transverse fold and protuberances. Thereby, the volume of this family was increased up to three genera (Heikkilä & Kaila, 2010), and the name Amphisbatidae was synonymised with Lypusidae (Nieukerken et al., 2011). ...
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... and Staudinger & Wocke (1871). Lypusa(Zeller, 1852) is a Palaearctic moth genus which, until recently, was supposed to comprise a single widespread species L. maurella (Denis & Schiffermüller, 1775)(Heikkila & Kaila 2010). Elsner et al. (2008 established that it actually comprises a species group.Elsner et al. (2008) found on Mt Nanos the species L. tokari, only few kilometres away from where Mann (1854) recorded L. maurella. ...
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A checklist of the Microlepidoptera species recorded in Slovenia is presented. Each entry is accompanied by complete references, and remarks where appropriate. Until now, the data on Microlepidopteran fauna of Slovenia have not been compiled, with the existing information scattered in literature, museums and private collections throughout Europe. The present checklist is based on records extracted from 290 literature sources published from 1763 (Scopoli) to present. In total, 1645 species from 56 families are listed. Izvleček. SEZNAM METULJČKOV (MICROLEPIDOPTERA) SLOVENIJE – Predstavljen je seznam vrst metuljčkov, zabeleženih v Sloveniji. Za vsako vrsto so podane reference, kjer je bilo smiselno, pa tudi komentar. Do sedaj podatki o metuljčkih Slovenije še niso bili zbrani, obstoječi podatki pa so bili razpršeni v različnih pisnih virih, muzejskih in zasebnih zbirkah po Evropi. Predstavljeni seznam temelji na podatkih iz 290 pisnih virov, objavljenih od 1763 (Scopoli) do danes. Skupaj je navedenih 1645 vrst iz 56 družin.
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The superfamily Gelechioidea (Lepidoptera: Obtectomera) has a high species diversity. It consists of more than 18,400 described species and has a global distribution. Among it, large numbers of species were reported to be economically important to people's production and life. However, relationships among families or subfamilies in Gelechioidea have been exceptionally difficult to resolve using morphology or single gene genealogies. Multiple gene genealogies had been used in the molecular phylogenetic studies on Gelechioidea during the past years, but their phylogenetic relationships remain to be controversial mainly due to their limited taxa sampling relative to such high species diversity. In this paper, 89 ingroup species representing 55 genera are sequenced and added to the data downloaded from GenBank, and six species representing four closely related superfamilies are chosen as outgroup. The molecular phylogeny of Gelechioidea is reconstructed based on the concatenated data set composed of one mitochondrial marker (COI) and seven nuclear markers (CAD, EF‐1ɑ, GAPDH, IDH, MDH, RpS5, wingless). The phylogenetic results, taking into consideration of the comparative morphological study, show that the clade of Gelechioidea is strongly supported and separated from other superfamilies, which further proves its monophyly. Five families are newly defined: Autostichidae sensu nov., Depressariidae sensu nov., Peleopodidae sensu nov., Ashinagidae sensu nov. and Epimarptidae sensu nov. Meanwhile, a monophyletic “SSABM” clade considered to be closely related is proposed for the first time, consisting of Stathmopodidae, Scythrididae, Ashinagidae, Blastobasidae and Momphidae. Moreover, geometric morphometric analyses using merged landmark data set from fore and hind wings of 118 representative species are conducted. The phenetic tree shows that the monophyly and phylogenetic relationships correspond with the results of molecular phylogeny largely, which well proves its importance and potential application in both phylogenetic reconstruction and species identification.
