Tineid Genera of Australia (Lepidoptera)

By GS Robinson and ES Nielsen

The introductory chapters of this book give a detailed review of the phylogeny, morphology, classification and biology of Tineidae on a worldwide scale. Detailed morphological treatment of each genus is complemented by illustrations of wing patterns, head structure and head vestiture, venation, and male and female genitalia of representative species.

• Language: English
• Publisher: CSIRO PUBLISHING
• Release date: Jan 1, 1993
• ISBN: 9780643105805

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Chapter 1

Phylogeny and Family Definition

Systematic Position of the Tineidae

The ditrysian Lepidoptera comprise some 30 superfamilies of which 26 form the Apoditrysia, a monophyletic group defined by the possession of a tortricid-type sternum II (Minet 1986; Nielsen 1989). The remaining ‘primitive’ ditrysian superfamilies are the Tineoidea, Gracillarioidea, Yponomeutoidea and Gelechioidea. All appear to be monophyletic (Robinson 1988a; Nielsen 1989). However, the precise relationship of these superfamilies is still uncertain. Bar the Tineoidea, all have the segmentation of the maxillary palpus reduced from a ground-plan state of five to only four segments. This reduction could be considered a synapomorphy of all Ditrysia excluding Tineoidea (Robinson 1988a).

Many of the constituent groups of the Yponomeutoidea and Gelechioidea, but not all, have distinct scattered spines on the abdominal terga. This feature may be a synapomorphy, albeit homoplasious, of the two groups or a feature lost again in the apoditrysian lineage. In the Tineoidea, Yponomeutoidea and Gracillarioidea the valval apodeme of the male is elongated, and muscle m4, the valval protractor, runs from the apodeme ventrally to insert on the vinculum close to the saccus. In the Gelechioidea and Apoditrysia, by contrast, the valval apodeme is strongly reduced. In the Gelechioidea, muscle m4 runs from the reduced apodeme or from its base either laterally or dorsally to the tegumen; this configuration is similar to that found in many but not all Apoditrysia (m4 runs to the vinculum in some Sesiidae and Zygaenidae, for example) (Kuznetsov and Stekolnikov 1976, 1977, 1984). The reduction of the apodeme and rotation of m4 may represent synapomorphies of Gelechioidea and Apoditrysia, but this relationship needs further investigation. A tentative phylogenetic hypothesis for the basal Ditrysian superfamilies is shown in Fig 5.

The monophyly of the Tineoidea is supported by five synapomorphies (Robinson 1988a); the corresponding plesiomorphic state is given in parentheses:

Fig 5 Tentative phylogeny of the basal superfamilies of Lepidoptera–Ditrysia. Possible autapomorphies supporting each clade are indicated.

1  frons with erect scales (as opposed to smooth-scaled).

2  labial palpus with lateral bristles (bristles absent).

3  proboscis short, with galeae disassociated, not coiled (proboscis elongate and coiled). Dugdale (1988) has noted also that the tineoid haustellum is not as strongly melanised as in other groups, and we can confirm this.

4  female genitalia with greatly elongated telescopic ovipositor (ovipositor short).

5  ovipositor with ‘ventral rods’ (‘rods’ absent).

It should be noted, however, that characters one and two occur in their apomorphic state in many Incurvarioidea, and may prove to be ground-plan characters of the Tineoidea if the Incurvarioidea are found to be the sister-group of the Ditrysia.

The superfamily Tineoidea contains four families: Tineidae, Eriocottidae, Acrolophidae and Psychidae. The Pseudarbelidae and Arrhenophanidae were considered by Robinson (1988a) to be subordinate groups within the Psychidae. Nielsen and Common (1991) included the Galacticidae within the Tineoidea; this placement requires confirmation, with clarification of the phylogenetic position of the group.

The monophyly of Acrolophidae + Psychidae is supported by three synapomorphies (Robinson 1988a):

1  antennal pecten absent (present).

2  proboscis rudimentary or absent (short, with galeae disassociated).

3  apophyses of metathoracic furca forming a bridge to the secondary furcal arms (furcal apophyses short, paramedial, not forming a bridge).

The monophyly of Eriocottidae + Acrolophidae + Psychidae is strongly supported by seven synapomorphies (Robinson 1988a):

1  antenna with strong tendency to bipectination (antenna filiform).

2  maxillary palpus reduced or absent (maxillary palpus five-segmented).

3  vein R5 terminating on forewing termen (terminating on costa).

4  retinaculum of male arising from wing membrane between costa and Sc (retinaculum arising from Sc).

5  female frenulum with supernumerary bristles (frenulum of only 2–4 bristles).

6  apex of sacculus in male genitalia with one or more thorn-like sensilla (sensilla absent).

7  in male genitalia, valva with basal pulvillus (a spinose membranous lobe on inner surface) (pulvillus absent).

