Abstract

INTRODUCTION

The Scopulini - the genus Scopula Schrank and its close relatives - is the largest tribe of the geometrid subfamily Sterrhinae, with about 900 described species. Species are found worldwide, although the group is mostly tropical.

The group is diverse in many aspects of morphology and ecology. The moths are typically moderate to small in size compared to other geometrids, with recurrent sexual dimorphism in wing size, shape and pattern. Males usually possess secondary sexual characters, such as coremata on the 2nd and 8th sternites and hair pencils on the hind legs. In the most extreme cases, male hindleg tarsi are absent; the swallowed tibia with a hair pencil is not used for walking but primarily for scent production and distribution (Hashimoto, 1992). Most species are nocturnal, straw-coloured and cryptic, yet there are a number of deviations, especially in the tropical lineages and in the groups that have adopted a diurnal mode of life. For example, species of the diurnal African genus Aletis Hübner are brightly coloured and are considered to be model species that are mimicked in appearance by species of the butterfly genus Euphaedra Hübner (Nymphalidae) (Staude & Curle, 1997).

The biology of most of the species is unknown. In the better-known western Palaearctic, Japanese, and Nearctic faunas, caterpillars of many species are polyphagous feeders on low herbs, although monophagy is not unusual (McGuffin, 1967; Sugi, 1987; Ebert, 2001). Some specialization in larval host-plants is evident at the generic level; for example, the species of Problepsis Lederer tend to feed on Oleaceae and those of Antitrygodes Warren on Rubiaceae (Holloway, 1997; Robinson et al., 2002). In South-east Asia a few species of Scopula have adopted an unusual way of life: the adults feed on mammalian sweat, tears and blood seeping from wounds (Bänziger & Fletcher, 1985), while species of Zythos Fletcher have been recorded visiting carrion (Holloway, 1997). A few species are of minor pest status, attacking tobacco (Sannino & Balbiani, 1984; Sannino & Espinosa, 1999) and groundnut (Satpathi, 1995). Typically, larvae are narrow and stick-like, and rest at an angle of 45 ° (Sugi, 1987; Ebert, 2001).

There has been only one published study on the systematics of the tribe, a treatment based on Palaearctic material (Sterneck, 1941). Other publications deal with generic or species descriptions, focusing mainly on regional faunas (Covell, 1970; Hausmann, 1994; Yazaki, 1996; Karisch, 2001). Lack of a global view of the tribe's taxonomy has resulted in a relatively restricted generic classification. This is problematic because it can result in an unstable system in a group where new species are constantly being described and ascribed to genera that have traditionally been recognized in each region. A further consequence of regional classification has been a tendency towards describing new genera on anad hoc basis, where new species are identified that do not fit the existing system. Many generic concepts appear to have originated in this way and their status needs to be revised. A similar problem affects the systematics of the Macariini (Scoble & Krüger, 2002).

There has hitherto been no attempt to synthesize and evaluate the findings of these regional studies, or to view the systematics of the tribe from a global perspective. A recent cladistic analysis by Sihvonen & Kaila (2004) attempted to redress this problem for the tribes of Sterrhinae, in the course of which Scopulini was recovered as a monophyletic entity. It was beyond the scope of the analysis, however, to resolve generic relationships within Scopulini.

This is the first published study of the generic relationships of the tribe on a worldwide scale. Its objectives are as follows: (1) to present a testable hypothesis of the tribe's phylogeny; (2) to find characters that can be used to delimit natural, synapomorphy-based groups, and (3) to classify the extant, described fauna. It is hoped that this approach will lead to a more stable system in a species-rich group where generic concepts have hitherto been vague.

Classification of the Scopulini: historical background

I will not discuss in detail here the characters that appear in the original generic descriptions. Most of the latter were made in the 19th century, at a time when methods and generic concepts differed from ours. They often appear to be superficial; in many cases, names were introduced in the form of simple lists.

As already mentioned, the literature dealing with the generic classification and systematics of the tribe is limited, with most of the data drawn from checklists and similar treatments. Careful examination of a few of these (Covell, 1983; Wiltshire, 1994; McQuillan & Edwards, 1996; Viidalepp, 1996; Leraut, 1997; Hacker, 1999) reveals that there appear to be several well-recognized, possibly monophyletic groups within Scopulini such as Scopula, Problepsis and Zythos, although we are left with the problem of inferring whether the order of genera in the checklists reflects their phylogenetic relationships. I now summarize the historical classification of Scopulini in chronological order. Only papers with a broader systematic importance are discussed.

Pierce (1914) arranged the British fauna into tribes based on a detailed examination of genitalic structures, a novel approach at that time. Acidaliinae (Scopulini) consisted of one genus, Acidalia(the author was not mentioned in this context, see Appendix under Scopula), and the species referred to it are nowadays considered to belong to Scopula. He considered Acidalia a natural genus due to the peculiar structures of the male and female genitalia. These include inter alia extreme fusion of the male genitalia, presence of coremata at the articulation of the sacculus with the tegumen, cerata and mappa on the male's eight sternite, and a spine-like signum on the female genitalia.

In his monograph on the Palaearctic Geometridae, Prout (1912-16) brought together 21 genera that he collectively called the Acidalia group (Scopulini). The group was characterized, among other features, by having one or two large areoles, in contrast to the Cosymbia group (Cosymbiini) where this structure was either absent or if present, single and very small. In the Acidalia-group male antenna were fasciculate and valve simple, whereas in the Cosymbia-group they were bipectinated and valve complex. The Acidalia-group was divided into sections according to the number of forewing areoles. The two-areole group contained, for example, Rhodostrophia Hübner (Rhodostrophiini), as well as genera that are nowadays not considered to be closely related to it, including Somatina Guenée (Scopulini).

In the context of these groups, he discussed generic relationships. For example, the one-areole genus Problepsis was considered to be a straight derivative of two-areole genus Somatina. These two genera were differentiated from Acidalia by their larger size, and most species by the absence of spurs on male hind tibia, these being replaced by a hair pencil.

Prout described four new Scopulini genera -Glossotrophia, Antilycauges, Holarctias and Oar- the latter three being monotypic. The descriptions were based on external morphological features; for example, the lack of medial spurs on the female hind leg in Glossotrophia was considered exceptional. Antilycauges was characterized by having numerous short, fasciculated sensilla in the male antenna, an elongated wing cell (areola), and palpi which were stout and rough scaled. Holarctias was differentiated from closely related taxa, namely section Pylarpe Herrich-Schäffer of genus Scopula, by having a hairy head and body, flat front, small eyes and moderately long palpi with attenuated terminal segments.

Oar was considered an abnormal genus within the Acidalia-group and possibly a derivate of Emmiltis Hübner (Sterrhini) (see Sihvonen & Kaila, 2004), due to a shortened proboscis. Furthermore, palpi were covered with moderate hair scales and male antenna were characterized by numerous long, fasciculated sensilla. The monotypic genus Stigma Alphéraky was considered the closest relative of Oar, but differentiated from the latter by having palpi covered with long, erect hair scales. Finally, Cinglis Guenée was associated with the Cosymbia-group (Cosymbiini) because of its bipectinated antenna and complex valve, although the systematic position was considered preliminary.

In a supplement to his monograph, Prout (1934-39) redefined the generic concepts of the Scopulicae (Scopulini), largely based on external characters, that he had adopted in his earlier work. This redefinition was based on utilization of genitalic structures, relying on Sterneck's revisions of Palaearctic Sterrhinae (Sterneck's manuscript was widely distributed prior to its publication in 1941 and cited by Prout, see below) and onPierce (1914).

Prout included seven genera in the Scopula-group (Scopulini) in the Palaearctic region, characterized by two socii and absence of uncus and gnathos. Further, the valves were described as ‘flowing out’ (ausgeflossene), the male 8th sternite as having mappa and cerata and the female hind leg as having two pairs of spurs except in Glossotrophia. At the generic level, no delimiting characters were given to Scopula, whereas Glossotrophia was considered a natural genus and easy to delimit on the basis of one pair of spurs in the female hind leg. It was noted, however, that the apical portion of the sacculus of the valva (fibula) is melanized in a number of species in Scopula, as well as in Glossotrophia, thus obscuring the generic boundaries. Holarctias, Oar and Stigma were characterized by absence of cerata, and as having varying degrees of valve differences relative to Scopula. Although the aberrant wing venation of Cinglis was noted, Prout concluded that this genus is appropriately combined with Scopulini as it shares the typical condition of the male 8th sternite.

Prout (1929-35) subsequently treated the African Scopulini. As with the Palaearctic material, the generic descriptions emphasized differences in wing venation, male antennae, and hindtibial spurs. The majority of the genera that he associated with Scopulini are still treated as such, except Discomiosis Prout and Tricentroscelis Prout, which are currently associated with Rhodostrophiini (Sihvonen & Kaila, 2004). Zygophyxia Prout, on the other hand, was seen as a relative of Sterrha(=Idaea[Sterrhini]), although it is nowadays considered to be within the concept of the Scopulini (Hacker, 1999; Sihvonen & Kaila, 2004). The monotypic African genera Isoplenia Warren and Isoplenodia Prout were considered to be close relatives of Epicosymbia Warren, based on the pectinated male antennae and two-areole condition of the forewing.

Prout noted the structural similarity between Scopula and Epicosymbia, especially in features of the male genitalia, although the well-developed uncus of Epicosymbia was interpreted as an unusual Scopulini feature. The generic status of Lissoblemma Warren was questioned, and Prout speculated that it would perhaps be better placed as a Section within Somatina, having in common with many Somatina species the bipectinated male antenna and absence of male hind leg spurs. Currently, Lissoblemma is associated with Rhodostrophiini (Sihvonen & Kaila, 2004).

Monotypic Leucoxena was considered a branch of Scopula but retained as a separate genus as the hindwing veins Sc + R₁ and Rs were stalked for a longer distance than in the typical Scopula. Prout (1929-35) was the first to discuss the Sterrhinae connection of brightly coloured, diurnal species of two-areole genus Aletis and one-areole Cartaletis Warren, on the basis of wing venation characters and genitalic structures, although he still treated them as Oenochrominae.