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A definitive species list is the foundation of biodiversity and conservation work. As we deal with massive climatic changes in the Anthropocene, knowing which species make up our diverse ecosystems will be critically important if we wish to protect and restore them. The Lepidoptera, moths and butterflies, are the fourth-largest insect order in terms of global diversity, with approximately 158,000 described species. Here we report the distributions of 5431 species that occur in Canada and Alaska, as well as 53 species that have been reported from the region but not yet verified. Additionally, 19 species are listed as interceptions or unsuccessful introductions, and 52 species are listed as probably occurring in the region. The list is based on records from taxonomic papers, historical regional checklists, and specimen data from collections and online databases. All valid species and their synonyms, and all Nearctic subspecies and synonyms are included, except for butterfly subspecies (and their synonyms) that have never been reported from the region. The list is presented in taxonomic order, with the author, date of description, and original genus provided for each name.
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For the kingdom Animalia, 1,552,319 species have been described in 40 phyla in a new evolutionary classification. Among these, the phylum Arthropoda alone represents 1,242,040 species, or about 80% of the total. The most successful group, the Insecta (1,020,007 species), accounts for about 66% of all animals. The most successful insect order, Coleoptera (387,100 species), represents about 38% of all species in 39 insect orders. Another major group in Arthropoda is the class Arachnida (112,201 species), which is dominated by the mites and ticks (Acari 54,617 species) and spiders (43,579 species). Other highly diverse arthropod groups include Crustacea (66,914 species), Trilobitomorpha (19,606 species) and Myriapoda (11,885 species). The phylum Mollusca (117,358 species) is more diverse than other successful invertebrate phyla Platyhelminthes (29,285 species), Nematoda (24,783 species), Echinodermata (20,509 species), Annelida (17,210 species) and Bryozoa (10,941 species). The phylum Craniata, including the vertebrates, represents 64,832 species (for Recent taxa, except for amphibians): among these 7,694 described species of amphibians, 31,958 species of "fish" and 5,750 species of mammals.
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The subfamily Periacminae is characterized by the structure of the valva, with sacculus sharply separated from the remaining part of the valva. The subfamily includes two tribes: Periacmini Lvovsky, 2005 (with four genera: Periacma Meyrick, 1894; Irepacma Moriuti, Saito et Lewvanich, 1985; Ripeacma Moriuti, Saito et Lewvanich, 1985, and Epiracma Wang et Li, 2005) and Meleonomini Lvovsky, trib. n. with two genera Meleonoma Meyrick, 1914 and Phaulolechia Diakonoff, 1951. A key to the genera based on the characters of the moths’ appearance and genitalia is provided. The following new synonymy is established: Meleonoma Meyrick, 1914 (= Acryptolechia Lvovsky, 2010, syn. n.), and Phaulolechia Diakonoff, 1951 (= Variacma Wang, 2006, syn. n.). Four new species of Meleonoma are described from Nepal: M. nepalella sp. n., M. jubingella sp. n., M. dierli sp. n., M. montana sp. n.; and M. meyricki sp. n. is described from China. Eight new combinations are established: Meleonoma malacobyrsa (Meyrick, 1921) comb. n., Meleonoma facunda (Meyrick, 1910), comb. n., Meleonoma torophanes (Meyrick, 1935), comb. n. (all from Acryptolechia), Meleonoma flavimaculata (Christoph, 1882), comb. n. (from Euteles), Meleonoma peditata (Wang, 2006), comb. n. (from Cryptolechia), Phaulolechia saccata (Wang, 2006), comb. n., Phaulolechia subrotunda (Wang, 2006), comb. n., and Phaulolechia isotyra (Meyrick, 1938), comb. n. (all from Variacma). Meleonoma facialis Li et Wang, 2002 originally described from China was recorded in the Russian Far East, Nepal and Indonesia (Java).