The remaining group, the Tineidae, is the basalmost lineage of the Tineoidea, and thus of the ditrysian Lepidoptera. Robinson (1988a) noted that most Tineidae possess a single complete or almost complete annulus of scales on each flagellomere whereas the ditrysian ground-plan state appears to be the presence of two annuli that may be more or less disordered. The latter character-state is expressed in the Eriocottidae + Acrolophidae + Psychidae clade. However, several groups of Tineidae, including a disproportionate number of Australian genera, have two annuli of scales (e.g. Eudarcia Clemens) or a disordered pattern that appears to be derived from a pair of annuli (e.g. Timaea Walker). So the Tineidae lack a strong synapomorphy to support their monophyly, and they may therefore represent a grade rather than a clade. Among the Tineidae isolated instances of incongruence occur involving the characters that define the Tineoidea or the Eriocottidae + Acrolophidae + Psychidae clade. Examples include the absence of ventral ovipositor rods in Erechthias Meyrick (q.v.), the presence of a broad, triangular retinaculum arising from between the costa and Sc in Timaea and Mesopherna Meyrick, the smooth frons in Hieroxestinae, lack of lateral bristles on the labial palpi of some Opogona Zeller species, and a strongly developed and apparently functional proboscis in Timaea, Mesopherna and Metapherna. These apparent conflicts may represent examples of character convergence or reversal (homoplasy), but some might, on further investigation, contradict the monophyly of the Tineidae.

The phylogeny hypothesised for the Tineoidea is shown in Fig 6; the Tineidae are represented by a single lineage, but this is, as argued above, a debatable interpretation.

Subfamily Classification

Of 15 currently recognised subfamilies within the Tineidae, 11 occur in Australia. These are fully characterised below, with notes on composition, distribution and biology. The remaining four subfamilies are Siloscinae, Euplocaminae, Stathmopolitinae, and Teichobiinae. The subfamily Siloscinae contains three genera with 12 Afrotropical species (Gozmány and Vari 1973), although we have seen a further unnamed species from Borneo that may also be referable to this subfamily. Siloscines are characterised by erect spicular scales on the forewing and by the saccus of the male genitalia being articulated; the group is convincingly monophyletic. Euplocaminae contains the single genus Euplocamus Latreille, with 12 species occurring in the Palaearctic and Oriental regions. Stathmopolitinae contains the monobasic genus Stathmopolitis Walsingham from the Canary Island. Teichobiinae contains the single genus Psychoides Bruand, with three Palaearctic species.

We have not accounted for one family-group name in the Tineidae. The subfamily Amydriinae was proposed by Dietz (1905) for a ‘rag-bag’ of genera that included Setomorpha Zeller (see below). Amydria Clemens itself is currently associated with the Myrmecozelinae, but relationships here are so confusing (see Remarks for Perissomasticinae) that the correct placement or synonymy of the name Amydriinae must remain in doubt for the present.

The proliferation of family-group names within the Tineidae is helpful only insofar as some of them represent monophyletic entities, while the monophyly of the family Tineidae itself is in doubt. Most were erected in the context of regional faunistic studies, and their original definition antedates phylogenetic systematics. Some of the names have become widely used and widely accepted, particularly in the context of studies of Holarctic Tineidae. Zagulajev’s five monographs in the Fauna SSSR series, for example, deal with the Tineinae, Nemapogoninae, Scardiinae, Myrmecozelinae and Meessiinae respectively. Gozmány and Vari (1973) allocate the Afrotropical tineid genera to subfamilies. Petersen and Gaedike (passim) have been much more conservative, however, and have avoided the use of suprageneric categories in most of their papers with the exception of a few recent ones. Similarly, Davis (1983, 1984) has not used subfamily categories in checklists of the Nearctic and Neotropical Tineidae.

Fig 6 Phylogeny of the Tineoidea (s. str.). Autapomorphies supporting each clade are listed in the text (after Robinson 1988a).

In this work we have adopted suprageneric categories to accommodate the tineid genera occurring in Australia. In so doing we have looked for evidence of monophyly in each of the 15 recognised tineid subfamilies and, perhaps surprisingly, have found it in 13. The Myrmecozelinae, however, appear to be little more than a ‘rag-bag’ assemblage of taxa. We have retained it as a convenient collection-point for genera that cannot be accommodated elsewhere. However, some monophyletic groups of genera are discernible within the Myrmecozelinae, for example the tribe Haplotineini (Robinson 1984a). The monophyly of the Meessiinae is equally unconvincing, although lichenivory may be a synapomorphy of some of the presently-included genera (see below).

The erection of numerous tribal names within some tineid subfamilies has, with a few exceptions, been of only limited value. Most represent small, apparently monophyletic groups but many are monogeneric. We have listed such names in the synonymy of the recognised subfamilies. No such names are involved in the case of the four subfamilies that do not occur in Australia (see above).

