Olethreutine Moths of Australia: (Lepidoptera: Tortricidae)

By Marianne M. Horak and with contributions by Furumi Komai

Olethreutine moths often have fruit-boring larvae and this economically important group includes many horticultural pests such as codling moths, Oriental fruit moths and macadamia nut borers. This volume is the first reference to describe the 90 olethreutine genera present in Australia.

It provides generic definitions, a key to genera, generic descriptions, and illustrations of adults, heads, venation, genitalia of both sexes and other diagnostic structures of all genera. Summaries of biology and distribution and a checklist for all named Australian species are given for each genus. Importantly, it includes a comprehensive reorganisation of olethreutine classification, based on generic revisions, with a worldwide impact.

The volume contains copious illustrations (two species per genus where possible) to convey generic concepts, and to allow identification of this economically important group. Nearly all olethreutine genera present in Australia extend into Asia and beyond, so the book will be relevant to horticultural pests throughout Asia, and crucial to an understanding of olethreutine evolution worldwide.

The diverse Australian olethreutine fauna is particularly rich in enarmoniine and grapholitine genera, several new to science and adding significantly to the concepts of these two tribes. Given the wealth of biological information, the book will be important for ecological work on phytophagous insects well beyond Australia.

• Language: English
• Publisher: CSIRO PUBLISHING
• Release date: Jun 28, 2006
• ISBN: 9780643099371

Book preview

1. Phylogeny of the Olethreutinae

The Tortricidae are the only family in the homogeneous and undoubtedly monophyletic superfamily Tortricoidea. The group is easily recognisable by a combination of character states such as: an unscaled proboscis, presence of ocellus and chaetosema, porrect or upcurved labial palpi (but never horn-shaped with a long, slender, curved terminal segment), minute maxillary palpi generally not visible on undissected specimens, usually rough head scaling but with the scaling on the lower frons very short and upwardly oriented, ‘tortricoid’ sternum 2 with broad apodemes and at most very weak sternal rods, and flat, leaf-like ovipositor lobes. The flat ovipositor lobes of the female are a derived feature (apomorphy) characterising the family (Dugdale 1988; Horak and Brown 1991), with only few exceptions that are due to secondary modification associated with specialised oviposition. Upwardly oriented scaling on the lower part of the frons and the anal fork with straight prongs, found in most tortricid caterpillars, are possible further apomorphies for the family. The anal fork serves to eject frass from the shelter and tends to be lost in groups where caterpillars feed as borers.

Three subfamilies, Tortricinae, Chlidanotinae and Olethreutinae, have become generally accepted within the Tortricidae (Horak 1998). The latter two are considered monophyletic based on several unique, derived character states whereas the Tortricinae are paraphyletic, comprising highly derived groups as well as very generalised, plesiomorphic taxa. All Olethreutinae share two unique features: 1, highly modified male genitalia with juxta and caulis fused into a single structure instead of loosely hinged as in other tortricids, and 2, each segment of the antennal shaft with only a single row of scales instead of two rows. Two rows of scales per antennal segment are considered to be the groundplan state for the Ditrysia (Robinson and Nielsen 1993). Some tortricine Sparganothini (not occurring in Australia) also have antennae with a single ring of scales per segment, but they differ in the details of their structure and are clearly independently derived. Olethreutines can usually also be recognised by the presence of a cubital pecten (Fig. 14), but this structure is absent in a few olethreutine genera and present in some generalised tortricines. Whilst male olethreutine genitalia are highly derived, the wing venation in this subfamily is more generalised overall than in both the Tortricinae and Chlidanotinae. M-stem and chorda are retained at least in some species of most olethreutine genera, in contrast to the large majority of the other two subfamilies. The historical development of the tribal classification of the Olethreutinae is discussed in detail in the general remarks for each of the six tribes here recognised.

The aim of this revision was primarily to define the olethreutine genera present in Australia in cladistic terms, based on derived characters that would provide clear generic limits. This will allow identification at least to genus and the assignment of unnamed and yet-to-be-discovered species to the correct genus. Examination of the type species and as wide a range of species as possible for each genus, including all taxa present in Australia, was the necessary basis for such a study. On the other hand, there was no need to resolve difficult species complexes as they do not impinge on generic definitions. Generic boundaries are subjective depending on the chosen level of inclusiveness even in a cladistic framework based on monophyletic groupings that comprise all the progenies of the taxon that acquired a given derived feature. An entirely pragmatic approach was chosen, guided by practicalities and accepted usage wherever possible, resulting in the fewest possible nomenclatural changes. After the genera were individually assembled and defined they were used as building blocks for a higher classification by searching for synapomorphies, i.e. shared derived characters, linking them into groups. This process was very successful for the Olethreutini, building on the monumental monograph of the south-east Asian fauna by Diakonoff (1973), and for the Eucosmini. It was less satisfactory for the Enarmoniini. Though several pairs of sister genera were identified, it was not possible to identify cohesive higher groupings, and the genera were arranged in alphabetical order within the tribe. Komai (1999) had already provided a phylogeny for the Palaearctic Grapholitini and extended this for all Australian genera of the tribe.