Prout's (1920–41) monograph of Indo-Australian fauna was the first comprehensive treatment of Scopulini published in the region. He again adopted the approach he had used with the Palaearctic material. Antitrygodes was considered both as an offshoot of Somatina and as closely related to Problepsis, as it had only one areole present in the forewing. This compact group of cryptically coloured moths differed further from Somatina by having a less serrated hindwing margin, although the male genitalia were found to be similar to Scopula.

When Turner (1908) described Autanepsia, he considered it to be closely related to Rhodostrophia because the only species of Autanepsia had two areoles. Prout (1920–41) speculated whether Autanepsia should be treated as a section of Australian genus Dithalama Meyrick or Somatina, because it shared with them all the diagnostic characters (apart from its two terminal spurs, absent in the two latter genera). Hairy species of Dasybela Turner from Tasmania were noted to have superficial similarity with species of Holarctias from the Northern Hemisphere. The compact, small, primarily Malaysian genus Nobilia Walker (Zythos Fletcher) was diagnosed as having moderately short labial palps, hind tibia of males shortened and without spurs, and two-areole forewings. Monotypic Lipomelia Warren, while smaller than Nobilia, was diagnosed as having the same features.

Janse (1933–35) did not discuss the generic relationships of Scopulini, but his work is significant due to the detailed descriptions and illustrations that he provided. Species of Problepsis, for example, were shown to have two fused, setose broad lobes, a unique feature that has been shown to support monophyly of the genus (Sihvonen & Kaila, 2004). The genera immediately preceding and following Scopula in his publication included those that are still associated with the tribe, except Discomiosis(Rhodostrophiini) (Sihvonen & Kaila, 2004).

Sterneck (1941) published the only detailed hypothesis on the relationships of the Scopulini, based on a detailed morphological examination of a large number of taxa from across the Palaearctic region. The study was not limited to Scopulini, but covered all Palaearctic Sterrhinae. He laid the basis for further studies, and the detailed structural descriptions and illustrations that he provided are still widely used (e.g. Hausmann, 1994). He recognized three main lineages within Scopulini. Holarctias and Oar were treated separately as they lacked cerata and mappa on the male 8th sternite. Cinglis was treated in isolation because it had the unusual conditions of fasciculate male antennae with ventrolateral sensilla arranged in multiple rows, and aberrant wing venation (i.e. R₅ being unstalked and rising directly from the areole). The remaining six genera were divided into two groups on the basis of number of forewing areoles, aedeagus shape, and cerata structures. The first group included Problepsis and Somatina and the second group Antilycauges, Scopula, Glossotrophia and Stigma.

When Hausmann (1994) described two new genera, namely Scopuloides and Pseudocinglis, he emphasized the quantitative differences of proboscis length and wing venation characters, in addition to number of hindtibial spurs. Scopuloides was considered a close relative of Glossotrophia, sharing with it two spurs in the female hind leg, although it was noted that species of Scopuloides also had several features in common with Scopula. Pseudocinglis, although externally similar to Glosstorophia, was considered to be morphologically intermediate between Cinglis and Antilycauges on the basis of genital and wing venation characters.

Holloway (1997) provided explicit descriptions of Sterrhinae tribes and genera in his treatment of the Bornean fauna, treating the species from that area within a broader taxonomic context. The Problepsini and Aletini were also considered to be within the concept of Scopulini. While he did not discuss generic relationships or apomorphic features, the order in which he presented the genera can be inferred to reflect affinities between them. Differences in the structures of the male's 2nd and 8th sternite, socii and signum were often used in generic delimitations.

In addition to these works, a few taxonomic treatments of single genera, focusing on regional faunas, have been published. These include revisions of Nearctic species of Scopula(Covell, 1970), various studies on Glossotrophia(Hausmann, 1993a,,b, 1994) and Sundanean species of Zythos(Yazaki, 1996).

DEFINITION OF THE STUDY GROUP

The definition of the tribe Scopulini, as used here, follows the cladistic analysis of Sterrhinae by Sihvonen & Kaila (2004). In that paper we described how only a few of the putative characters delimiting the tribe effectively defined parts of it. It was found to include the Aletini and Problepsini, as had already been suggested by Holloway (1996, 1997). The generic relationships within the tribe were largely unresolved. Monophyly was found to be supported by two homoplastic synapomorphies: the absence of a medial ridge on the epinotum of the male metathorax (two occurrences within Sterrhinae) and the ovoid and granulate signum with spines pointing away from the centre (two occurrences and three reversals within Sterrhinae). The Sterrhini were recovered as the sister group of the Scopulini.

Currently, the tribe comprises 25 genera, 12 of which are monotypic (Sihvonen & Kaila, 2004).

MATERIAL AND METHODS

STUDY MATERIAL

This was obtained from the following institutions: Australian National Insect Collection, CSIRO Division of Entomology, Canberra, Australia (ANIC); The Natural History Museum, London, UK (BMNH); Finnish Museum of Natural History (Zoological Museum), Helsinki, Finland (ZMH), and Zoologische Staats-Sammlung, München, Germany (ZSBS).

SPECIMEN PREPARATION AND PHOTOGRAPHY

The genitalia and abdomen were prepared following methods described by Hardwick (1950). The male vesica was everted via the caecum that was cut open by placing the aedeagus inside a hypodermic syringe (Sihvonen, 2001).

Before preparing the head and thorax, the wings were removed and prepared (as described below), or glued to a piece of plastic with a temporary pin with original labels. The head and thorax, still attached to the original pin, were placed in a 10% potassium hydroxide (KOH) solution and kept at room temperature for 12–15 h. Following maceration, the original pin was removed and replaced by the temporary one. Scales were removed with a brush, except the male secondary sexual characters such as scent pencils of the hind tibia. The metathorax was separated from the mesothorax at the intersegmental membrane using forceps.

All preparations of genital, abdominal and other structures were preserved temporarily in glycerol, which allowed examination from various perspectives. These were subsequently transferred into ethanol tubes for 24 h to remove the glycerol and then mounted on slides in Euparal (Hardwick, 1950). Glass props were used to support the coverslip. Thoracic preparations were mounted laterally, and metathoracic preparations were mounted with the posterior margin facing the coverslip. All were stained with Chlorazol black.

The wing venation slides were prepared as follows. One pair of wings was removed either by gentle downwards pressure or by lifting with fine forceps. The wings were then placed in a dish containing 99.5% ethanol and scales removed from both surfaces with delicate brushes. They were placed on a slide in a drop of ethanol, which was replaced by a drop of euparal, and covered with a coverslip. The slides were left unstained.

Various structures were prepared using a Wild M5 stereomicroscope (maximum magnification 50×). External characters of adult and immature stages were examined using a Wild M10 (512×). Adult and genital structural characters were examined using a stereomicroscope and a Leitz Diaplan phase contrast compound microscope (1560×). Photographs were taken using a Leica DC300 digital camera and Image Manager software. Image files were processed and the final plates arranged using CorelDraw.

OUTGROUPS

Idaea aversata L. and I. emarginata L., from the Sterrhini, the putative sister group (Sihvonen & Kaila, 2004), were used in order to root the tree and polarize the characters (Farris, 1972; Nixon & Carpenter, 1993).

INGROUP

The study material included 90 species, covering the worldwide geographical range of the Scopulini (listed in Table 1). Their identity was checked against external or genital examination of type specimens, apart from a few species found in the Holarctic region, whose identity is well established.

In a phylogenetic analysis of a species-rich group, such as the Scopulini, it is impractical to include all described species. As a consequence, those included in the analysis were chosen according to the following guidelines. First, all type species of the nominal genera were included. If material was not available, I attempted to include species that were morphologically similar. Second, more than one species was included from each genus (unless monotypic) in order to reduce the number of characters that could possibly appear as autapomorphic.

Third, an attempt was made to cover the tribe's morphological diversity as exhaustively as possible. For this purpose 650 species were studied, including genitalia, covering over 70% of the described fauna (see Appendix). In addition, many more were examined externally. Finally, I attempted to include taxa that came from areas where the Scopulini have received comparatively little attention, in particular the Neotropics and the rest of the Southern Hemisphere. The type species of each included genus are illustrated as follows: imagos are shown in Figures 1–25, male genitalia in Figures 30–54, male 8th sternite in Figures 55–79 and female genitalia in Figures 80–99.

Figure 1–25.

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Type species of genera included in the phylogenetic analysis. The numbers on the figures refer to the numbers of the character states. 1. Idaea aversata. 2. Somatina anthophilata. 3. Dithalama cosmospila. 4. Isoplenodia arrogans. 5. Zythos turbata. 6. Lipomelia subusta. 7. Problepsis ocellata. 8. Scopula ornata. 9. Ignobilia urnaria. 10. Antitrygodes divisaria. 11. Epicosymbia dentisignata. 12. Glossotrophia confinaria. 13. Isoplenia trisinuata. 14. Cartaletis libyssa. 15. Aletis helcita. 16. Leucoxena lactea. 17. Stigma kuldschaensis. 18. Dasybela achroa. 19. Pseudocinglis eurata. 20. Cinglis humifusaria. 21. Autanepsia poliodesma. 22. Antilycauges pinguis. 23. Oar pratana. 24. Zygophyxia tornisecta. 25. Scopuloides fucata. Scale bars = 5.0 mm.

Figure 1–25.

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Type species of genera included in the phylogenetic analysis. The numbers on the figures refer to the numbers of the character states. 1. Idaea aversata. 2. Somatina anthophilata. 3. Dithalama cosmospila. 4. Isoplenodia arrogans. 5. Zythos turbata. 6. Lipomelia subusta. 7. Problepsis ocellata. 8. Scopula ornata. 9. Ignobilia urnaria. 10. Antitrygodes divisaria. 11. Epicosymbia dentisignata. 12. Glossotrophia confinaria. 13. Isoplenia trisinuata. 14. Cartaletis libyssa. 15. Aletis helcita. 16. Leucoxena lactea. 17. Stigma kuldschaensis. 18. Dasybela achroa. 19. Pseudocinglis eurata. 20. Cinglis humifusaria. 21. Autanepsia poliodesma. 22. Antilycauges pinguis. 23. Oar pratana. 24. Zygophyxia tornisecta. 25. Scopuloides fucata. Scale bars = 5.0 mm.

Figure 30–54.