Interrelationships of the monophyletic subfamilies of the Tineidae are obscure. It has not proved possible to identify any convincing synapomorphies of any combination of groups, and further work is required to elucidate their phylogeny. Earlier explanations of relationships, such as that of Zagulajev (1968), have a vague phenetic basis and are unconvincing narratives.

Chapter 2

Morphology

Head

The head vestiture of many Tineidae is often characteristic, consisting of erect piliform scales on the occiput, vertex and frons (Fig 171). The scales tend to be grouped into tufts, reflecting the distribution of the underlying scale-bases, and may form more or less conspicuous whorls. However, there are some exceptions to this pattern. For example, semi-appressed forward-directed scales are exhibited by Setomorpha (Fig 351); the frons of Hippiochaetes is smooth-scaled; scales of many Hieroxestinae are appressed and plate-like, and, on the occiput, directed posteriorly.

Tineidae possess neither externally visible ocelli nor chaetosemata. The cranium (head capsule) of most Tineidae is strongly marked with sutures or sulci (Figs 7, 8). These scale-free lines may be membranous, and in some taxa broad and clearly flexible, but in some genera the head capsule is strongly sclerotised and most sutures may be entirely fused and indiscernible. In Tiquadra, for example, only the occipital suture remains.

The transfrontal sulcus separates the vertex from the frons and is usually straight and broad, running between the antennal bases. In Oenoe, however, it is strongly reflexed dorsally, and in Dryadaula it appears ‘W’-shaped. In some taxa a U-shaped zone of flexible membrane in the upper half of the frons, bounded dorsally by the transfrontal sulcus, encloses an oval or circular sclerite. This facial sclerite may possibly be protruded to assist in eclosion. However, there is no direct evidence for this suggestion.

The epicranial suture runs longitudinally from the middle of the transfrontal sulcus to the posterior margin of the head. It may be only weakly developed posteriorly. The occipital suture is transverse and runs parallel with the posterior margin of the head, separating the occiput from the vertex. In a few taxa the occipital suture may be displaced caudally and merge with the posterior dorsal margin of the cranium. A suture originates at each end of the occipital suture and runs diagonally to the middle of the posterior cranial margin to form a common junction with the epicranial suture; we refer to these sutures as the ‘oblique scale-free lines’ of the occiput.

The disposition of scale-bases on the head capsule is dictated by the presence of sutures which separate the scale-bases into discrete fields. Within these fields the scales are evenly distributed in most taxa, but in Hieroxestinae the scale-bases are arranged in irregular transverse rows. Hieroxestines also have the scale-pattern modified to accommodate the ‘brow-ridge’ in Opogona (q.v.) and the frontal tuft in Amphixystis. There is considerable variation in the distribution of scale-bases on the frons. Typical or plesiomorphic may be the pattern seen in Edosa, a roughly circular mediofrontal field (corresponding to the ‘protrusible sclerite’ – above) separated by the U-shaped membranous suture from a pair of boot-shaped lateral fields (Fig 430). Modifications to this pattern include loss of the U-shaped suture and thus a complete frontal scale-field (Hapsiferinae, Setomorphinae, Hieroxestinae), reduction of the fields (Tenaga), loss of the ventral region of the ‘U’ (Timaea, Acridotarsa), and reduction of the ‘U’ and enlargement of the lateral fields (Mimoscopa, Mesophema, Metapherna, Erechthiinae, Harmaclona, Gerontha, etc.).

The compound eye of tineids is unmodified, the facets of regular size. Differentiated facets and a horizontally divided or partly divided eye are exhibited by some species of Roeslerstam-miidae that resemble tineids superficially. The eye may be emarginate postero-dorsally to accommodate the antennal base, a feature particularly marked in some Hieroxestinae. The size of the eye relative to that of the cranium varies greatly. Among Australian genera small eyes are found in, for example, Dryadaula and Amphixystis with an interocular index of about 0.7. Large eyes are found in, for example, Eudarcia males with an index of about 1.4. The smallest eyes that we have seen are in representatives of the Afrotropical-Asian genus Machaeropteris (index 0.50–0.55) and the largest in Australian Metapherna (index 2.2).

The mouthparts of many Tineidae are strongly developed, but reduction of one or more of the component structures occurs frequently, and apparently in many independent lineages. Extreme reduction may be observed in Harmaclona (Fig 570) and the Setomorphinae, in which only a short rudiment remains of the galea and maxillary palpus. In Tenaga even these are absent (Fig 133). But in all Tineidae that we have examined the labial palpus is fully developed. Faucheux and Chauvin (1979b) have reviewed the structure of the mouthparts of representatives of Monopis, Trichophaga, Tinea and Tineola, based on SEM studies.