Table 1. Characters used in cladistic analysis

A cladistic analysis using WinClada (Nixon 1999) and NONA (Goloboff 1999) was run on a matrix (Appendix 1) of 126 morphological characters (Table 1) for 73 olethreutine genera and two outgroup taxa. The plesiomorphic tortricine Williella sauteri Horak from New Caledonia and Templemania animosana Busck (Anacrusini) from Mexico (coll. J. A. Powell) were used as outgroups. All Australian olethreutine genera were included except for the Grapholitini as work on their revision was not based on a DELTA matrix. Also, Komai (1999) had already demonstrated their monophyly and provided a phylogeny for the tribe. Grapholita zapyrana (Meyrick) was subsequently scored as a single exemplar species to include at least one representative of the tribe. All genera were scored to represent the character variation across the genus that resulted in numerous polymorphies. All multistate characters were treated as unordered. A heuristic search using multiple TBR + TBR (multi*max*) with 10 000 maximum trees retained, 1000 replications and 10 starting trees per replication, produced two trees with a length of 1283, consistency index of 21 and retention index of 47. The strict consensus tree is given as Table 2. Ordering, weighting or deletion of selected characters significantly changed the topography of the resulting trees but was not further pursued.

In the consensus tree (Table 2), the entire tribe Olethreutini, apart from Sycacantha Diakonoff and Atriscripta, gen. nov., clusters on a single branch with both the Microcorsini and Bactrini included as subordinate and paraphyletic respectively polyphyletic taxa. The majority of both Eucosmini and Enarmoniini are each clustered as a monophyletic group. A basal grade below the Olethreutini contains the single grapholitine taxon (Grapholita zapyrana), two genera of the Enarmoniini (Loboschiza Diakonoff and Irianassa Diakonoff) and the only two isolated genera of the Olethreutini, Sycacantha and Atriscripta, gen. nov. A second grade between the Olethreutini and the terminal Eucosmini and Enarmoniini consists first of four enarmoniine genera (Anthozela Meyrick, Oriodryas Turner, Eucosmogastra Diakonoff and Pternidora Meyrick), followed by four eucosmine genera (Epiblema Hübner, Coenobiodes Kuznetzov, Eucosmophyes Diakonoff, Herpystis Meyrick). The eucosmine absconditana-group is the sister taxon to the enarmoniine clade. The Microcorsini appear as a paraphyletic group subordinate within the Olethreutini, with Demeijerella Diakonoff as the sister taxon to Collogenes Meyrick. The Bactrini also are subordinate within the Olethreutini, as well as polyphyletic with Lobesia Guenée between Endothenia Stephens and the sister taxa Bactra Stephens and Syntozyga Lower.

Table 2

The failure of the analysis to recover several well accepted groupings based on unique apomorphies casts some doubt on the adequacy of the matrix. It was scored for all characters encountered within each genus as a very effective basis for descriptions and keys and to facilitate future work on the Papuan and Oriental fauna. However, it possibly contained too much polymorphism for a phylogenetic analysis, and time constraints did not allow rescoring a more suitable matrix based on exemplar taxa. The resulting phylogenetic hypothesis therefore has to be regarded as very preliminary and insufficiently conclusive to make changes to the higher classification. For example, several probable synapomorphies linking Bactra and Endothenia with some Olethreutini and the presence of a tibial pencil in Endothenia suggest that the highly derived Bactrini are merely subordinate within the Olethreutini rather than a separate tribe. The cladistic analysis indeed places the Bactrini within the Olethreutini but is considered too flawed to provide sufficient corroboration to sink a generally accepted tribe. A more rigorous cladistic analysis, reassessing some of the characters and their states, and including molecular information, may resolve this question in the near future.

2. Morphology

Morphological structures are here discussed largely from a taxonomic angle, with emphasis on diagnostic characters including external features and the sclerotised parts of the genitalia. An in-depth treatment of lepidopteran morphology is provided by Kristensen (2003) and a summary of tortricid morphology by Horak (1991).

Head (Figs 1–9)

The head capsule of Olethreutinae, as of all Tortricidae, lacks prominent sutures, but there is usually a scale-free band between the antennal sockets, with the vertex above and the frontoclypeus (termed frons in the following) below. The tentorial pits (Fig. 6) are prominent on the denuded head. The tortricid head scaling is quite distinct, with anteriorly oriented scales on the vertex and upper frons, but upwardly oriented, small, appressed scales over the lower of the frons (Figs 1–3). According to Zimmerman (1978), upwardly directed scales on the lower part of the frontoclypeus occur among microlepidoptera only in the Tortricidae, many Cossidae and some Tineidae. The anteriorly oriented scales on the vertex and upper frons are usually erect and long, rarely short and closely appressed.

The compound eye is large and spherical, with a band of microtrichia along its margin. Only at very high magnification are short, sparse setae visible. The ocellus and chaetosema are usually well developed, adjacent to each other behind the antenna, next to the eye margin (Fig. 4). Ocellus size varies greatly, but at least a vestigial scar is always present. The chaetosema is a pin-cushionlike domed area with narrow, modified hair scales.

The antenna is filiform, reaching to about the middle of the forewing or to slightly beyond. The scape and pedicel are unmodified, densely scaled, with a small intercalary sclerite between the two segments. The flagellum always has scales across its dorsal surface and usually an unscaled band of variable width along its ventral surface bearing the majority of sensilla. More rarely, the flagellum is scaled all round. Each olethreutine flagellar segment has a single ring of scales (Fig. 8), usually ventrally interrupted, whereas all other Tortricidae, except for some Spargonothini not present in Australia, have two rings of scales per flagellar segment (Fig. 7). The sensory setae, termed cilia in taxonomic language, vary particularly in the male from nearly invisible on the scaled antenna to as long as the diameter of the flagellum. In some genera of the tribe Eucosmini and the tortricine tribe Archipini the flagellum in the male may be notched at or near its base, with a deep dorsal excavation (Fig. 9) involving several flagellar segments. This secondary sexual modification has not only been developed at least twice within the family, it also has been lost again many times within genera.