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Male genitalia and aedeagus of type species of genera included in the phylogenetic analysis. The numbers in the figures refer to the numbers of the character states. 30. Idaea aversata(slide number) PS704. 31. Somatina anthophilata(slide number) BMNH GEO20369. 32. Dithalama cosmospila PS707. 33. Isoplenodia arrogans BMNH GEO20527. 34. Zythos turbata BMNH GEO20371. 35. Lipomelia subusta BMNH GEO20530. 36. Problepsis ocellata BMNH GEO20370. 37. Scopula ornata PS692. 38. Ignobilia urnaria BMNH GEO20531. 39. Antitrygodes divisaria BMNH GEO20375. 40. Epicosymbia dentisignata BMNH GEO20365. 41. Glossotrophia confinaria BMNH GEO20382. 42. Isoplenia trisinuata BMNH GEO20363. 43. Cartaletis libyssa BMNH GEO20372. 44. Aletis helcita BMNH GEO20360. 45. Leucoxena lactea BMNH GEO20376. 46. Stigma kuldschaensis PS859. 47. Dasybela achroa PS897. 48. Pseudocinglis eurata PS894. 49. Cinglis humifusaria PS648. 50. Autanepsia poliodesma BMNH GEO20526. 51. Antilycauges pinguis BMNH GEO20381. 52. Oar pratana PS714. 53. Zygophyxia tornisecta BMNH GEO20380. 54. Scopuloides origalis PS58 (male of S. fucata type species of the genus was not available for study). Figures are not to scale.

Figure 30–54.

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Male genitalia and aedeagus of type species of genera included in the phylogenetic analysis. The numbers in the figures refer to the numbers of the character states. 30. Idaea aversata(slide number) PS704. 31. Somatina anthophilata(slide number) BMNH GEO20369. 32. Dithalama cosmospila PS707. 33. Isoplenodia arrogans BMNH GEO20527. 34. Zythos turbata BMNH GEO20371. 35. Lipomelia subusta BMNH GEO20530. 36. Problepsis ocellata BMNH GEO20370. 37. Scopula ornata PS692. 38. Ignobilia urnaria BMNH GEO20531. 39. Antitrygodes divisaria BMNH GEO20375. 40. Epicosymbia dentisignata BMNH GEO20365. 41. Glossotrophia confinaria BMNH GEO20382. 42. Isoplenia trisinuata BMNH GEO20363. 43. Cartaletis libyssa BMNH GEO20372. 44. Aletis helcita BMNH GEO20360. 45. Leucoxena lactea BMNH GEO20376. 46. Stigma kuldschaensis PS859. 47. Dasybela achroa PS897. 48. Pseudocinglis eurata PS894. 49. Cinglis humifusaria PS648. 50. Autanepsia poliodesma BMNH GEO20526. 51. Antilycauges pinguis BMNH GEO20381. 52. Oar pratana PS714. 53. Zygophyxia tornisecta BMNH GEO20380. 54. Scopuloides origalis PS58 (male of S. fucata type species of the genus was not available for study). Figures are not to scale.

Figure 55–79.

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Male 8th sternite of type species of genera included in the phylogenetic analysis. The numbers in the figures refer to the numbers of the character states. 55. Idaea aversata(slide number) PS704. 56. Somatina anthophilata(slide number) BMNH GEO20369. 57. Dithalama cosmospila PS707. 58. Isoplenodia arrogans BMNH GEO20527. 59. Zythos turbata BMNH GEO20371. 60. Lipomelia subusta BMNH GEO20530. 61. Problepsis ocellata BMNH GEO20370. 62. Scopula ornata PS692. 63. Ignobilia urnaria BMNH GEO20531. 64. Antitrygodes divisaria BMNH GEO20375. 65. Epicosymbia dentisignata BMNH GEO20365. 66. Glossotrophia confinaria BMNH GEO20382. 67. Isoplenia trisinuata BMNH GEO20363. 68. Cartaletis libyssa BMNH GEO20372. 69. Aletis helcita BMNH GEO20360. 70. Leucoxena lactea BMNH GEO20376. 71. Stigma kuldschaensis PS859. 72. Dasybela achroa PS897. 73. Pseudocinglis eurata PS894. 74. Cinglis humifusaria PS648. 75. Autanepsia poliodesma BMNH GEO20526. 76. Antilycauges pinguis BMNH GEO20381. 77. Oar pratana PS714. 78. Zygophyxia tornisecta BMNH GEO20380. 79. Scopuloides origalis PS58 (the male of S. fucata. the type species of the genus. was not available for study). Figures are not to scale.

Figure 55–79.

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Male 8th sternite of type species of genera included in the phylogenetic analysis. The numbers in the figures refer to the numbers of the character states. 55. Idaea aversata(slide number) PS704. 56. Somatina anthophilata(slide number) BMNH GEO20369. 57. Dithalama cosmospila PS707. 58. Isoplenodia arrogans BMNH GEO20527. 59. Zythos turbata BMNH GEO20371. 60. Lipomelia subusta BMNH GEO20530. 61. Problepsis ocellata BMNH GEO20370. 62. Scopula ornata PS692. 63. Ignobilia urnaria BMNH GEO20531. 64. Antitrygodes divisaria BMNH GEO20375. 65. Epicosymbia dentisignata BMNH GEO20365. 66. Glossotrophia confinaria BMNH GEO20382. 67. Isoplenia trisinuata BMNH GEO20363. 68. Cartaletis libyssa BMNH GEO20372. 69. Aletis helcita BMNH GEO20360. 70. Leucoxena lactea BMNH GEO20376. 71. Stigma kuldschaensis PS859. 72. Dasybela achroa PS897. 73. Pseudocinglis eurata PS894. 74. Cinglis humifusaria PS648. 75. Autanepsia poliodesma BMNH GEO20526. 76. Antilycauges pinguis BMNH GEO20381. 77. Oar pratana PS714. 78. Zygophyxia tornisecta BMNH GEO20380. 79. Scopuloides origalis PS58 (the male of S. fucata. the type species of the genus. was not available for study). Figures are not to scale.

Figure 80–99.

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Female genitalia of type species of genera included in the phylogenetic analysis. The numbers in the figures refer to the numbers of the character states. 80. Idaea aversata(slide number) PS828. 81. Somatina anthophilata(slide number) BMNH GEO20472. 82. Dithalama cosmospila PS832. 83. Zythos turbata BMNH GEO20483. 84. Problepsis ocellata BMNH GEO20485. 85. Scopula ornata PS835. 86. Ignobilia urnaria(insert: signum) BMNH GEO20532. 87. Antitrygodes divisaria BMNH GEO20480. 88. Epicosymbia dentisignata BMNH GEO20462. 89. Glossotrophia confinaria BMNH GEO20476. 90. Isoplenia trisinuata BMNH GEO20464. 91. Cartaletis libyssa BMNH GEO20468. 92. Aletis helcita BMNH GEO20469. 93. Leucoxena lactea BMNH GEO20478. 94. Pseudocinglis eurata PS895. 95. Cinglis humifusaria PS644. 96. Antilycauges pinguis BMNH GEO20479 (insert: corpus bursae spines). 97. Oar pratana PS804. 98. Zygophyxia tornisecta BMNH GEO20474. 99. Scopuloides fucata PS59. Figures are not to scale.

Figure 80–99.

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Female genitalia of type species of genera included in the phylogenetic analysis. The numbers in the figures refer to the numbers of the character states. 80. Idaea aversata(slide number) PS828. 81. Somatina anthophilata(slide number) BMNH GEO20472. 82. Dithalama cosmospila PS832. 83. Zythos turbata BMNH GEO20483. 84. Problepsis ocellata BMNH GEO20485. 85. Scopula ornata PS835. 86. Ignobilia urnaria(insert: signum) BMNH GEO20532. 87. Antitrygodes divisaria BMNH GEO20480. 88. Epicosymbia dentisignata BMNH GEO20462. 89. Glossotrophia confinaria BMNH GEO20476. 90. Isoplenia trisinuata BMNH GEO20464. 91. Cartaletis libyssa BMNH GEO20468. 92. Aletis helcita BMNH GEO20469. 93. Leucoxena lactea BMNH GEO20478. 94. Pseudocinglis eurata PS895. 95. Cinglis humifusaria PS644. 96. Antilycauges pinguis BMNH GEO20479 (insert: corpus bursae spines). 97. Oar pratana PS804. 98. Zygophyxia tornisecta BMNH GEO20474. 99. Scopuloides fucata PS59. Figures are not to scale.

PHYLOGENETIC METHODS

The character set summarized in Table 2 was analysed using NONA v. 2.0 (Goloboff, 1999) and all characters were weighted equally. The command sequence used for the matrix, using TBR branch swapping, was hold*; hold/50; mult*200. The initial results were then submitted to more exhaustive searches as recommended in the manual: max*; swap*; altswap*; mswap*2. The trees were saved with sv*, read back into NONA and the command best issued to delete suboptimal cladograms (Nixon & Carpenter, 1996). The final cladograms were saved with sv*, and used for calculation of a strict consensus tree. Final trees, a strict consensus tree and character optimizations were studied using WinClada v.1.00.08 (Nixon, 2002).

Table 2.

Data matrix for the cladistic analysis of Scopulini, based on characters from adult morphology and ecology. Missing data are represented by ‘?’. Characters coded from the literature are within grey tint boxes. Numbers in parentheses after each taxon name indicate the literature reference as follows: (1) Hausmann (1994); (2) Wiltshire (1986); (3) Janse (1933–35); (4) Turner (1908); (5) Prout (1929–35); (6) Prout (1920–41)

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Table 2.

Data matrix for the cladistic analysis of Scopulini, based on characters from adult morphology and ecology. Missing data are represented by ‘?’. Characters coded from the literature are within grey tint boxes. Numbers in parentheses after each taxon name indicate the literature reference as follows: (1) Hausmann (1994); (2) Wiltshire (1986); (3) Janse (1933–35); (4) Turner (1908); (5) Prout (1929–35); (6) Prout (1920–41)

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CHARACTER DESCRIPTIONS

Where possible, the terminology of the general morphology, including wing venation and thoracic sclerites, was adopted from Scoble (1992), genital terminology from Covell (1970), Klots (1970) and Sibatini (1972). In doubtful cases, descriptive terms were used and were accompanied by illustrations.