The pilifers are strongly developed triangular processes in many genera, bearing stiff mesally-directed setae. Their reduction to a pad of setae arising from only a slight swelling on the labrum has occurred in Acridotarsa, Moerarchis, Thomintarra and in the Setomorphinae. Curiously, the setae are deciduous in Acridotarsa and the Setomorphinae. Reduction in the number of setae or their complete absence may be observed in, for example, all four Australian genera here placed in the Meessiinae. Pilifers are entirely absent in Trachycentra, Phereoeca and Harmaclona.

The mandible is strongly developed in many Tineidae, forming a well-sclerotised but nevertheless often inconspicuous process held transversely beneath the labrum. The shape of the mandible is fairly consistent among the genera examined, scimitar- or kukri-shaped, but in some groups it is less broadened apically and best described as banana-shaped or elongately rectangular. It may often be almost half the width of the frons, so that the two mandibles span almost the complete width of the head capsule. The mandible is particularly conspicuous in Dryadaula and some Hieroxestinae. It is apparently absent in some groups, for example Harmaclona and Tenaga.

Figs 7, 8 Tineidae, head terminology; dorsal aspect (7) and frontal aspect (8).

The maxillary palpus is more strongly developed in the Tineidae than in any other ditrysian group. Its reduction occurs frequently in separate lineages and such reduction is strongly correlated with reduction of the galea. Its complete loss is rare but may be seen in Tenaga. Fully developed, the maxillary palpus is five-segmented and in life is held folded. The ‘elbow’ of the fold, the third segment, has dark scales on its outer surface in many taxa. In a macerated head preparation the maxillary palpus frequently exceeds the labial palpus in length. In some species of Eudarcia the palpus is six-segmented, apparently through division of the third segment. The disposition of sensilla on the maxillary palpus is characteristic, although not always simple to discern by light microscopy.

Faucheux and Chauvin (1980b) have described an SEM study of the sensilla on the maxillary palpus of representatives of Monopis, Trichophaga, Tinea and Tineola. Their interpretation of the palpus as six-segmented is based on their counting the rudiment of the cardo plus stipes as the first segment. They found sensilla campaniformia on the cardo-stipes rudiment and on the first segment; there was also a single campaniform sensillum on the third segment of Tinea pellionella. They recorded sensilla chaetica (our trichoid sensilla) on the fourth and fifth segments but these were absent from the fourth segment in Trichophaga. They found trichoid sensilla on the terminal segment of the much-reduced maxillary palpus of Tineola, in which genus the homology of the segments is uncertain. They also recorded sensilla basiconica on the fifth segment situated on an apical and a subapical process.

Our examination of maxillary palpi was by light microscopy only; the first segment bears a small group of up to five or six sensilla trichodea on its inner surface; these are absent in some Scardiinae. In many groups the second segment bears sensilla that we identified originally as sensilla coeloconica but which Faucheux and Chauvin have shown to be sensilla campaniformia; these are not always obvious in light microscopy. The third segment appears to be scale-bearing only. The fourth and fifth segments are densely packed with sensilla trichodea interspersed with microtrichia, and the fifth segment usually bears conspicuous apical and subapical processes bearing sensilla basiconica, the latter not visible in conventional microscopy. All these groups of sensilla, presumed to represent the ground-plan distribution, are subject to reduction or loss in particular taxa, such reduction appearing fairly consistent within genera and frequently correlated with shortening of the palpus. Reduced segmentation of the maxillary palpus occurs in many genera: for example, Phereoeca is four-segmented as are Mesopherna and Metapherna. The palpus is three-segmented in Harmaclona, but is so small that some variation in this number might be expected. Reduction almost to a rudiment, with segmentation ill-defined (one to three segments) is to be seen in Lindera and Setomorpha. Further structural modification is unusual, but noteworthy is the spine-like development of the trichoid sensilla on the fourth segment in Asymplecta, and the enlarged terminal segment which carries a whorl of scales in Tinissa.

The galeae in most Tineidae are short, disassociated, not as strongly sclerotised as in other Lepidoptera families with a functional proboscis and, if rolled up, are rolled in a haphazard fashion. However, in Timaea, Mesopherna and Metapherna the galeae are more elongate than in other Tineidae and apparently form a functional proboscis (Figs 474–476). Reduction or loss of the galeae is common-place. They are lost completely in Tenaga, for example, and are reduced to short, stumpy rudiments in Tineola and Harmaclona. We have not studied the galeal sensilla in detail. However, most species have at least one trichoid sensillum on the outer galeal surface at about one-quarter its length. The sensilla styloconica on the outer surface of the distal half of the galea are usually shallow warts, but in Mimoscopa and Metapherna they are conspicuously protuberant. Faucheux and Chauvin (1979a) examined the galeal rudiment of Tineola and found that the apex bore a pair of sensilla chaetica (our trichoid sensilla) and four sensilla basiconica. The sensilla on the galea of representatives of four tineine genera have been described by Faucheux and Chauvin (1980b); they record 1–5 sensilla chaetica on the outer surface of the galea along with numerous regularly spaced sensilla basiconica. A small number of sensilla basiconica are also scattered along the food groove.