Olethreutine mouthparts (Figs 5, 6) conform to those of other higher Lepidoptera, with the maxillae joined to form a proboscis. The narrow, unscaled labrum has a lateral pair of pilifers. The basal portion of the maxilla, the cardo and stipes, bear the grooved galea and the maxillary palpus. The small olethreutine maxillary palpus (Fig. 5) is hardly visible on the fully scaled head and varies from four-segmented and scaled, to reduced to a minute, naked remnant. The two grooved galeae form a naked proboscis, entirely unscaled. The labial palpi are well developed, three-segmented, with an invaginated vom Rath’s organ subapically on the last segment (Fig. 6). Relatively long, horizontally held, sinuate labial palpi with the second segment sinuate and distally widened, and the relatively short terminal segment angled forward or downward (Figs 1, 2), is the most common configuration for olethreutines and the plesiomorphic condition for the family. More rarely, the labial palpi are upcurved, with a rather slender and curved second segment, and a short but porrect or apically rising third segment (Fig. 3). However, upcurved olethreutine labial palpi never have a long, drawn-out tip tapering to a point as in the horn-shaped gelechioid palpi.

Thorax and appendages (Figs 10–17)

The thorax scaling may be smoothly appressed above or the scales may be raised to form a single or bipartite transverse crest posteriorly. Presence or absence of this raised scale tuft, the posterior crest, is a taxonomically informative character, but it is also a very ephemeral structure easily lost without the damage being apparent.

Figs 1–9. Tortricid head structure. 1, Epiphyas postvittana (Tortricinae). 2, Crocidosema plebejana (Olethreutinae) (arrow: maxillary palpus). 3, Cydia pomonella (Olethreutinae). 4, ocellus (oc) and chaetosema (ch). 5, pilifer (pi) and maxillary palpus (mp). 6, descaled head with mouthparts, tentorial pits (tp) and vom Rath’s organ (vR). 7, tortricine antennal flagellum (E. postvittana). 8, olethreutine antennal flagellum (Dudua aprobola). 9, flagellum with antennal notch (arrow) (Strepsicrates semicanella).

The tegulae (Fig. 10), curved, crescent-shaped plates pertaining to the mesothorax and covering the bases of the forewings, have a scale tuft from their downcurved anterior corner (termed anteroventral corner in descriptions) that may be lengthened to form a long scale pencil, extending posteriorly beneath the wing bases.

In the Olethreutinae the legs are well developed and typical of ditrysian Lepidoptera, with the fore tibia with an epiphysis with bristles along its inner surface, the mid tibia with a pair of apical spurs, and the hind tibia with two pairs of spurs, one medial and one apical. The legs are densely scaled, with the scaling on the mid tibia modified with a spiny appearance, due to parallel, grooved, pointed scales of decreasing size packed into sheaves, each enveloped by the outermost, largest scale. Several olethreutine groups have a modified hind tibia in the male, often with a narrow, long, eversible scale pencil from the base of the inner surface (Figs 15, 16), and sometimes with large lateral scale fringes and/or various large clumps of modified scales along the inner surface (Fig. 17). The fivesegmented tarsus usually has spines near the apex of segments 1–4, reduced in some groups.

Olethreutine forewing shape (Figs 10–12) ranges from triangular to subrectangular to subovate, and from rather wide to quite narrow, with the hindwings roughly as broad as the forewings. The forewing index (the maximum width of forewing divided by its maximum length) quantifies the wing ratio. The termen varies from nearly at right angles to the costa to strongly oblique and may be more or less straight or variably sinuate with an indentation beneath the apex. In some enarmoniine genera the apex is strongly projecting and followed by a deep notch, producing a so-called ‘falcate’ apex (Figs 474, 475). The hindwing shape is sometimes sexually dimorphic, usually with an expanded anal area in the male, often folded or rolled along the anal margin (Fig. 13). The forewing shape is sexually dimorphic only in a few cases (Lobesia, Archilobesia), but the base of the costa is expanded in the males of some groups and folded over the wing as a so-called costal fold (Figs 10, 12), usually enclosing modified scales. Wing coupling is effected by frenular bristles from the base of the hindwing fitting into the retinaculum beneath the forewing. Females have three (Fig. 14) (rarely two) frenular bristles, males a single composite one (Fig. 13). The retinaculum is a membranous hook arising from the subcosta (Sc) or a spur of Sc in males and a row of erect scales behind the base of the cubitus (CuA) in most females. The forewing scales are usually smoothly appressed, but raised scale tufts occur in several groups and can be of taxonomic significance. The olethreutine hindwing nearly always has a cubital pecten (Fig. 14), a row of hair scales on or just behind the base of CuA, sometimes modified (Crocidosema plebejana Zeller, Fig. 76), in contrast to nearly all other tortricids that lack a cubital pecten. The hindwing scaling is variously modified in the male, particularly along the anal margin, but also including very small and widely spaced scales resulting in a semi-translucent wing (Fig. 667), fields of melanic scales above and/or beneath (Fig. 48), and scale pencils in diverse positions (Figs 13, 25–27, 39).