There are 141 characters in total: 41 are from wings, 50 from the head, thorax, and abdomen, 49 from genitalia, and one from ecology. One hundred and four characters were binary and the 37 multistate characters were treated as unordered. I attempted to minimize a priori assumptions regarding the value of characters as a source of systematic information; consequently, all characters that could be coded unequivocally were included. Many, such as the shape of the base of the male metafurca (Ch. 54, Figs 109, 110), are novel and demonstrate that detailed morphological studies can yield new and phylogenetically significant characters.

Figure 109–117.

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Characters used in the cladistic analysis. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 109. Base of male metafurca (posterior view). Idaea aversata(slide number) PS766. 110. Base of male metafurca (posterior view). Cartaletis variabilis(slide number) BMNH GEO20575. 111. Membranous sac at posterior part of male metathoracic coxa (dorsal view). Scopula silonaria BMNH GEO20433. 112. Opening on dorsal sclerite of male metathorax (dorsal view). Scopula ternata PS718. 113. Hair pencils of tibia on outer and inner margin of male hindleg. Ignobilia urnaria BMNH GEO20579. 114. Pretarsus of male hindleg. Zythos turbata BMNH GEO20443. 115. Pretarsus of male hindleg. Problepsis achlyobathra BMNH GEO20724. 116. Pouch on male 2nd sternite with lateral invaginations. Antitrygodes parvimacula PS883. 117. Lateral appendices of socii. Problepsis plagiata PS709.

Figure 109–117.

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Characters used in the cladistic analysis. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 109. Base of male metafurca (posterior view). Idaea aversata(slide number) PS766. 110. Base of male metafurca (posterior view). Cartaletis variabilis(slide number) BMNH GEO20575. 111. Membranous sac at posterior part of male metathoracic coxa (dorsal view). Scopula silonaria BMNH GEO20433. 112. Opening on dorsal sclerite of male metathorax (dorsal view). Scopula ternata PS718. 113. Hair pencils of tibia on outer and inner margin of male hindleg. Ignobilia urnaria BMNH GEO20579. 114. Pretarsus of male hindleg. Zythos turbata BMNH GEO20443. 115. Pretarsus of male hindleg. Problepsis achlyobathra BMNH GEO20724. 116. Pouch on male 2nd sternite with lateral invaginations. Antitrygodes parvimacula PS883. 117. Lateral appendices of socii. Problepsis plagiata PS709.

Two autapomorphic qualitative characters (Chs. 0 and 58) were included in the analysis. This was because these characters were present in one monotypic genus only, and I wanted to alert lepidopterists to search for these features outside the taxa studied. Furthermore, the inclusion of autapomorphic characters does not affect the calculation of consistency indices (Yeates, 1992). The data set is given in Table 2.

Head

• 0. Pilifer: (0) present; (1) absent.




• 1. Sensilla styloconica at distal end of proboscis: (0) present; (1) absent (morphologically unidentifiable).




• 2. Proximal pigment layer of compound eyes: (0) circular; (1) semicircular (figured in Sihvonen & Kaila, 2004).




• 3. Two pores on anterior part of dorsal sclerite between antennae: (0) absent (morphologically unidentifiable); (1) present.




• 4. Two extensions on anterior part of dorsal sclerite between antennae: (0) absent; (1) present (Fig. 100). Similar protuberant, rounded projections dorsad to antennal opening have been reported from Ennominae (Young & McQuillan, 2003).




• 5. Scales between antennae: (0) flat scales only; (1) both flat and hair scales.




• 6. Scales among chaetosemata: (0) absent (Fig. 103); (1) present (Fig. 104).




• 7. Male antennal segments dorsally: (0) unicolorous; (1) bicolorous.




• 8. Ventrolateral sensilla on proximal part of male flagellomere: (0) single row; (1) multiple rows.




• 9. Ventrolateral sensilla on medial part of female flagellomere: (0) present; (1) absent.




• 10. Ventrolateral sensilla on proximal part of female flagellomere: (0) undifferentiated; (1) multiple rows; (2) single row.

Figure 100–108.

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Characters used in the cladistic analysis. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 100. Extensions on the anterior part of dorsal sclerite between antennae. Somatina microphylla(slide number) PS711. 101. Structure of iridescent scales. Zythos turbata. 102. Structure of iridescent scales. Problepsis conjunctiva. 103. Chaetosemata. Zythos avellanea. 104. Scales among chaetosemata. Cartaletis variabilis. 105. Dark patch at forewing transverse median line that is separated by vein CuA1 (veins drawn on top). Problepsis conjunctiva. 106. Dark patch at forewing M2 (veins drawn on top). Problepsis centrophora. 107. Anterior and posterior ventral lamina of metathorax metafurca. Scopula imitaria PS779. 108. Pouch on posterior margin of male metathorax coxa. Problepsis ocellata(slide number) BMNH GEO20421.

Figure 100–108.

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Characters used in the cladistic analysis. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 100. Extensions on the anterior part of dorsal sclerite between antennae. Somatina microphylla(slide number) PS711. 101. Structure of iridescent scales. Zythos turbata. 102. Structure of iridescent scales. Problepsis conjunctiva. 103. Chaetosemata. Zythos avellanea. 104. Scales among chaetosemata. Cartaletis variabilis. 105. Dark patch at forewing transverse median line that is separated by vein CuA1 (veins drawn on top). Problepsis conjunctiva. 106. Dark patch at forewing M2 (veins drawn on top). Problepsis centrophora. 107. Anterior and posterior ventral lamina of metathorax metafurca. Scopula imitaria PS779. 108. Pouch on posterior margin of male metathorax coxa. Problepsis ocellata(slide number) BMNH GEO20421.

Wing pattern

See Figures 26–29 for a few examples of how wing pattern homologies have been interpreted.

Figure 26–29.

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Interpretation of wing pattern homologies. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 26. Glossotrophia confinaria. 27. Problepsis aegretta. 28. Somatina chalyboeta. 29. Antitrygodes divisaria. Abbreviations: t.a., transverse anterior line. t.m., transverse median line. t.p.l., transverse posterior line. s.t.s.l., subterminal shade line. a.t.s.l., adterminal shade line. t.l., terminal line. Figures are not to scale.

Figure 26–29.

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Interpretation of wing pattern homologies. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 26. Glossotrophia confinaria. 27. Problepsis aegretta. 28. Somatina chalyboeta. 29. Antitrygodes divisaria. Abbreviations: t.a., transverse anterior line. t.m., transverse median line. t.p.l., transverse posterior line. s.t.s.l., subterminal shade line. a.t.s.l., adterminal shade line. t.l., terminal line. Figures are not to scale.

• 11. Forewing costa: (0) narrow; (1) wide, dark brown; (2) wide, grey.




• 12. Light green colour on wings: (0) absent; (1) present.




• 13. Reticulate pattern: (0) absent; (1) present (Fig. 10).




• 14. Elongated marking on hindwing inner margin: (0) absent; (1) present (Fig. 10).




• 15. Wings unicolorous (without irroration): (0) no; (1) yes, orange; (2) yes, light yellow.




• 16. Distribution of iridescent scales: (0) absent; (1) scattered uniformly; (2) restricted to a few rows including discal spots; (3) restricted to a few rows excluding discal spots.




• 17. Structure of iridescent scales: (0) absent; (1) glossy, longitudinally grooved (Fig. 101); (2) glossy, smooth (Fig. 102).




• 18. Iridescent scales of cilia: (0) absent; (1) present.




• 19. Forewing discal spot unicolorous, dark: (0) yes (Fig. 23); (1) no; (2) absent.




• 20. Forewing discal spot structure: (0) spot; (1) cryptic (Fig. 29); (2) lunular (Fig. 5); (3) ocellate, semiround; (4) ocellate, round (Fig. 7); (5) absent.




• 21. Forewing discal spot with wide brown margin: (0) absent; (1) present (Fig. 105).




• 22. Hindwing discal spots: (0) present, dark spot; (1) present, ocellate; (2) present, concolorous with wings; (3) absent.




• 23. Pale markings on anterior side of hindwing discal spot: (0) absent; (1) present (Fig. 5).




• 24. Dark patch on hindwing margin at vein M₂: (0) absent; (1) present (Fig. 29).




• 25. Transverse median line: (0) present, distinct (Fig. 13); (1) present, obscure (Fig. 7); (2) absent.




• 26. Dark patch at forewing transverse median line separated by vein CuA₁: (0) absent; (1) present (Fig. 105).




• 27. One dark patch at forewing M₂: (0) absent; (1) present (Fig. 106).




• 28. Transverse posterior line: (0) continuous; (1) spotted.




• 29. Two dark marks on forewings medial of transverse posterior line: (0) absent; (1) present (Fig. 8).




• 30. Shade on forewing termen between transverse anterior and transverse median lines: (0) absent; (1) present (Fig. 28).




• 31. Shade on forewing between transverse median and transverse posterior lines: (0) absent; (1) present (Fig. 23).




• 32. Dark mark at forewing termen: (0) absent; (1) present (Fig. 8).




• 33. Forewing margins wide: (0) no; (1) yes (Fig. 15).




• 34. Forewing margins with white spots: (0) no; (1) yes (Fig. 14).




• 35. Hindwing margins wide: (0) no; (1) yes.




• 36. Hindwing margins with white spots: (0) no; (1) yes.




• 37. Terminal line at forewing apex: (0) present, not smoothly curved around apex; (1) present, smoothly curved around apex (Fig. 19); (2) absent. A smoothly curving terminal line that continues around the forewing apex has been considered a synapomorphy of Glossotrophia(Hausmann, 1993c).




• 38. Terminal line continuity at forewing margin: (0) discontinuous; (1) continuous; (2) absent.




• 39. Terminal line width at forewing margin: (0) equally wide; (1) wider at vein ends; (2) absent.




• 40. Spot at vein end at forewing margin: (0) absent; (1) present.




• 41. Subterminal shade line: (0) present, continuous; (1) present, discontinuous (Fig. 7); (2) absent.




• 42. Subterminal shade line and terminal line: (0) do not touch each other on fore and hindwings; (1) touch each other at fore and hindwings (Fig. 6).




• 43. Adterminal shade line: (0) present (Figs 26–29); (1) absent.




• 44. Fringe: (0) chequered, dark spot at vein end; (1) unicolorous; (2) chequered, white spot at vein end (Fig. 20).

Wing scales

• 45. Shape at forewing discal spot area: (0) margins parallel; (1) tapering towards apex.