The labial palpus is invariably three-segmented in Tineidae, and is fairly consistent in terms of its length relative to the height of the head capsule, and in the relative length of the three segments. The position in which the labial palpus is held varies between groups. In some Hapsiferinae and Hieroxestinae it is distinctly recurved; it is drooping in some Meessiinae. A group of sensilla campaniformia (Faucheux and Chauvin 1979a, 1980a), is usually present on the inner surface of the basal segment. The second segment may be swollen (e.g. Trachycentra, Thomintarra) but in most genera is subcylindrical and curved dorsad. Coarse lateral bristles occur on the second segment in most tineid genera. They are usually arranged in an irregular line of up to 15 bristles (but in some genera there are far fewer – only one or two in Asymplecta or Phaeoses). Additionally there may be a terminal whorl of up to about 15 bristles from the upper outer surface of the segment. Bristles appear to be shed easily during life, and their number and distribution on the palpus can be accurately ascertained only by examination of the large sockets in a macerated preparation. Finer ventral bristles occur in some taxa; however, their socket diameter is considerably less than that of the coarse bristles and similar to that of normal scales. Coarse bristles appear to be absent in several genera – Metapherna, Harmaclona, Trachycentra, Tiquadra and Opogona. In addition to bristles, the labial palpus may have a ventral brush of scales (e.g. Trachycentra), but in some genera such as Opogona the palpus is smooth-scaled. The terminal segment is slightly to strongly flattened in many Tineidae and may additionally be greatly swollen (Parochmastis, Tiquadra). A more or less conspicuous vom Rath’s organ is present in many genera, consisting of sensilla coeloconica (Faucheux and Chauvin 1980a) lining a subapical groove or pit. The organ is particularly large in Acridotarsa. It is apparently absent in several genera (e.g. Morophaga). Faucheux and Chauvin (1980a) studied the distribution and number of sensilla of the vom Rath’s organ in Monopis, Tinea, Tineola and Trichophaga. They found that in Trichophaga, although no vom Rath’s organ is discernible, the sensilla coeloconica were still present but scattered over the surface of the segment. They found the vom Rath’s organ of Tineola to be minute, with a single sensillum. This does not accord with our observations of two individuals (by light microscopy) in which the vom Rath’s pit is large and apparently contains numerous sensilla.

The tineid antenna provides a further suite of characters. The scape is usually stout and irregularly vase-shaped, but is distinctly slender and elongated in Erechthiinae and Hieroxestinae. An antennal pecten is present in many genera, consisting of anything from two or three (e.g. Ectropoceros) to about thirty (e.g. Metapherna) stout bristles. A pecten is absent in Australian Dryadaulinae, Setomorphinae, Hapsiferinae, Harmaclona, Gerontha and Hieroxestinae. In the three species of Scardiinae represented in Australia the pecten is very dense, consisting of more than 40 bristles. The scape is smooth-scaled in most genera, but in Eudarcia, for example, elongate dorsal scales form a sparse eye-cap. The pedicel is always slightly larger than the basal flagellomeres, simple and barrel-shaped, with the exception of males of some Erechthias species in which it is modified to form part of the notch at the base of the flagellum. The basal flagellomeres are modified in males of Comodica and in some Erechthias species. In the latter they form a shallow notch and in the former the basalmost three flagellomeres are fused and subtend a dorsal process. This encloses a deep notch with internal spines.

The flagellomeres are cylindrical or approximately barrel-shaped in almost all genera. However, in groups in which the annulus of scales is incomplete ventrally the flagellomeres are irregular in cross-section (e.g. Morophaga). The flagellomeres are conspicuously ciliated to finely pubescent in all genera examined. However, in some groups the cilia are very short or subparallel to the surface of the flagellomere and hidden or almost completely hidden by the covering of scales. The ciliation of the basal flagellomere is very sparse, particularly on the dorsal surface, compared with subsequent segments. The cilia may be very long and whip-like (e.g. Gerontha). There is some sexual dimorphism in all genera examined, females having fewer and shorter cilia than males. Types of cilia cannot be adequately differentiated by light microscopy, and neither can other types of sensilla on the flagellomeres. However, SEM studies such as that of Faucheux (1989) on Trichophaga have shown that the ‘cilia’ consist of a mixture of sensilla trichodea, sensilla basiconica and sensilla chaetica. Additionally there are sensilla auricillica, sensilla styloconica, sensilla squamiformia, and sensilla coeloconica on the flagellomeres. Some of the latter are of the ‘palisaded’ type, distinguishable under high-power light microscopy. A single campaniform sensillum is present on the pedicel. The flagellomeres in most genera are completely covered by one or two annuli of overlapping scales. A notable exception is Morophaga in which the ventral surface of the antenna is scale-free (Fig 123). Most tineid genera have one annulus of scales per flagellomere (e.g. Tinea), but a disproportionate number of the Australian genera have two annuli (e.g. Eudarcia and Parochmastis) or a disrupted pattern of scale -cover that may be derived from the two-annuli condition (e.g. Mesophema). The width of the scales may vary considerably between genera, affecting the radial density of the scale-bases. In Trichophaga, Faucheux (1989) found that a typical flagellomere (the tenth) had a scale annulus consisting of 13 longitudinal rows of 5–6 scales. In genera with broad scales, such as Acridotarsa, the number of longitudinal rows is plainly fewer; in genera with very narrow scales, such as Edosa, the number is greater.