The olethreutine forewing pattern (Figs 10, 11) is derived from several basic elements, variously developed and modified. Several dark, parallel, outwardly oblique, transverse bands and derivations thereof constitute one of the basic pattern elements, present in all tortricid subfamilies but with various interpretations of their number (Brown and Powell 1991; Baixeras 2002; Razowski 2003b). Traditionally, these so-called transverse fasciae have been referred to by specific names, but as homology of a given pattern element with the relevant fascia is often unclear and there is no consensus nomenclature, no fascia names are used here. Costal strigulae (pairs of short transverse white marks along the costa) constitute the second basic pattern element, also found in all three tortricid subfamilies. Pairs of costal strigulae frame the five dark fasciae (Brown and Powell 1991; Baixeras 2002), but as some of the strigulae are usually obliterated, especially in the basal half, doubled or confluent, no attempt at numbering has been made here. Strictly speaking, the white markings are the strigulae, but on pale wings the dark spacers in between are often more conspicuous and for ease of descriptions have been referred to as dark strigulae. A complex, often metallic marking in the tornus consisting of several parallel, longitudinal, black dashes framed by an erect silvery band on each side if fully developed, is variably termed ocellus, speculum or ocelloid patch (Fig. 11), the term here used to differentiate it from the ocellus proper.

Olethreutine wing venation is typically heteroneurous with a reduced number of radius branches (R-branches) in the hindwing, but otherwise only little modified in the generalised condition. The vein nomenclature adopted here conforms to that of Common (1990), Nielsen and Common (1991) and the earlier volumes of the series ‘Monographs on Australian Lepidoptera’, with the radial sectors numbered as R2, R3, R4 and R5 rather than the phylogenetically correct Rs1, Rs2, Rs3 and Rs4.

Figs 10–13. Olethreutine adult males and wing venation (ap, apex; cf, costal fold; ch, chorda; cs, costal strigulae; cv, closing vein; el, expanded anal lobe; fb, frenulum bristle, male; Ms, M-stem; op, ocelloid patch; sp, scale pencil; tg, tegulae; to, tornus). 10, Spilonota sp. 11, Grapholita zapyrana. 12, forewing, Spilonota constrictana, male. 13, hindwing, Holocola zopherana, male, with single frenulum bristle, M3 and CuA1 fused into single vein and 3A absent.

Fig. 14. Olethreutine female hindwing with cubital pecten (cp) and three frenulum bristles (fb).

The underived forewing has a full complement of free veins, with the discal cell apically closed by the closing vein (discocellular vein) and with the media stem (M-stem) and the stem of R4 and R5 (chorda) well developed. If present, the chorda delineates a small accessory cell (Fig. 12), but M-stem and/or chorda may be reduced or absent. R4 usually runs to the costa and R5 to the termen, but rarely R5 ends at or before the apex. R4 and R5 are stalked in several groups, R3 is often approximated to R4 but only rarely are the two veins stalked or R3, R4 and R5 all borne on the one stalk. Males of Lobesia Guenée often have a pterostigma (Fig. 31) and several of the R branches may be modified in Metaselena. Presence of three free M branches and two free branches of the cubitus anterior (CuA branches) is the underived condition, with the relative position of M2 and M3 of taxonomic significance, varying from widely distant and parallel at base to having their bases closely approximated and M2 curved towards M3. In Aglaogonia, gen. nov. and Ancylophyes Diakonoff (both Enarmoniini) one forewing vein is missing. With very few exceptions the cubitus posterior (CuP) is present near the wing margin, but never as a fully developed vein. The anal veins 1A and 2A are fused distally, forming a basal loop.

The underived olethreutine hindwing also has a full complement of free veins, but the M-stem is never developed and the cell is closed only by vestigial cross-veins. The subcosta is fused with the first radius branch (Sc+R1), with R1 often visible as a cross vein between the bases of Sc and the single radial sector (Rs). The three M branches and the two CuA branches are free from the discal cell in the generalised olethreutine hindwing, with some of these veins becoming approximated, stalked or even fused in more derived wings. With few exceptions Rs and M1 are closely approximated and parallel at their base (Fig. 13) or the two veins are stalked, and the rare cases where these two veins are somewhat distant right from their bases could possibly denote the ancestral character state (Horak 1984). Whether M3 and CuA1 are stalked or not has long been regarded as the crucial difference between Eucosmini and Olethreutini, and despite several exceptions this rule still holds to a surprising extent, especially if one considers the least derived members of each genus. The cubitus posterior and all anal veins, the distally fused 1A+2A and 3A, are all present in the underived olethreutine hindwing, though often not as tubular veins. Presence of 3A in nearly all genera except the majority of Eucosmini seems to be taxonomically informative.

Figs 15–17. Olethreutine male hind tibia. 15, tibial pencil (tp) hidden in scaling. 16, exposed tibial pencil. 17, modified hind tibia of Cryptophlebia ombrodelta.

Pregenital abdomen

The olethreutine pregenital abdomen comprises eight segments in the male and seven segments in the female, though in more derived groups the ostium often is fused with or located within the accordingly modified seventh sternite. The sternum of the first segment (S1) is lacking and the configuration of S2 is of the tortricoid type with well-developed ventral apodemes (Figs 224, 465) and at most weak remnants of sternal rods (venulae). The anterolateral processes are usually well developed (Figs 465, 944), but absent in some genera (Fig. 977). A pair of round dorsal depressions, often called dorsal pits (Figs 312, 314), are present on T2 in several genera of the Olethreutini, and also occurs in at least two tortricine groups. The structure of the eighth segment in the male, especially the shape of its hind margin, is taxonomically relevant in some groups.