• 46. Surface at forewing discal spot area: (0) longitudinally grooved; (1) smooth.

Wing shape

• 47. Forewing termen rounded:(0) no; (1) yes.

Wing venation

• 48. Forewing areoles: (0) one, formed by R veins; (1) two, formed by R veins. Features of wing venation have been used extensively in the systematics of the Scopulini (e.g. Prout, 1912-16; Forbes, 1948; Hausmann, 1994).




• 49. Point of origin of forewing vein R₅: (0) proximal areole, stalked with veins R₂-R₄; (1) distal areole, distinctly separate from R₂-R₄; (2) distal areole, stalked with R₂-R₄; (3) proximal areole, distinctly separate from R₂-R₄; (4) stalked with M₁. I am aware that characters 49–51 are variable and the division of these into distinct character states may turn out to be artificial if more taxa and material were to be studied.




• 50. Point of origin of forewing vein M₁: (0) proximal areole; (1) distal areole.




• 51. Hindwing veins Sc+ R₁and Rs: (0) fused shortly; (1) separate, but connected via a crossvein (R₁?). 51: 1 was earlier considered to be a synapomorphy for the Orthostixinae (Hausmann, 2001).

Thorax

• 52. Anterior ventral lamina of metathorax metafurca: (0) present (Fig. 107); (1) absent.




• 53. Posterior ventral lamina of metathorax metafurca: (0) present (Fig. 107); (1) absent.




• 54. Base of metathorax metafurca: (0) margins constricted (Fig. 109); (1) lateral margins basally opened or straight (Fig. 110).




• 55. Pouch on posterior margin of male metathorax coxa: (0) present (Fig. 108); (1) absent. Pouch absent in females.




• 56. Membrane sac at posterior part of male metathorax coxa: (0) absent; (1) present (Fig. 111).




• 57. Lateral processes of male metathorax coxa: (0) elongated (Fig. 109); (1) undifferentiated (Fig. 110). Processes absent in females.




• 58. Opening on male metathorax dorsal sclerite: (0) present (Fig. 112); (1) absent.




• 59. Medial ridge in epinotum: (0) present; (1) absent.




• 60. Male hindleg outer margin hair pencil of tibia: (0) present (Fig. 113); (1) absent.




• 61. Male hindleg inner margin hair pencil of tibia: (0) present (Fig. 113); (1) absent.




• 62. Male hindleg distal spurs of tibia: (0) absent; (1) one; (2) two. The number of male and female hindleg spurs has been used extensively in systematics of the Scopulini (e.g. Janse, 1933-35). Intraspecific variation has been reported to occur in the number of spurs in a few Glossotrophia species (Hausmann, 1993c).




• 63. Female hindleg medial spurs of tibia: (0) absent; (1) two; (2) one.




• 64. Apical spines on male hindleg tarsus: (0) absent; (1) present.




• 65. Thick spines on anterior side of male hindleg tarsus: (0) absent; (1) present.




• 66. Male hindleg tarsomeres: (0) not fused; (1) tarsomeres 1–5 fused; (2) tarsomeres 2–5 fused; (3) tarsomeres 4–5 fused.




• 67. Male hindleg claws: (0) absent (Fig. 115); (1) one (Fig. 114); (2) two, separate.




• 68. Male hindleg pretarsus arolium: (0) absent; (1) present.




• 69. Male hindleg pretarsus pulvillus: (0) present (Figs 114, 115); (1) absent.

Abdomen

• 70. Three parallel rows of white spots: (0) no; (1) yes (Fig. 15).




• 71. Pouch on male 2nd sternite anteromedially: (0) round (Fig. 116); (1) posterior margin elongated; (2) absent. The surface of this pouch was found to be variable between species. It is smooth in the majority of taxa. It is not unusual for it to be round, with a granulate surface (e.g. Scopula eburneata) or have star-shaped invaginations (e.g. Scopula achrosta). In a few species, such as Aletis helcita and Stigma kuldschaensis, it is covered with stout setae. Due to the quantitative nature of this variation I was unable to code it unambiguously.




• 72. Pouch on male 2nd sternite anteromedially: (0) with one chamber (Fig. 116); (1) absent; (2) with two chambers.




• 73. Pouch on male 2nd sternite anteromedially with lateral elongation: (0) laterally not invaginated; (1) laterally invaginated (Fig. 116); (2) absent.




• 74. Extensions on male 2nd sternite anterolaterally: (0) present (Fig. 116); (1) absent.




• 75. Setae laterally on 6th male segment: (0) absent; (1) present.




• 76. Pouch between male 7th and 8th sternite: (0) absent; (1) two pouches (Fig. 59); (2) one pouch (Fig. 66).




• 77. Male 8th sternite anterior margin: (0) unmodified; (1) trifurcate, blunt (Fig. 61); (2) trifurcate, round (Fig. 64); (3) bifurcate (Fig. 74); (4) elongated medially (Fig. 78).




• 78. Male 8th sternite with two parallel grooves: (0) absent; (1) present (Fig. 64).




• 79. Cerata: (0) absent; (1) rudimentary (Fig. 72); (2) fully developed (Fig. 62). There exists intraspecific variation in the shape and length of the cerata in many species of Scopula and Glossotrophia(Hausmann, 1999b). For this reason I did not include quantitative features of these structures in this study.




• 80. Cerata fused with mappa: (0) no; (1) yes (Fig. 61).




• 81. Cerata with setae: (0) no; (1) yes (Fig. 78).




• 82. Cerata bifurcate: (0) no; (1) yes.




• 83. Mappa: (0) absent; (1) setose (Fig. 75); (2) bare (Fig. 76).




• 84. Posterior margin of mappa separate: (0) no; (1) yes (Fig. 65).




• 85. Posterior part of male 8th sternite sclerotized: (0) no; (1) yes (Fig. 59).




• 86. Posterolateral appendices on male 8th sternite: (0) present (Fig. 56); (1) absent.




• 87. Male 8th tergite: (0) unmodified, membranous; (1) modified.




• 88. Male 8th tergite posterior margin: (0) unmodified; (1) concave, with two round lateral lobes (Fig. 118); (2) concave, with two sclerotized lateral spines (Fig. 119); (3) concave, with two sharp lateral projections (Fig. 120); (4) convex (Fig. 121).




• 89. Male 8th tergite laterally constricted: (0) no; (1) yes (Fig. 122).




• 90. Male 8th tergite with two membranous appendices on posterolateral margin: (0) no; (1) yes (Fig. 121).

Figure 118–132.

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Characters used in the cladistic analysis. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 118. Posterior margin of male 8th tergite. Scopula submutata(slide number) PS693. 119. Posterior margin of male 8th tergite. Dithalama cosmospila PS707. 120. Posterior margin of male 8th tergite. Somatina eurymitra PS888. 121. Posterior margin of male 8th tergite. Zythos avellanea PS879. 122. Posterior margin of male 8th tergite. Antitrygodes divisaria(slide number) BMNH GEO20375. 123. Apex of coremata. Scopula rhodocraspeda BMNH GEO20700. 124. Position of sacculus. Zythos obliterata BMNH GEO20591. 125. Dorsally fused anterior margins of tegumen (dorsal view). Zythos turbata BMNH GEO20371. 126. Plate-shaped sclerotization at proximal end of ductus ejaculatorius. Antilycauges pinguis BMNH GEO20381. 127. Sclerotization of vesica. Scopula rivularia BMNH GEO20684. 128. Dorsal diverticulum of vesica with sharp sclerotization at apex. Problepsis plagiata PS709. 129. Parallel folds of vesica. Zythos turbata BMNH GEO20371. 130. Signum. Somatina indicataria PS834. 131. Bare zone of signum. Zythos obliterata BMNH GEO20592. 132. Flexible lamella antevaginalis divided into two parts. Zythos avellanea PS880.

Figure 118–132.

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Characters used in the cladistic analysis. Species are mentioned in their generic combinations prior to analysis. The numbers in the figures refer to the numbers of the character states. 118. Posterior margin of male 8th tergite. Scopula submutata(slide number) PS693. 119. Posterior margin of male 8th tergite. Dithalama cosmospila PS707. 120. Posterior margin of male 8th tergite. Somatina eurymitra PS888. 121. Posterior margin of male 8th tergite. Zythos avellanea PS879. 122. Posterior margin of male 8th tergite. Antitrygodes divisaria(slide number) BMNH GEO20375. 123. Apex of coremata. Scopula rhodocraspeda BMNH GEO20700. 124. Position of sacculus. Zythos obliterata BMNH GEO20591. 125. Dorsally fused anterior margins of tegumen (dorsal view). Zythos turbata BMNH GEO20371. 126. Plate-shaped sclerotization at proximal end of ductus ejaculatorius. Antilycauges pinguis BMNH GEO20381. 127. Sclerotization of vesica. Scopula rivularia BMNH GEO20684. 128. Dorsal diverticulum of vesica with sharp sclerotization at apex. Problepsis plagiata PS709. 129. Parallel folds of vesica. Zythos turbata BMNH GEO20371. 130. Signum. Somatina indicataria PS834. 131. Bare zone of signum. Zythos obliterata BMNH GEO20592. 132. Flexible lamella antevaginalis divided into two parts. Zythos avellanea PS880.

Male genitalia

• 91. Uncus: (0) present; (1) absent.




• 92. Socii: (0) absent; (1) fused at apex, dorsal of tuba analis (Fig. 40); (2) fused at apex, ventral of tuba analis (Fig. 117); (3) separate (Fig. 43). The fused, setose structure lying ventrad of the tuba analis in Problepsis has been considered to be a subscaphium (Holloway, 1997). Based on its position, general appearance and the fact that the tuba analis is visible dorsad of this fused structure, I consider the structure to be socii. In Epicosymbia, socii are fused, but lie dorsad of the tuba analis. I also consider Cinglis and Pseudoscinglis to have socii, contrary to Hausmann's (1994) interpretation. The paired, asymmetrical structures found on the tegumen margin in Zythos are here considered to be socii, although the usual setae are absent.




• 93. Socii with two lateral appendices: (0) no; (1) yes (Fig. 117).




• 94. Socii asymmetrical: (0) no; (1) yes.




• 95. Socii widest at apex: (0) no; (1) yes.




• 96. Socii setae: (0) absent; (1) present.




• 97. Socii crossed: (0) no; (1) yes.