The length of the antenna varies both within and between genera. A markedly short antenna, 0.3x the length of the forewing, is found in Harmaclona, and a long antenna, 1.1x the length of the forewing, in Tenaga.

Thoracic Appendages

Variation in the structure of legs among the Tineidae is, for the most part, unremarkable. In most genera the legs are smooth-scaled, with the exception of the hind tibia which carries elongate suberect scales on its upper surface. The upper surfaces of the fore- and mid-tibia and tarsi are usually dark-scaled. The mid tibia of Scardiinae has oblique dark and light stripes on the outer surface. A foretibial epiphysis is present in most genera but may be absent as in, for example, Eudarcia. The tibial spur pattern is invariably 0–2–4. The scale vestiture of some Hieroxestinae is distinctly bristly, superficially resembling that of Stathmopodidae. Other groups may have bristly scales, particularly on the ventral surface of the tarsi (e.g. Tineola). Some genera have true spines or setae that survive maceration. We have not searched exhaustively for these, but note fine sparse setae in Tinissa, small tarsal spines in Monopis, strong tibial and tarsal spining in Timaea, and tarsal spining in Gerontha, Lindera, Tiquadra and Parochmastis. The hind legs of Gerontha are greatly elongated and the joints apparently possess a ‘spring-locking’ function such that they are held rigid even after maceration (Fig 514).

Fig 9 Tineidae, wing venation terminology.

The wings of Tineidae vary in shape. They may be very narrow as, for example, in Oenoe with forewing and hindwing indices of 0.17 and 0.09. Typically, they are broader than this, with a forewing index of 0.25–0.30 and a hindwing index of 0.20–0.25. Generally both wings are subovate although the shape of the anterior margin of the hindwing varies greatly and may be concave in some genera. The forewing apex may be produced and upturned (e.g. some Erechthiinae). The termen is usually convex but in Trachycentra is markedly concave giving the wing a falcate appearance. There is a wide variety of forewing patterns, from the uniform dirty brown of some Tinea to the striking metallic transverse strigulae of some Moerarchis and the complicated patterns of some Erechthias that make use of particoloured scales. In most genera the pattern elements are transverse, but in Mimoscopa and Timaea, for example, they are longitudinal. Marked sexual dimorphism is rare but is exhibited by Timaea in the case of pattern, by Gerontha in the case of shape, and by Setomorpha in the case of venation. The hindwing bears no pattern to speak of although it may be tinted with brown or grey, the pigmentation often concentrated apically and close to the veins. In some Scardiinae there are distinct dark speckles in the apical third of the hindwing. In some species of Gerontha the hindwing is semihyaline. There is considerable variation in the type of scale-cover of the forewing. Rough or raised scaling is frequently encountered; examples occur in Acridotarsa (rough scaling), Gerontha (rough scaling and patches of elongate erect scales) Oenoe and Monopis (tufts of raised scales) (Figs 16, 17). Marked variation in scale size and texture occurs in some Opogona species which have patches of broad, smooth metallic scales separating the dark and pale areas of the wing. Specialised sex-scales are unusual but we have noted such scaling in some Erechthiinae; several examples from this group are figured by Zimmerman (1978).

The wing scales in Tineidae are relatively long and slender. The wing scales are frequently multi-layered with the upper layer consisting of relatively long and slender scales with deeply dentate apical margins and the lower layer of shorter, broader and apically scalloped or rounded scales. There is some variation in relative length, overall shape and in the shape of the simpler apical margin which is usually dentate. The scales examined have all been normal-type scales (Nielsen and Common 1991) but a detailed study of scale ultrastructure is outside the scope of this work. Obvious scale dimorphism is observed in the tineine genera with a hyaline forewing spot (‘fovea’) (Figs 336, 337) and Edosa (Fig 416). The ultrastructure of the scales examined is very uniform; the majority of tineids have one row of simple, fine perforations, windows, in the inter-ridge area (Figs 12, 13). Exceptions are observed in Craniosara which have multiple, fine windows, Setomorpha with multiple, relatively large windows (Fig 14) and Tiquadra with a single row of relatively large windows (Fig 15). Nomenclature for wing scales is that proposed by Downey and Allyn (1975).