Olethreutine males are characterised by various secondary sexual modifications of the pregenital abdomen and its scale cover. These structures are often part of a functional unit involving the legs or scale pencils from the thorax or hindwing, and if they are present they may be taxonomically informative. Modified scaling on S2 is found in shallow folds laterally in Sycacantha Diakonoff and Lobesia (Fig. 222), and medially either as simple patches (Metaselena, Fig. 580) or complex tufts. Eversible deep pockets containing modified scales occur usually laterally on the abdomen between two segments (some Lobesia, Proschistis Meyrick, Pammenopsis Kuznetsov (Fig. 942)). Oxysemaphora males have longitudinal lateral folds with modified scales to hold the scale pencil from the hindwing anal margin. Males of some genera have lateral and/ or dorsal brushes of modified scales, often long and mane-like (Sorolopha Lower, Cryptophlebia Walsingham). The complex coremata of the eighth segment in the Grapholitini (Fig. 948) are discussed there.

Male genitalia (Figs 18–20)

Olethreutine male genitalia are unmistakable, their fused juxta, caulis and anellus complex immediately identifies their subfamily position and provides a convincing derived character (apomorphy) indicating that all Olethreutinae must have had a single common ancestor with exactly this structure. In other words, this unique modification strongly supports the monophyly of the Olethreutinae and, without exception, is a diagnostic feature for the entire subfamily. In the ground plan for the family, i.e. the condition hypothesized for an ancestral tortricid, the valvae and the teguminal appendages would all have been well developed and involved in clasping the female abdomen during copulation. The functional unit of the fused juxta, caulis and anellus complex, in conjunction with strongly sclerotised, rigid valvae, obviously allowed reduction of the teguminal appendages that have been reduced and lost repeatedly in many olethreutine groups. Whilst the male genitalia provide excellent diagnostic characters at the species and generic levels, parallel reduction of the teguminal appendages can result in superficially very similar structures erroneously suggesting a closer relationship.

The dorsal tegumen (tergum 9) and ventral vinculum (sternum 9) are laterally articulated to form a ring from which arise the various, mostly paired processes of the genitalia. Modification of the tegumen usually goes hand in hand with reduction of some of its appendages, in extreme cases resulting in a simple wide band. The two lateral, basalmost portions of the tegumen, the so-called pedunculi, frequently have medial apodemes, attachments for the tergal flexor muscle of the valva (m4 in Kuznetsov and Stekolnikov 1973, 1977) (Figs 219, 478, 480, 482). The vinculum always is a more or less narrow band in the Olethreutinae.

The full complement of teguminal appendages in the Olethreutinae, as in all lower Ditrysia, is a paired uncus, a pair of hairy socii and paired gnathos arms (Figs 19, 20). There is little doubt that the ancestral (plesiomorphic) olethreutine uncus was a paired structure that has become fused into a single process, variously modified and/or reduced repeatedly, even within genera. The membranous, hairy, plesiomorphic olethreutine socii lobes are greatly modified in many groups, with a particular derived character state often diagnostic: like the long, slender, arm-like socii of Sorolopha, the bifid socii of Sycacantha, or the rigid, strongly sclerotised socii of Spilonota and related genera. On the other hand, the socii have been reduced repeatedly to small setose patches or entirely lost in many Grapholitini. Relatively few olethreutines have well-developed, strongly sclerotised gnathos arms, either joined at the tip (Aterpia Guenée) or upcurved, and fused with the ventral wall of the anal tube (Fig. 20), forming a subscaphium. Often the gnathos is reduced, with only lateral remnants contiguous with the posterior tegumen margin.

The valvae, paired clasping organs, are strongly sclerotised in the Olethreutinae, with a large membranous portion at the base of their inner surface to allow muscle movement and somewhat misleadingly termed ‘basal excavation’. The valvae are articulated with the vinculum and the base of the pedunculi, and their dorsal margin (costa) proximally ends in a muscle attachment, the costal process (basal process (Komai 1999), costal hook). Numerous names have traditionally been used for various parts of the lepidopteran valva, with little consistency across the groups. These terms are here restricted to sacculus (the reinforced basal portion of the ventral margin often ending in a free process) and the cucullus (the distal part of the valva usually delineated by a ventral emargination). Additional lobes and processes of more derived valvae are simply described rather than referred to with terms such as pulvinus etc. that imply homology. Olethreutine valvae are often characterised by prominent spines, single or in groups, with an associated nomenclature. Attempts to establish the homology of the various spine groups across tribal boundaries were unsuccessful, and descriptive terms were chosen instead.

The diaphragm, the membrane closing the body cavity, extends between the tegumen and the vinculum and is perforated by the aedeagus and the anus. The aedeagus is supported from below by sclerotised portions of the diaphragm, the basal juxta fused with the caulis and the anellus that surrounds the aedeagus. In olethreutines, there are only rarely sclerotisations of the diaphragm above the aedeagus other than along the ventral surface of the anal tube, and a transtilla is never present. The olethreutine aedeagus is tube-shaped, and the membranous, eversible inner vesica often bears spines (cornuti) that may be shed in the bursa during copulation. A large group of deciduous cornuti, whose sockets are visible even if they have been shed, seems to be the ancestral condition. Progressive reduction of cornuti numbers and/or modification of at least some into fixed cornuti are both derived conditions.