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Gelechioidea are one of the most species rich and least studied superfamilies of Lepidoptera. We examine the interrelationships within the superfamily using the densest taxon sampling to date, combined with the most extensive ever morphological and molecular character data. We perform partitioned and combined analyses using maximum likelihood, Bayesian and parsimony approaches. The combined dataset consists of 155 exemplar species of Gelechioidea, representing nearly all subfamilies recognized in recent classifications. Parsimony analyses are performed with a dataset including 28 additional terminal taxa with only morphological data available. We use eight genes with a total of 6127 bp, and morphological data with 253 characters derived from larval, pupal, and adult morphology. The analyses of combined data yield more resolved trees and significantly better-supported groupings than either dataset when analysed alone. The recurrent monophyletic groupings in all our modelbased analyses support a revision of the family classification. Deeper relationships vary between analyses and data partitions, leaving them ambiguous. The place of the root remains a challenge for future research. We propose a revised classification and suggest the division of Gelechioidea into 16 families. We redefine Depressariidae Meyrick, 1883 for a monophylum that includes Acriinae, Aeolanthinae, Cryptolechiinae, Depressariinae, Ethmiinae, Hypercalliinae, Hypertrophinae, Peleopodinae, Oditinae, Stenomatinae, Carcina, and a diversity of predominantly New World taxa previously excluded from Lypusidae (Amphisbatidae s. authors) but left without family position. A monophyletic Oecophoridae s. s., including Deuterogoniinae and Pleurotinae, is obtained for the first time with significant support. Elachistidae s. l. is found to be polyphyletic, and Elachistidae is restricted to comprise Agonoxeninae, Elachistinae, and Parametriotinae. Batrachedridae are polyphyletic, with several genera pending further study. Apart from the core Batrachedra, the taxa previously included in this family are grouped in an expanded Pterolonchidae, together with Coelopoetinae and Syringopainae. Lypusidae s. s. and Chimabachidae form a monophylum; Chimabachinae is united with Lypusidae as a subfamily, stat. n. Our results contradict the subfamily classifications of several families, notably Lecithoceridae and Autostichidae, but due to insufficient sampling of taxa we refrain from comprehensive taxonomic conclusions on the subfamily level, and encourage focused studies to resolve these groups.
Article
The broad-winged moths (Oecophoridae sensu lato) belong to the family complex Gelechiiformes with 4 (Kuznetzov and Stekolnikov, 2001) or 6 (Sinev, 1992) superfamilies. The classification of these moths was rather constant in the XX century, but at the turn of the century most researchers concluded that broad-winged moths in the traditional sense (Oecophoridae sensu lato) were a polyphyletic group and that a major revision of their classification was necessary. Many classifications of these families have been proposed recently (Common, 1990; Minet, 1990; Scoble, 1992; Sinev, 1992; Leraut, [1993], 1997; Heppner, 1998; Hodges, 1999; Kuznetzov and Stekolnikov, 2001; Lvovsky, 2002; Kaila, 2004; Bucheli and Wenzel, 2005). This communication presents one more classification variant, based on analysis of the results of the preceding researchers and personal views of the author.
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This is the first comprehensive, reliable, well-illustrated book covering the enormous diversity of Australian moths, summarising our knowledge of them by the acknowledged experts in the field. The text includes nomenclature and a wealth of information on distribution, larval food plants, and the fascinating behaviour of these often colourful insects. There are authoritative accounts of moth structure, their life history, biology, population control, economic significance, evolution and geographical distribution. Additional features include a section on collecting and studying moths, a glossary, a detailed index and an extensive list of references.