Figs 10, 11 Tineidae, retinaculum. 10: Tinea pellionella Linnaeus; typical tineid male retinaculum: elongately triangular and rolled. 11: Tinea bivittatella Walker; broad and shallow lobe.

Typically, Tineidae have a full complement of free forewing veins (Fig 9). Sc is invariably present, but is strongly displaced anteriorly in Dryadaula and Eudarcia. All branches of R terminate anterior to or at the apex; exceptions to this are some specimens of Acridotarsa in which R5 terminates just posterior to the apex, and Trachycentra and Mesopherna in which R5 terminates well posterior to the forewing apex. A pterostigma is developed consistently in relatively few genera – Trichophaga and Tineola - but occurs sporadically among species of Tinea and Monopis. R1 is absent in Crypsithyris and absent or vestigial in Opogona and Amphixystis. Absence of any other branch of R is rare, but occasional loss of R4 occurs, possibly through fusion with R5 (e.g. Xeringinia). Stalking of pairs of R branches occurs in many genera; the commonest pairing is R4 + 5, for example in Thomintarra, but R3 + 4 is frequent also, for example in Morophaga. R2 appears always to be free. More complicated stalking sequences, involving R3, R4, R5 and, in some cases, M1 may be seen in several genera, notably Tenaga and all Hieroxestinae excluding Asymplecta. Such stalking is subject to sexual dimorphism in Setomorpha (q.v.). Although all three M branches are usually present, one may be absent in such genera as Eudarcia and Analytarcha. Stalking of Ml and M2 occurs in, for example, Oenoe; M2 and M3 are stalked in Crypsithyris. M3 and CuA1 are stalked in Setomorpha and some Monopis species. Loss (or fusion) of either CuA1 or CuA2 occurs in Eudarcia, Oenoe and Parochmastis. CuP is present in all Tineidae but is tubular only proximally and distally. A1 and A2 typically form a broad basal loop but are fused for their distal two-thirds. However, A2 is weak in many genera and may be obsolete, for example in Dryadaula. Modifications to the forewing cross-veins occur in many genera, notably involving weakening of sections at the end of the cell. A novel M-Cu basal cross-vein has been developed in Moerarchis. More dramatic modifications to venation occur in those genera with a hyaline forewing spot – Monopis, Crypsithyris and Crypsithyrodes.

Figs 12–15 Tineidae, ultrastructure of normal type scales of forewing. 12: Moerarchis australasiella (Donovan), . 13: Ectropoceros pterocosma (Meyrick), . 14: Setomorpha rutella Zeller, . 15: Tiquadra atomarcha (Meyrick), . 12 and 13 show ‘normal’ small interridge windows, while 14 and 15 have different types of large windows.

As in the forewing, the hindwing venation of Tineidae is typically complete, with all veins free. Sc + R1 and Rs are separate and subparallel; Sc + R1 is displaced anteriorly in Eudarcia. Stalking of Ml and M2 is common and may be seen in, for example, Phereoeca and Tiquadra. M3 is lost in some genera, for example Dryadaula. The full complement of posteriormost veins – CuP, 1A + 2A and 3A – seems to be expressed only in large species with broad hindwings. 3A is absent in most groups with narrow hindwings.

Specialisations of the wing membrane are infrequent. Hyaline spots, possibly providing evaporative surfaces for pheromones, occur in some species of Erechthias, Comodica and Gerontha. The ‘hyaline’ spots of Monopis, Crypsithyris and Crypsithyrodes appear not to be of this type as they are covered with unpigmented thin scales (Fig 337).

There is considerable variation in the frenulo-retinacular wing-coupling of Tineidae. The female retinaculum consists of a few semi-erect scales on Cu. It does not appear to be a particularly effective device and is not strongly developed. The male retinaculum is typically an elongately triangular tongue with a rolled apex arising from Sc (Fig 10). But in Dryadaula and Eudarcia it is a very broad and shallow lobe from Sc. A superficially similar lobe, but arising from the membrane between the costa and Sc and resembling the retinaculum in Psychidae, occurs in Timaea (Fig 11) and Mesopherna. In Metapherna and Mimoscopa the retinaculum is of the triangular tongue type but arises from a short anterior spur from Sc. The retinaculum is absent in Trichophaga, Gerontha and Moerarchis. It is also lost in a number of Oriental species of Edosa. The male frenulum is a single bristle in all Tineidae examined. It is thickened, strongly melanised and spatulate in Trichophaga and the closely related (extra-Australian) genus Ceratophaga. The female frenulum typically consists of two or three bristles, but this number is reduced to one in Dryadaula, Phaeoses and some species of Erechthias. Four to six bristles are found in several genera, for example Acridotarsa, Thomintarra and Lindera. In the extra-Australian Scardia the female frenulum has about 15 bristles and resembles that of many Psychidae.