• 98. Gnathos: (0) present; (1) absent.




• 99. Coremata: (0) absent; (1) present, apex turned ventrally (Fig. 123); (2) present, apex straight.




• 100. Ventral margin of tegumen: (0) unmodified (straight); (1) modified, dentate; (2) modified, round lobe.




• 101. Anterior margins of tegumen fused dorsally: (0) no; (1) yes (Fig. 125).




• 102. Transtilla: (0) anterior margin connected to valvula via a membrane; (1) anterior margin sclerotized to valvula; (2) absent.




• 103. Transtilla wide plate: (0) yes; (1) no.




• 104. Valva: (0) sacculus and valvula fused, line of fusion not visible; (1) sacculus and valvula fused, line of fusion visible; (2) sacculus and valvula separate.




• 105. Sacculus of valva: (0) symmetrical; (1) asymmetrical (Fig. 44).




• 106. Right sacculus of valva bifurcate: (0) no; (1) yes.




• 107. Valvula of valva ventral of sacculus: (0) no; (1) yes (Fig. 124).




• 108. Valvula of valva: (0) symmetrical; (1) asymmetrical.




• 109. Valvula apex of valva: (0) nondentate; (1) dentate.




• 110. Medial margin of sacculus upturned: (0) no; (1) yes.




• 111. Curved lobe on lateral margin of sacculus: (0) no; (1) yes (Fig. 39).




• 112. Vinculum size: (0) small; (1) enlarged.




• 113. Vinculum shape: (0) symmetrical; (1) asymmetrical (Fig. 43).




• 114. Vinculum margin: (0) convex (Fig. 41); (1) concave (Fig. 36).




• 115. Posteriorly directed lobe on top of vinculum with a medial ridge: (0) absent; (1) present (Fig. 49).




• 116. Anterior margin of juxta with wing-like processes: (0) absent; (1) apex free (Fig. 52); (2) apex fused to sacculus of valva; (3) one apex free, other apex fused.




• 117. Anterior margin of juxta with long processes laterally: (0) absent; (1) present.




• 118. Right arm of wing-like juxta processes longer: (0) no; (1) yes.




• 119. Only anterior margin of wing-like juxta processes sclerotized: (0) no; (1) yes.




• 120. Lobes of aedeagus: (0) absent; (1) present.




• 121. Round lobe on dorsal side of aedeagus: (0) absent; (1) present.




• 122. Distal part of aedeagus: (0) smooth; (1) with sclerotized spines (Fig. 128).




• 123. Apex of aedeagus with several separate spines: (0) no; (1) yes.




• 124. Vesica spiral: (0) yes; (1) no.




• 125. Sclerotizations of vesica: (0) present; (1) absent.




• 126. Plate-shaped sclerotization at proximal end of ductus ejaculatorius: (0) present (Fig. 126); (1) absent.




• 127. Long, curved structure on vesica: (0) absent; (1) present (Fig. 127).




• 128. Several teeth on vesica: (0) absent; (1) present, on a plate; (2) present, separate.




• 129. Long, curved cornutus at proximal end of ductus ejaculatorius: (0) absent; (1) present (Fig. 46).




• 130. Plate-shaped sclerotization on apical diverticulum: (0) absent; (1) present (Fig. 40).




• 131. Long diverticulum dorsally with sharp sclerotization at apex: (0) absent: (1) present (Fig. 128).




• 132. Parallel folds on vesica: (0) absent; (1) present (Fig. 129).

Female genitalia

• 133. Signum: (0) absent; (1) spines separate, point away from centre (Fig. 130); (2) spines fused, point away from centre (Fig. 86).




• 134. Fused signum spines visible at edges only: (0) no; (1) yes.




• 135. Bare zone at signum laterally at both sides of median ridge: (0) no; (1) yes (Fig. 131).




• 136. Long spines on inner surface of corpus bursae: (0) present, hollow (Fig. 80); (1) present, solid (Fig. 96); (2) absent.




• 137. Proximal part of corpus bursae sclerotized: (0) no; (1) yes.




• 138. Spirals of ductus bursae: (0) one (Fig. 80); (1) less than one.




• 139. Lamella antevaginalis flap-like, flexible: (0) no; (1) yes, one plate (Fig. 99); (2) yes, divided into two parts (Fig. 132).

Ecology

140. Species diurnal only: (0) no; (1) yes.

RESULTS

The initial search of 141 characters resulted in 20 equally parsimonious cladograms (L = 853, CI = 0.22; RI = 0.59). More efficient search strategies did not find any more equally parsimonious cladograms, nor did the ‘best’ command eliminate any of the original trees. A strict consensus tree of these appears in Figure 133(L = 876, CI = 0.21; RI = 0.58). Figure 134 shows one of the original trees (19 of 20) with the characters plotted. This is shown to give an indication of the supporting characters and to show how certain genera may be related, in contrast to the unresolved nodes of the strict consensus cladogram. The strict consensus cladogram is simplified to show the generic relationships within Scopulini in Figure 135.

Figure 133.

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Strict consensus of 20 equally parsimonious cladograms. based on 141 characters of adult morphology and ecology (L = 876, CI = 0.21; RI = 0.58). Species are mentioned in their generic combinations prior to analysis. Names on the right-hand margin indicate genera as recognized here.

Figure 133.

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Strict consensus of 20 equally parsimonious cladograms. based on 141 characters of adult morphology and ecology (L = 876, CI = 0.21; RI = 0.58). Species are mentioned in their generic combinations prior to analysis. Names on the right-hand margin indicate genera as recognized here.

Figure 134.

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Cladogram 19 of 20 based on 141 characters from adult morphology and ecology (L = 853, CI = 0.22; RI = 0.59). Numbers above hash marks indicate character numbers and below character states. Black bars indicate unique and white bars non-unique synapomorphies. Species are mentioned in their generic combinations prior to analysis. Names on the right-hand margin indicate genera as recognized here.

Figure 134.

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Cladogram 19 of 20 based on 141 characters from adult morphology and ecology (L = 853, CI = 0.22; RI = 0.59). Numbers above hash marks indicate character numbers and below character states. Black bars indicate unique and white bars non-unique synapomorphies. Species are mentioned in their generic combinations prior to analysis. Names on the right-hand margin indicate genera as recognized here.

Figure 135.

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A simplified cladogram of Figure 133 transformed to show generic relationships within the Scopulini. Taxa of uncertain generic association (incertae sedis) are shown in grey.

Figure 135.

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A simplified cladogram of Figure 133 transformed to show generic relationships within the Scopulini. Taxa of uncertain generic association (incertae sedis) are shown in grey.

In total, 22 characters were found to support the monophyly of the Scopulini (Fig. 134, Table 3). This is partially artificial since only two taxa from the putative sister group (Sterrhini) were used as outgroup. Cladistic analysis of Sterrhinae tribes has revealed that many of the synapomorphies supporting monophyly of the Scopulini would not do so if more outgroup taxa were included in the analysis (Sihvonen & Kaila, 2004).

According to this analysis there are two main lineages within Scopulini (Figs 133, 135; for supporting characters, see Diagnoses). The first includes Isoplenodia, Dithalama, Somatina and Zythos. Dithalama was found to be paraphyletic with regard to Isoplenodia, and the relationships of Somatina and Zythos are unresolved. One taxon, Somatina eurymitra, is incerta sedis(see Diagnosis of Somatina). The second lineage includes Lipomelia, Problepsis and Scopula. Lipomelia and Problepsis were recovered as sister groups. The sister group of Scopula is unknown, as there is a trichotomy in this lineage involving Lipomelia, Problepsis, Scopula, while two taxa (Somatina irregularis and Scopula haemaleata) are incertae sedis(see Diagnoses of Somatina and Scopula).

The relationships of species groups within Scopula are largely unresolved, but the species groups themselves often have many supporting synapomorphies. Many characters showed extensive homoplasy, for example the presence of a posteriorly directed lobes on top of the vinculum (Ch. 115, CI = 0.05; RI = 0.37, Fig. 49) and the presence of sclerotized spines on the distal part of the aedeagus, i.e. carina (Ch. 122, CI = 0.09; RI = 0.28).

DISCUSSION

GENERIC CLASSIFICATION OF THE SCOPULINI

The classification proposed here diverges markedly from previous classifications (see ‘Systematic Treatment’ and Appendix). This is not surprising if we consider that there has not been a previous attempt to study systematically the material throughout the morphological and geographical range of the tribe.

The generic concepts adopted here inevitably result in groupings that contain externally different species. This is most pronounced within Scopula, as it is considered to include species from previously recognized genera such as Glossotrophia, Antitrygodes and Aletis. However, from the morphological point of view, the recognized genera appear to represent rather well the material I have examined. In most instances the main groups were fairly apparent from morphological study and comparison, and these groupings were confirmed later by cladistic analysis.

CHARACTERS AND TAXON SAMPLING

The cladistic analysis was based on 141 characters, coded from adult morphology and ecology. Although the analysis covered the morphology very extensively, much promising phylogenetic information was observed that could not be incorporated into the data matrix. In most cases this was due to the quantitative nature of the morphological variation, which prevented unambiguous coding. For example, the shape of the pouch, situated on the posterior margin of the male metathoracic coxa (Ch. 55, Fig. 108), was found to be an area of extensive variation, in addition to the shape of the vesica, and length and arrangement of antennal sensilla in males. A further example is the shape of the outer membrane of the anterior opening of the aedeagus that encloses the ductus ejaculatorius. Unfortunately, this membrane was routinely removed before I realized its possible phylogenetic significance (Rose, 1985). Due to the shortage of material, immature stages were not included in the study, and they certainly need to be studied more extensively in the future.

Many characters previously used extensively in systematics of the Scopulini were found to be homoplastic. One example is the number of forewing areoles. Problepsis and Scopula have been considered to be one-areole genera, whereas the two-areole condition was found to be true for Somatina(e.g. Sterneck, 1941; Prout, 1920-41). However, two areoles are also present in a few taxa of Problepsis and Scopula(Ch. 48, CI = 0.14; RI = 0.76). In addition, the putative features of Glossotrophia(i.e. the absence of medial spurs of the female hindtibia and the presence of a dark terminal line that curves around the forewing apex) appeared homoplastic, as already noted by Hausmann (1993c). Clearly, if species are associated with certain genera on the basis of one homoplastic character alone, nonmonophyletic groupings result.