Female genitalia (Figs 21, 22)

The female reproductive system of the Olethreutinae is of the ditrysian type with a separate copulatory opening (ostium bursae) in addition to the ovipore for oviposition. The ovipore and anus are close together between the typically tortricid, broad, flattened, and usually hairy ovipositor lobes that are a diagnostic feature for the family Tortricidae, and usually allow sexing of totricid moths in a reliable manner. Rarely, the ovipositor lobes are modified: either reduced in a telescopic ovipositor (Figs 492, 626) or with conspicuously enlarged papillae in Collogenes Meyrick (Fig. 144). The ovipositor lobes and the 8th tergite have paired long apodemes as muscle attachments (the apophyses posteriores and apophyses anteriores), further lengthened if the ovipositor is telescopic. The ostium bursae is located in the 8th sternite (S8) in generalised olethreutines (Fig. 21) but is displaced anteriorly in several groups, ending up either in a deep excavation of the hindmargin of S7 (Fig. 22), invaginated behind its hindmargin or located within S7. In its original position in the ventral membranous portion of the eighth segment, the entrance to the ostium is usually surrounded by some sort of sclerotisation, referred to as the ‘sterigma’. The posterior portion of the sterigma behind the ostium is also referred to as ‘lamella postvaginalis’ and that anterior to it as ‘lamella antevaginalis’ — practical but somewhat incorrect terms as the aperture concerned is strictly the copulatory orifice and not the vagina. In tortricids of the other two subfamilies, the sterigma is connected to the base of the apophyses anteriores by a well-sclerotised band, a connection absent in all olethreutines (Figs 21, 22) except the Microcorsini (Figs 132, 144). Absence of this connection is considered the derived condition, with the Microcorsini sharing the ancestral condition with the rest of the family. The bursa copulatrix is usually differentiated into a narrow ductus bursae and the corpus bursae proper. In generalised olethreutines the corpus bursae has two signa that are often reduced to one or entirely lost. The ductus seminalis connects the ductus bursae with the vagina and nearly always has a diverticulum (bulla seminalis).

The structure of the female genitalia, i.e. the bursa copulatrix and associated structures, often provides more pertinent information on higherlevel relationships than the male genitalia that are most informative at the generic and specific levels. Frequently, portions of the ductus bursae are sclerotised and may be taxonomically informative. A small and at least ventrally split ring just below the ostium, the colliculum (Fig. 21), seems part of the tortricid groundplan and is variously modified, lengthened or reduced in the olethreutines. Several enarmoniine genera are characterised by a sclerite in the bursa ‘neck’, the transition between the corpus bursae and the ductus bursae (Fig. 470). The structure of the signa, if they are present, is highly diagnostic, and Diakonoff (1973) very successfully subdivided the Olethreutini on the basis of signum shape. However, the signa are frequently reduced from two to one or completely lost, even within a genus. In some Eucosmini the bursa wall is strongly spinulose, except near the signa, and parallel with these spinules becoming larger, the signa seem to decrease in size and eventually to disappear (Figs 754–756). Deciduous cornuti retained in the corpus bursae may provide valuable information if all available males have shed their cornuti.

Figs 18–20. Olethreutine male genitalia. 18, Eucosmophyes (ae, aedeagus; an, anellus; be, basal excavation; ca, caulis; co, cornuti; cu, cucullus; ju, juxta; so, socii; vi, vinculum). 19, Cnecidophora; cp, costal process; ca, caulis; gn, gnathos; so, socii; un, uncus). 20, Fibuloides (an, anellus; cu, cucullus; gn, gnathos; pe, pedunculus; pun, paired uncus; sa, sacculus).

Figs 21–22. Olethreutine female genitalia. 21, Ophiorrhabda (bn, bursa ‘neck’; co, colliculum; cb, corpus bursae; db, ductus bursae; la, lamella antevaginalis; lp, lamella postvaginalis; si, signum). 22, Fulcrifera (aa, apophyses anteriores; ap, apophyses posteriores; bs, bulla seminalis; di, diverticulum; ds, ductus seminalis; lp, lamella postvaginalis; os, ostium).

Figs 23–24. Microcorsini, wing venation, males. 23, Cryptaspasma sordida, slide ANIC 2747. 24, Collogenes loricata, slide ANIC 13768.

Figs 25–27. Olethreutini, wing venation, males. 25, sOxysemaphora hacobiani, slide ANIC 1224. 26, Cnecidophora ochroptila, slide ANIC 1228. 27, Sorolopha cyclotoma, slide ANIC 1280.

Figs 28–33. Olethreutini, wing venation, males. 28, Sycacantha exedra, slide ANIC 1265. 29, Atriscripta arithmetica, slide ANIC 1267. 30, Demeijerella chrysoplea, slide ANIC 1266. 31, Lobesia physophora (pt, pterostigma), slide ANIC 1260. 32, Gnathmocerodes euplectra, slide ANIC 2717. 33, Rhectogonia electrosema, slide ANIC 1285.

Figs 34–39. Olethreutini, wing venation, males. 34, Ophiorrhabda phaeosigma, slide ANIC 2699. 35, Dudua phyllanthana, slide ANIC 1256. 36, Trachyschistis hians, slide ANIC 1244. 37, Archilobesia drymoptila, slide ANIC 1257. 38, Euobraztsovia chionodelta, slide ANIC 1280. 39, Eremas leucotrigona, slide ANIC 1269.

Figs 40–45. Olethreutini, wing venation, males. 40, Zomariana doxasticana, slide ANIC 17119. 41, Podognatha vinculata, slide ANIC 1281. 42, Temnolopha mosaica, slide ANIC 1259. 43, Diakonoffiana tricolorana, slide ANIC 1262. 44, Megalota uncimacula, slide ANIC 1264. 45, Costosa australis, slide ANIC 1270.

Figs 46–51. Olethreutini, wing venation, males. 46, Metrioglypha phyllodes, slide ANIC 1272. 47, Dactylioglypha tonica, slide ANIC 2467. 48, Statherotis sp., slide ANIC 1276. 49, Aterpia protosema, slide ANIC 17115. 50, Proschistis polyochtha, slide ANIC 2700. 5l, Rhodacra pyrrhocrossa, slide ANIC 17090.