Figs 16, 17 Tineidae, scale tufts on forewings. 16: Oenoe sp., , scale tufts along dorsum. 17: Monopis meliorella (Walker), , tuft of vertical scales.

Supplementary wing-coupling devices, involving respectively microtrichia and specialised scaling along opposing wing surfaces, have been noted in Gerontha and Harmaclona (see below and Fig 496).

Pregenital Abdomen

The pregenital abdomen of Tineidae consists typically of eight segments in the male and seven in the female (Fig 18). The first sternum is lacking and the first tergum consists of a sclerotised frame enclosing a trapezoidal membrane. The latter may be partially sclerotised. T I is usually fused with T II, at least medially. There is no prespiracular tergosternal bar of sclerotisation between S II and T I and thus the abdominal base is of the ‘tineoid’ type (sensu Kyrki 1983). Usually slender and elongate apodemes arise from close to the anterior margin of S II. The supporting sclerotisation of these apodemes is variable; typically the medial anterior region of S II is heart-shaped or H-shaped and more strongly melanised than the remainder of the sternum. Sclerotisation from the apodeme base along the lateral margin of S II may be enhanced (venulae). The melanised anterior region of S II occupies about half its length; there is a group of sensory bristles just posterior to the demarcation of the modified and unmodified regions. The second sternum is highly modified and bears a pair of tympanal organs in Harmaclona. In some extra-Australian species of Dryadaula and in all Comodica species there is a pair of large hair-pencils on the second sternum.

The sterna and, particularly, the terga are generally wider in females than in males. Their sclerotisation may be uniform, but in many groups the sclerotisation and melanisation of the terga is greater medially and anteriorly so that they appear superficially T-shaped. T II is often more strongly melanised than subsequent terga. Their shape varies from rectangular to trapezoidal; posteriorly, the terga may become very narrow. Scaling on the abdomen is random in most groups, but the scales are arranged in irregular transverse rows in Oenoe and the Hieroxestinae. Other modifications include: the presence of a pair of lateral grape-like pleural glands between segments II and III in males of some Eudarcia species; a pair of hair-pencils in a glandular pouch in a similar position in males of some species of Erechthias; specialised scale-patches on S IV in Xeringinia; lateral slits that accommodate fringe-scales from the hindwing costa in both sexes of Niditinea; posteriorly directed socketed microtrichia on the sterna of Sarocrania; setose terga with medial wrinkles in Thomintarra; ‘honeycomb’- patterned patches in the anterior corners of the sterna in both sexes of Ectropoceros; shallow pouches in the intertergal membrane in Amphixystis.

Modifications of the hind segments of the abdomen are commonplace. Males of 13 of the Australian genera have shallow coremata in the pleural membrane of the eighth segment. The morphological affinity of these structures is uncertain, but in many cases they are connected by sclerotisation to the margins of the sternum and would appear to be derived by invagination of the sternal margin. Nevertheless, their overall homology is not proven and their structure and function needs further investigation. The presence or absence of such structures is variable within genera (Oenoe, Tinissa, Morophaga, Tiquadra, Erechthias), and has been shown to be highly homoplasious (Robinson 1986b). The eighth sternite in males of several genera is modified by the anterior extension of the corners, apparently to form an apodeme that may function in connection with extrusion of the coremata in the pleural membrane (e.g. Analytarcha and some Erechthias species). However, such an extension also occurs in Parochmastis which has no coremata.

Complex reduction and modification of the terminal two segments has occurred in males of Dryadaula, and the eighth segment is reduced to a narrow torque in males of Lindera and Setomorpha. The eighth tergite is strongly melanised and inverted T-shaped in some species of Niditinea, and the eighth sternite is divided longitudinally in some Hapsiferinae. Modification of the abdomen is less usual in females. The seventh segment is greatly reduced in most species of Tinissa to accommodate the corethrogyne, a dense tuft of extremely fine deciduous hair (Fig 23). A similar corethrogyne is invaginated into three intersegmental pouches in Perissomasticinae.

Fig 18 Tineidae, abdominal pelts. 18a: Edosa talantias (Meyrick), , slide ANIC 1147. 18b: Mimoscopa ochetaula Meyrick, , slide ANIC 1108. tp: tuberculate plate.

Male Genitalia

The male genitalia of Tineidae (Figs 19–21) exhibit a remarkable range of variety and provide, as in many other groups of Lepidoptera, a suite of