The number of analysed taxa is potentially adequate to recover the major groupings of the Scopulini and reveal the morphological variation within them. The number need not be increased markedly in future studies. The geographical coverage of taxa analysed was also extensive and adequate to recover the generic relationships. However, more taxa and characters are going to be needed to establish the relationships of species within the large genus Scopula; these were largely unresolved in the analysis.

FROM CLADOGRAM TO CLASSIFICATION

Transforming the information provided by cladograms into a formal classification is problematic because there is no objective way to define what constitutes a category, such as a genus. For the classification to have validity, one should attempt to delimit the category at the generic level and above so that they maximise their value in communication. If the categories are too narrowly or widely delimited, they may fail to adequately convey the relatedness of species.

This project has been undertaken with the aim of providing a more stable classification for the Scopulini. The problem with previous classifications has been the lack of global perspective, resulting in relatively restricted generic schemes. The present approach has involved drawing together information from earlier studies and adding new observations. As a result, many new generic synonyms and species combinations are proposed. A similar approach was used by Scoble & Krüger (2002) when they classified the world fauna of Macariini.

Groups of species are here recognized as genera if they are monophyletic and supported by synapomorphies that are unique or have low homoplasy. I have emphasized similarities rather than differences. I have also tried to avoid making new generic synonyms if the material studied is somehow contradictory or insufficient. One could argue that instead of proposing new generic synonyms for Scopula(see Appendix), I could have retained many of the traditional genera. However, had I done so, it would have either rendered the remaining taxa paraphyletic or split lineages into a large number of small, often new, genera. Furthermore, a classification where genera are very narrowly delimited can lead to the frequent addition of new genera when new species with slightly differing character combinations are discovered.

In the present paper, some genera are recognized that contain externally very different species, while others are more homogeneous. This is because I have tried to identify and name only monophyletic groupings rather than artificial classes. It may be argued that this approach can result in a reduction both of taxonomic resolution and of the overall comprehensiveness of the classification. However, as species groups within genera can still be recognized, identified and named, this need not be the case.

Finally, it is worth emphasizing that the majority of the genera of Scopulini were described in the 19th century. Generic taxonomy, for example, was based on comparison of external features alone. It is only natural that with recent advances, such as high-magnification microscopes, synapomorphy-based phylogenetic reasoning and parsimony software, results may be obtained that contradict the earlier classifications.

Consequently, the proposed classification has one large genus, Scopula, that contains 800 valid species, i.e. over 85% of the described fauna. Problepsis, Zythos and Somatina are middle-sized genera that include together over 100 species. Dithalama tentatively contains four species, whereas Lipomelia is monotypic. Isoplenodia is tentatively retained as a valid, monotypic genus although its recognition makes Dithalama paraphyletic. More material on Dithalama and Isoplenodia is needed to better resolve the relationships. A few species are incertae sedis, but they are retained in their traditional generic positions until further evidence is available (see Appendix).

FUTURE RESEARCH

Although this work has hopefully clarified many problems endemic to the systematics of the Scopulini, further large-scale research is needed to test the validity of the results and to establish a more precise understanding of generic relationships. Emphasis should be placed on the study of immature stages, in addition to a detailed morphological study of adults. The use of molecular characters is likely to be useful when attempting to solve intra- and intergeneric relationships, although it is difficult to obtain material that extensively covers the study group. It would be important also to study possible associations of a few Neotropical genera, i.e. Acratodes Guenée, Haemalea Hübner, Leptostales Möschler, Proutoscia Schaus, and Pseudasellodes Warren (Sihvonen & Kaila, 2004) with the Scopulini.

As is evident from this study, species-level revisions of many genera in the tribe are still needed (see also Hausmann, 1999b). No doubt many species are still misplaced, although they can be associated with related species with the help of the generic diagnoses provided here. For example, Somatina is a nonmonophyletic assemblage of species, whose generic association with Scopula, Dithalama and Problepsis needs to be revised.

Most importantly, a worldwide taxonomic revision of Scopula, the largest genus of the tribe, is essential. Many species of this genus are difficult to identify based on external features alone, and the lack of comprehensive identification literature has resulted in extensive synonymy. This practice still continues. If the defect is not corrected, future research on this group will be bedevilled with nomenclatorial problems arising from synonymies. The example of Scopula floslactata is a case in point. In addition to the valid species name, this species was described 12 (!) times between 1763 and 1951. One can only imagine how much time it has taken for later authors to correct mistakes.

I hope that this study and the proposed classification will provide a solid basis for future research on Scopulini systematics and taxonomy. Comprehensive identification tools, whether in the form of literature or internet-based databases, are urgently required. Once completed, they will provide a basis for applied studies on these moths, including the reconstruction of evolutionary issues such as the evolution of lacriphagy (Bänziger & Fletcher, 1985) and the shift from nocturnal to diurnal modes of life.

SYSTEMATIC TREATMENT

DIAGNOSES FOR THE TRIBE SCOPULINI AND ITS GENERA

A generic key to the male Scopulini is given, although it must be borne in mind that many characters are homoplastic. Use of a key that relies on a single or a few possibly homoplastic characters can result in arbitrary generic or even tribal combinations because no all-inclusive external features have yet been allocated that can be applied in order to separate species between Scopulini, Sterrhini and Cosymbiini. A key to the females is not provided due to the lack of material. A less formal approach to identify the Scopulini and component genera is more effective than a dichotomous key. The reader is therefore referred to characteristic tribal and generic features that are italicized in the diagnoses (see also Table 3). Many species can be assigned to genera by focusing on these few italicized characters alone. It must be remembered that even though some genera and species can be recognized from external features, in most cases it is necessary to study the genitalia. Important structures relating to generic placement are found also in the male coxa, hindleg and 8th sternite and tergite.

SCOPULINI DUPONCHEL, 1845

Definition of the tribe is based on a cladistic analysis of the Sterrhinae by Sihvonen & Kaila (2004), where two synapomorphies were found: the signum of the female genitalia being ovoid, granulate, with spines pointing away from the centre (Fig. 130), and the absence of a medial ridge on the epinotum of the male metathorax (character state ‘present’ is figured in Sihvonen & Kaila, 2004). In that analysis neither of these characters was considered to be unique to the tribe, see Results. In the present study it was found that they were unique. In addition, 20 other characters, 18 of which are unique, are proposed as possible tribal synapomorphies (Table 3). The male 8th tergite with the posterior margin concave with two round lateral lobes (88: 1, Figs 118, 122), may be a true synapomorphic character. Also characteristic are the absence of uncus and gnathos, and the presence of socii.

Wings

Wingspan c. 11–70 mm, usually 25-40 mm. Shape usually triangular, forewing apex elongated or termen occasionally rounded; hindwing margin often concave between vein ends or elongated at M₃. Facies very variable; many are straw-coloured, with numerous wavy transverse lines running from forewing to hindwing, and with small, dark discal spots(e.g. Figs 8, 21); ocellate discal markings and various other fasciae on white or grey background are also common; iridescent scales present in many species; forewings with fine striae and wide, dark medial band are not unusual; underside often fuscous, above patterns weakly repeated underside; underside ground colour uniform ochreous in some; transverse lines variably developed, usually all present, occasionally absent; sub- and adterminal shaded lines usually present, occasionally pronounced; terminal line continuous, discontinuous or absent; fringes unicolorous or chequered. Forewings with one or two areoles; R₅ from proximal or distal areole, either free or stalked with R₂-R₄ or stalked with M₁. Hindwing Sc + R₁ and Rs fused shortly or Sc and Rs separate but connected via R₁.

Head

Labial palpus upturned, 3-segmented, organ of vom Rath distinct. Pilifer present, rarely absent. Proboscis fully developed; variably shaped sensilla styloconica at distal end, absent in many species. Frons convex. Two openings or extensions on anterior part of dorsal sclerite between antennae either present or absent. Compound eye round to elliptic; proximal pigment layer circular or semicircular. Scales between antennae flat or flat and hair-like. Male antenna usually fasciculate, occasionally bipectinate to two-thirds; if bipectinate, proximal rami sometimes elongated; sensilla in single or multiple rows. Female antenna usually fasciculate, simple, occasionally sensilla arranged in single or even multiple rows. Antennal segments dorsally uni- or bicolorous. Chaetosemata usually from naked surface, occasionally from scaled surface.

Thorax

Foreleg with well-developed epiphysis; tibial spurs absent; thick spines on tarsus usually absent. Midleg with two tibial spurs, symmetrical or asymmetrical; tarsus normally developed, 5-segmented, thick spines usually absent, sometimes present. Male hindleg tibia often with outer and/or inner hair pencil proximally (Fig. 113), or absent; distal spurs two, one or absent; reduction in tarsus length common; all or part of tarsus segments may be fused; yet usually fully developed, 5-segmented; thick spines usually absent; two apical spines present in most species; usually two claws, symmetrical or asymmetrical, occasionally one or absent; arolium and pulvillus usually present. Female hindleg tibia usually with 2 + 2 spurs, occasionally with one medial spur or absent altogether; hindleg tarsus normally developed. Anterior and posterior ventral lamina of male metafurca usually present, occasionally absent; margins of metafurca basally opened or straight(Fig. 110); pouch on posterior margin of metathorax coxa usually present(Fig. 108), semiround, occasionally pronounced; medial ridge absent on epinotum of male metathorax dorsal sclerite; lateral processes of male coxa elongated in many species, variable in size and shape, rarely absent; absent in females.

Abdomen

Male 2nd sternite usually with distinct, round pouch(Fig. 116) between tympanal organs, posterior margin often invaginated, occasionally with two chambers, surface smooth, with fine ornaments or stout setae; anterolateral extensions present or absent. Male 6th segment occasionally with setae. Membranous sac between male 7th and 8th sternite usually present(Fig. 66). Male 8th sternite very variable; anterior margin convex, concave, tri- or bifurcate; cerata and mappa present in many species(Figs 62, 76); posterior margin membranous or sclerotized; posterolateral appendices occasionally present. Male 8th tergite variable; posterior margin usually concave, with two round lateral lobes(Figs 118, 122). Female abdomen simple, without modifications.