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Genera and previously described species of Nearctic Scythrididae are revised for the first time, based on the study of adult structures. About 90 percent of the Nearctic fauna known in collections consists of undescribed species. The supraspecific taxa treated in this work encompass less than half of the Nearctic species diversity. Only six new species are described, all within the largest and structurally most diverse genus. The status of all nominal species is revised. Valid species are redescribed and their features illustrated. General problems in the systematics of the Scythrididae are discussed. A description of adult features of the family Scythrididae is providad. Extra-limital genera are briefly reviewed. A key to the Nearctic genera and informal supraspecific lineages is provided.Six genera, including three new, are treated: Areniscythris Powell, 1976, Arotrura Walsingham, 1888, Asymmetrura gen. nov., Neoscythris gen. nov., Rhamphura gen. nov., and Scythris s. str. Hübner, [1825]. Areniscythris includes a single described species, Areniscythris brachypteris Powell, but is defined more broadly to account for a number of undescribed species. Arotrura is divided into nine informal species groups with the following included species: Arotrura atascosa sp. nov., Arotrura balli sp. nov., Arotrura divaricata (Braun) comb, nov., Arotrura eburnea Walsingham, Arotrura formidabilis sp. nov., Arotrura hymenata sp. nov., Arotrura longissima sp. nov., Arotrura oxyplecta (Meyrick) comb, nov., Arotrura powelli sp. nov., and Arotrura sponsella (Busck) comb. nov. Asymmetrura includes: Asymmetrura albilineata (Walsingham) comb. nov., Asymmetrura graminivorella (Braun) comb. nov., Asymmetrura impositella (Zeller) comb. nov. and type species, Asymmetrura matutella (Clemens) comb, nov., Asymmetrura reducta (Braun) comb, nov., and Asymmetrura scintillifera (Braun) comb. nov. Neoscythris includes: Neoscythris confinis (Braun) comb, nov., Neoscythris euthia (Walsingham) comb. nov., Neoscythris fissirostris (Meyrick) comb. nov. and type species, and Neoscythris planipenella (Chambers) comb. nov. Rhamphura includes: Rhamphura altisierrae (Keifer) comb, nov., Rhamphura ochristriata (Walsingham) comb. nov. and type species, Rhamphura perspicillella (Walsingham) comb. nov., Rhamphura suffusa (Walsingham) comb. nov., and the extra-limital Rhamphura immunis (Meyrick) comb. nov. from Peru. Scythris s. str. includes: Scythris immaculatella (Chambers) rev. stat., Scythris limbella (Fabricius), Scythris mixaula Meyrick, Scythris trivinctella (Zeller), and Scythris ypsilon Braun. A further eight species are phylogenetically distinct from Scythris s. str. but provisionally are only assigned to five informal monophyletic lineages until their cladistic relationships are more firmly established. These are: the Scythris basilaris lineage including Scythris basilaris (Zeller), Scythris eboracensis (Zeller), and Scythris fuscicomella (Clemens); the Scythris interrupta lineage including Scythris interrupta Braun; the Scythris inspersella lineage including Scythris inspersella (Hübner) and Scythris noricella (Zeller); the Scythris anthracina lineage including Scythris anthracina Braun; and the Scythris charon lineage including Scythris charon Meyrick. Three species are incertae sedis: Scythris inornatella (Chambers) comb, nov., Scythrispilosella (Zeller), and Scythris piratica Meyrick. Coleophora albacostella Chambers and Coleophora inornatella Chambers are transferred from the Coleophoridae. Scythris arizoniella (Kearfott) is transferred to the Coleophoridae [ Coleophora arizoniella (Kearfott) comb. nov.].The following new synonymy is proposed: Colinita Busck, 1907 = Arotrura Walsingham, 1888; Gelechia aterrimella Walker, 1864 and Scythris epilobiella McDunnough, 1942 = Scythris inspersella [Hübner, (1817)]; Scythris magnatella Busck, 1904 = Scythris noricella (Zeller, 1843); Scythris pacifica McDunnough, 1927 = Scythris immaculatella (Chambers, 1875); Coleophora albacostella Chambers, 1875 and Scythris hemidictyas Meyrick, 1928 = Neoscythris planipenella (Chambers, 1875).A cladistic definition of the family is presented for the first time. The monophyly of the Scythrididae is supported by the following synapomorphies: very narrow ductus bursae, broad ductus seminalis anastomosed with the oviduct and the corpus bursae, lack of signum, unique shape of the apophyses of the metathoracic furca, tarsomeres 1–4 with two subapical spurs, aedeagus ankylosed, and origin of forewing veins R4 and R5 on a common stalk with R4 extended to the costa and R5 to the termen. Relationships of the Scythrididae within the Gelechioidea are discussed. Based on the cladistic analysis of 52 structural characters, phylogenetic relationships of supraspecific taxa are inferred. Two cladograms, one for the genera and one for the species groups of Arotrura , are presented and used in deriving the classification.