Figs 52–55. Olethreutini and Bactrini, wing venation, males. 52, Gatesclarkeana sp. (Papua New Guinea), slide ANIC 2698. 53, Bactra venosana, slide ANIC 1252. 54, Syntozyga anconia, slide ANIC 1249. 55, Endothenia polymetalla, slide ANIC 3126.

Figs 56–61. Enarmoniini, wing venation, males. 56, Aglaogonia sp., slide ANIC 12960. 57, Anathamna sp., slide ANIC 3125. 58, Ancylis anguillana, slide ANIC 12961. 59, Ancylophyes monochroa, slide ANIC 1268. 60, Anthozela hemidoxa (Papua New Guinea), slide BM 24920. 61, Balbidomaga uptoni, slide ANIC 1274.

Figs 62–67. Enarmoniini, wing venation, males except 66. 62, Cyphophanes sp., slide ANIC 1240. 63, Eucosmogastra pyrrhopa, slide ANIC 2748. 64, Helictophanes uberana, slide ANIC 1273. 65, Helictophanes prospera, slide ANIC 1219. 66, Irianassa sp., female, slide ANIC 13810. 67, Loboschiza koenigiana, slide ANIC 1271.

Figs 68–73. Enarmoniini, wing venation, males except 71. 68, Metaselena lepta, slide GP 550. 69, Oriodryas olbophora, slide ANIC 1235. 70, Periphoeba trepida, slide ANIC 1295. 71, Pseudancylis acrogypsa, female, slide ANIC 17120. 72, Pternidora sp., slide ANIC 12931. 73, Tetramoera gracilistria, slide ANIC 1229.

Figs 74–75. Enarmoniini, wing venation, males. 74, Thysanocrepis crossota, slide ANIC 17116. 75, Toonavora aellaea, slide ANIC 1251.

Figs 76–81. Eucosmini, wing venation, males. 76, Crocidosema plebejana, slide ANIC 1209. 77, Epiblema strenuana, slide ANIC 13767. 78, Epinotia absconditana, slide ANIC 1263. 79, Coenobiodes melanocosma, slide ANIC 1247. 80, Melanodaedala scopulosana, slide ANIC 1286. 81, Rhopobota honesta, slide ANIC 1254.

Figs 82–86. Eucosmini, wing venation, males except additional female hindwing 83 and 86. 82, Fibuloides minuta, slide ANIC 17103. 83, Acroclita bryopa, slide ANIC 1257, slide ANIC 18806 (female hindwing). 84, Noduliferola neothela, slide ANIC 8664. 85, Heleanna chloreis, slide ANIC 17114. 86, Tritopterna capyra, slide ANIC 1279, slide ANIC 18807 (female hindwing).

Figs 87–93. Eucosmini, wing venation, males. 87, Spilonota constrictana, slide ANIC 1237. 88, Strepsicrates semicanella, slide ANIC 1218. 89, Holocola nr thalassinana, slide ANIC 1245. 90, Holocola deloschema, slide ANIC 1246. 91, Holocola zopherana, slide ANIC 1248. 92, Hermenias sp., slide ANIC 1231. 93, Eccoptocera australis, slide ANIC 1253.

Figs 94–97. Eucosmini, wing venation. 94, Herpystis sp., female, slide ANIC 13799. 95, Icelita indentata, male, slide ANIC 13785. 96, Eucosmophyes commoni, male, slide ANIC 13786. 97, Whittenella peltosema, female, slide ANIC 13798.

Figs 98–102. Grapholitini, wing venation, males except additional female hindwing 99 and 102. 98, Loranthacydia nr sinapichroa, slide ANIC 4942. 99, Notocydia nr niveimacula, slide ANIC 4920, slide ANIC 4921 (female hindwing). 100, Pammenemima tetramita, slide ANIC 8622. 101, Leguminivora sp., slide ANIC 19285. 102, Notocydia nr atripunctis, slide ANIC 4922, slide ANIC 4923 (female hindwing).

Figs 103–105. Grapholitini, wing venation, male fore- and hindwing, female hindwing. 103, Fulcrifera sp., slide ANIC 4929, slide ANIC 4933 (female hindwing). 104, Cydia pomonella, slide ANIC 4934, slide ANIC 4935 (female hindwing). 105, Apocydia pervicax, slide ANIC 4926, slide ANIC 4924 (female hindwing).

Figs 106–112. Grapholitini, wing venation, males. 106, Thaumatotibia zophophanes, slide ANIC 8605. 107, Thaumatotibia sp., hindwing only, slide ANIC 8607. 108, Cryptophlebia sp., slide ANIC 17026. 109, Gymnandrosoma gonomela, syntype [SAMA]. 110, Cryptophlebia ombrodelta, slide ANIC 8603. 111, Archiphlebia endophaga, slide ANIC 17010. 112, Archiphlebia rutilescens, slide ANIC 8608.

Figs 113–118. Grapholitini, wing venation, males except additional female hindwing of 117. 113, Acanthoclita sp. C, slide ANIC 8610. 114, Grapholita zapyrana, slide ANIC 8625. 115, Acanthoclita sp. B, slide ANIC 8612. 116, Microsarotis sp. (Thailand), slide F Komai 1120. 117, Pammenopsis barbata, slide ANIC 4925, slide ANIC 4936 (female hindwing). 118, Commoneria cyanosticha, slide ANIC 8635.

Figs 119–121. Grapholitini, wing venation. 119, Parapammene sp. A, male, slide ANIC 4941. 120, Parapammene sp. A, female, slide ANIC 4943. 121, Ixonympha sp., male fore- and hindwing, slide ANIC 4938; female hindwing, slide ANIC 4940.

Fig. 122. Posterior crest (pc), dorsally on thorax, Aterpia protosema.

3. Biology

Life histories

The Olethreutinae have by far their greatest diversity in the warmer and wetter parts of Australia, where rain or monsoon forests are their dominant habitats. Interpretation of label data in collections suggests that the great majority of species have several generations per year, especially in the moist tropics and subtropics. Species with a wide distribution and clearly several generations in the north are nevertheless rarely collected during winter in the temperate region, suggesting a slowing of their development during the cooler months. However, there may well be univoltine species in cooler and drier regions, especially among the Eucosmini in temperate southern latitudes or higher altitudes and possibly among the Grapholitini in seasonally dry habitats. No work has been done to establish whether any of the Australian Olethreutinae have a strict diapause to accomodate a seasonal food supply, such as availability of suitable fruit or seeds for some of the borers.

The flat tortricid ovipositor lobes are characteristic for the family and possibly an adaptation for a specific egg-laying mode. Nearly all tortricid eggs are round to oval, strongly flattened, dome-shaped to scale-like. In contrast to the Australian Tortricinae (Powell and Common 1985), there is very little information about olethreutine oviposition. Olethreutine eggs are usually deposited singly on the host plant, rarely in small, irregular clusters (Peterson 1965; van der Geest and Evenhuis 1991). In some species and genera with modified, telescopic ovipositors, the eggs are inserted into crevices in the host plant. None of the Australian species have ovipositors that would suggest a piercing mode of oviposition.

Feeding habits within the Olethreutinae are very diverse, ranging from leaf rolling and spinning foliage together to boring in shoots, fruit, seeds, stems, roots and woody branches. Some produce blister mines in leaves. The larvae of a few Australian species including Spilonota constrictana (Meyrick) live in a portable case made of hollowed flower buds or seed capsules spun together to form a tube (Fig. 730). Unlike some Tortricinae, none of the Australian Olethreutinae have been reared from dead plant tissue, i.e. leaf litter.

The ancestral tortricid larva is presumed to have been an external feeder, given the presence of an anal fork in all three tortricid subfamilies (Horak 1989). This structure serves to eject faeces from the feeding shelter and has frequently been lost in groups with internally feeding (i.e. boring) larvae. Had an internally feeding larva without the anal fork been ancestral, it would be very unlikely that later derived external feeders in all three subfamilies would have developed exactly the same type of anal fork. A considerable proportion of Australian olethreutine larvae are borers, often in seeds and fruit: one of the main reasons for their economic importance. However, both feeding modes, leaf rolling and boring, are found in several olethreutine tribes, and parallel shifts from one mode to the other have obviously occurred. It had been hoped that mapping internal and external feeding onto the cladogram resulting from this study would provide an indication of the shifts in feeding modes, but the cladistic analysis turned out to be too flawed to expect critical insights in this respect. However, the trends within tribes strongly suggest that: 1, external feeding is the more ancestral state within those tribes using both feeding modes, except possibly for the Grapholitini, and 2, switches between the two feeding modes have happened repeatedly, even within a tribe. In both the Microcorsini and Bactrini all larvae are borers, the former endosperm feeders in fruit and nuts and the latter predominantly in stems, often of monocotyledons. The known larvae of the Australian Olethreutini are nearly all leaf rollers or leaf tiers except for some species within Lobesia Guenée, Ophiorrhabda Diakonoff, Gnathmocerodes Diakonoff, and Gatesclarkeana Diakonoff, genera from four different genus-groups. The feeding mode is known for only a few enarmoniine genera, but it ranges from leaf rolling to fruit boring, and in Loboschiza koenigiana (Fabricius) both feeding modes occur in the same species. The larvae of Ancylis Hübner, probably the sister group to the remaining Enarmoniini, are usually leaf rollers, but one Australian species produces blotch mines in Banksia leaves. The Eucosmini also include external as well as internal feeders, with the more ancestral groups (Epinotia Hübner) predominantly leaf rolling, but with some borers even within Epinotia. The majority of Grapholitini larvae are internal feeders and, according to Komai’s (1999) hypothesis, this was the ancestral feeding mode of the Grapholita-group, with several subsequent reversals to external feeding. Almost all species of the remaining grapholitine genus-groups are borers as far as their biology is known. The Loranthacydia-group, the Dichrorampha-group (except for Pammenemima Diakonoff with leaf rolling larvae) and most species of the Cydia-group, in fact, all species with known biology in Australia, are borers. This poses the question as to whether internal feeding was the ancestral feeding mode for the entire tribe Grapholitini, though some species such as Grapholita molesta (Busck) have retained a well-developed anal fork.

Exceedingly little information is available on the biology of the pupal stage of Australian Olethreutinae. Most leaf rolling larvae pupate in their shelter, whereas boring larvae may pupate in the ground. Olethreutine pupae have rows of dorsal spines on the mobile abdominal segments and they partially extrude themselves from their cocoon before the adult emerges.

Adult olethreutines are mostly active early in the evening and at night, and they are successfully attracted to light. However, some Grapholitini fly during the day and may not readily come to light.

Larval host plants

For a worldwide study of larval food preferences of microlepidoptera, Powell (1980) analysed host plant usage of 540 tortricine and 841 olethreutine species. The data were mainly drawn from literature but included a list of food plants for the Australian fauna by I. F. B. Common, representing c. 950 Lepidoptera species. Even though the Holarctic region was heavily overrepresented in this sample, it is a useful departure point for looking at host plants of Australian Olethreutinae.

Powell (1980) found that 24% of tortricine and only 6% of olethreutine species were polyphagous, i.e. defined as feeding on three