Male genitalia

Genitalia capsule appearing ovoid when viewed ventrally, characterized by enlarged vinculum and usually short, separate valvula and sacculus of valva. Uncus and gnathos absent. Socii usually setose, separate; occasionally fused at apex, lying ventrally or dorsally of tuba analis. Ventral margin of tegumen straight or with various processes, sometimes with distinct teeth. Valvula and sacculus of valva usually separate(e.g. Fig. 41), fused in a few cases; sacculus lying ventrally of valvula; both processes often asymmetrical; valvula curved ventrally, weakly sclerotized, setose, sometimes strongly sclerotized, acute; sacculus wide at base, tapering towards apex, blunt or acute; lateral and ventral extensions in many species. Coremata present(Fig. 123) at lateral end of transtilla, often pronounced or even bent ventrally; lost easily, but surface of genitalia distinctly granulate. Dorsal part of juxta tunnel-shaped; ventral part often with wing-like processes(Fig. 52), often asymmetrical. Transtilla usually present, occasionally anterior margins of tegumen fused replacing it. Vinculum enlarged, often with posteriorly directed lobe on top(Fig. 49). Aedeagus shape and size variable, often with dorsal lobes or carina at apex; caecum small or moderate; outer membrane of aedeagus expanded to semiround sac covering proximal end of ductus ejaculatorius; vesica very variable; simple sac with moderate diverticula or very complex with numerous coiled, twisted diverticula; plate-shaped sclerotization at proximal end of ductus ejaculatorius usually present(Fig. 126); otherwise variably sclerotized, occasionally with distinct cornuti. Spermatophore elongated sac, size and shape variable; occasionally frenum distinct, curved structure.

Note: special care is needed to interpret homologies of the male genital structures. Socius, and especially valvula, sacculus and juxta often replace each other functionally. For example, the left arm of the juxta in Somatina anthophilata Guenée has functionally replaced the left sacculus of the valva, the latter being diminished in size.

Female genitalia

Papillae anales longitudinally grooved, setose, sometimes bilobed into rounded dorsal and shorter, narrower ventral part. Apophyses posteriores usually longer than apophyses anteriores. Lamella postvaginalis pronounced in some species, usually weakly developed or absent. Ostium bursae cup-shaped, a complex of structures, occasionally simple or virtually absent. Lamella antevaginalis usually flap-like(Figs 99, 132), flexible or undeveloped in those species that have a well-developed lamella postvaginalis. Ductus bursae membranous or variably sclerotized; ductus seminalis arises from ductus bursae, corpus bursae neck or corpus bursae. Corpus bursae sac-like, occasionally heavily folded; signum an ovoid field of separate spines(Fig. 130), pointing away from centre, either fused or absent along medial axis in some species; spines on inner surface of corpus bursae common.

Distribution and species diversity

Virtually cosmopolitan with 912 described species and many more undescribed.

Immature stages

Due to the shortage of material, it was not possible to include immature stages in the cladistic analysis. Patocka (2003: 270), who worked with Central European fauna only, was able to find only quantitative delimiting pupal features for Scopulini. He diagnosed the pupa as follows: the ‘front leg verges on antennae as far as the compound eye extends’. In the Rhodostrophini, by contrast, the ‘front leg verges on antennae beyond the compound eye’. In addition, cremaster setae D2 of Scopulini are closer to each other than in Rhodostrophiini.

The Glossotrophia species group of Scopula has the unique condition of a free and loped elongated proboscis extending beyond the distal part of the pupa (Hausmann, 2001; Patocka, 2003). Chaetotaxy and pupal characters are described under generic concepts, where data are available. The narrow larvae of many species rest in a stick-like posture at an angle of 45 °. Some specialization in larval host-plants can be seen: Oleaceae in Problepsis and Rubiaceae in some Somatina. However, frequent polyphagy in Scopula and the worldwide distribution of the tribe make generalizations difficult.

Biology

Species can be found in forested and open habitats. A few Scopula species are known to be of minor pest status (see Introduction).

Comments on systematics

In total, 22 characters were found to support the monophyly of Scopulini (Fig. 134, Table 3). This is partially artificial since only two taxa from the putative sister group, Sterrhini, were used as outgroup. It is known from the cladistic analysis of the tribes of Sterrhinae (Sihvonen & Kaila, 2004) that many of the characters that appear here as synapomorphies supporting the monophyly of Scopulini would not do so if more outgroup taxa were present. Scopulini also includes Aletini (Hampson, 1918) and Problepsini (Wiltshire 1990) as indicated by Holloway (1996, 1997). Both of these tribes share with Scopulini their characteristic structural features, including those of male 8th sternite, male and female genitalia and wing venation.

THEDITHALAMA, ISOPLENODIA, SOMATINAANDZYTHOSGROUP OF GENERA

This group of four genera is characterized by five synapomorphies (Figs 133-135): presence of two forewing areoles formed by R veins (48: 1, six occurrences and one reversal); forewing vein R₅ runs from the distal areole and is stalked with veins R₂-R₄(49: 2, eight occurrences); posterolateral appendices are absent on male 8th sternite (86: 1, five occurrences and six reversals); the medial margin of the sacculus is upturned (110: 1, four occurrences); lamella antevaginalis is flap-like, flexible, formed by one plate (139: 1, 13 occurrences and two reversals). Within the group, the relationships are not resolved. Somatina eurymitra is incerta sedis and recognition of Isoplenodia renders Dithalama paraphyletic. These three taxa form the sister group to Somatina+Zythos.

Comments on systematics

This group of genera, although not found to be supported by unique synapomorphies (Figs 134, 135), appears to be rather distinctive. Cerata are absent in most cases, and the species have two areoles in the forewings. In the other group of genera (i.e. Scopula, Lipomelia and Problepsis) the cerata are fully developed and in most species the forewings have only one areole.

DITHALAMA MEYRICK, 1888 (FIGS3, 32, 57, 82, 119, Appendix)

Diagnosis:

description is based on type species only (see ‘Comments on systematics’, below). Monophyly supported by five synapomorphies: transverse median line of wings is obscure (25: 1, 12 occurrences); posterior margin of male 8th tergite is concave, with two sclerotized lateral spines (88: 2, unique, Fig. 119); medial margin of sacculus is not upturned (110: 0, three occurrences); presence of posteriorly directed lobe on top of vinculum with a medial ridge (115: 1, 15 occurrences); apices of wing-like juxta processes are fused to sacculus of valva (116: 2, seven occurrences).

Imago:

wingspan 19–24 mm. Male antenna fasciculate, sensilla in single rows; female antenna fasciculate, simple. Pilifer present. Wing patterns variable, transverse lines ranging from almost straight to hardly recognizable; discal spot dark; terminal line discontinuous; wing patterns repeated on underside but indistinct. Forewing with two areoles; R₅ from distal areole, either stalked with R₂-R₄ or straight from accessory cell. Outer margin hair pencil of male hindleg tibia present, inner either present or absent; distal spurs of hindleg tibia absent; tarsomeres shortened but not fused; claws either two in number or absent. Female hindleg tibia with two pairs of spurs.

Male abdomen and genitalia:

pouch on male 2nd sternite round; anterolateral extensions present or absent. One membranous pouch between male 7th and 8th sternite. Male 8th sternite sclerotized; anterior margin elongated medially; laterally turned extensions or two blunt lobes on posterior margin; cerata, mappa and membranous posterolateral appendices absent(Fig. 57). Posterior margin of male 8th tergite concave with two sharp cerate-like projections or with two round lateral lobes (Fig. 119). Socii separate, setose. Ventral margin of tegumen with round lobes, separate or fused. Valvae symmetrical, variably sclerotized, anterior margin of sacculus medially upturned or levelled. Transtilla present. Anterior margin of juxta with wing-like processes, apex fused to sacculus of valva or absent. Vinculum enlarged. Aedeagus straight with one or several teeth, carina, at distal end; vesica simple, curved sac, without sclerotizations.

Female  genitalia:

lamella antevaginalis sclerotized, flap-like, flexible, medially elongated. Ostium bursae weakly sclerotized; ductus bursae wide, proximal part either membranous or sclerotized; ductus seminalis arises from ductus bursae. Corpus bursae elongated sac; signum either granulate, spines point away from centre or absent; long spines on inner surface of corpus bursae absent.

Distribution and species diversity:

Australia and Tasmania. Four species identified tentatively (McQuillan & Edwards, 1996). See Appendix under Dithalama.

Immature stages:

unknown.

Biology:

unknown, although D. cosmospila is suspected to live on Eucalyptus(Prout, 1920-41).

Comments on systematics:

the genus is in need of revision, as was already noted by Prout (1920-41). He speculated that Dithalama is closely related to Somatina and Autanepsia, and the latter monotypic genus should perhaps be combined with Dithalama. In this analysis Dithalama was recovered as paraphyletic with regard to Isoplenodia. The Dithalama+Isoplenodia connection, although not discussed earlier, is supported by a number of character absences (Fig. 134). In addition to Dithalama+Isoplenodia, the secondary loss of sclerotizations of vesica (125: 1), which is a derived feature in this analysis, has occurred outside of these taxa only in Tasmanian species Dasybela achroa and South African species Zygophyxia relictata, which were found to be unrelated to Dithalama. Absence of male hindleg claws (67: 0), pretarsus arolium (68: 0) and pulvillus (69: 1), on the other hand, are found also in a number of taxa in Zythos, Problepsis and Antitrygodes.

ISOPLENODIA PROUT, 1932 (FIGS4, 33, 58)

Diagnosis:

monophyly supported by eight synapomorphies: absence of pilifer (character and character state 0: 1, unique); ventrolateral sensilla on proximal part of male flagellomere are in multiple rows (8: 1, nine occurrences); absence of anterior ventral lamina of metathoracic metafurca (52: 1, six occurrences); absence of opening on male metathoracic dorsal sclerite (58: 1, character state ‘present’ shown in Fig. 112, unique); fusion of male hindleg tarsomeres 1–5 (66: 1, two occurrences); cerata are fully developed (79: 2, three occurrences, two reversals); mappa is bare 83: 2 (four occurrences); ventral margin of tegumen is unmodified, straight (100: 0, three reversals).

Imago:

wingspan c. 16 mm. Head smooth-scaled; male antenna fasciculate; sensilla arranged in multiple rows, proximal row elongated. Pilifer absent. Wings light brown, transverse lines weakly expressed; discal spot dark; terminal line discontinuous, equally wide; underside light fuscous. Forewing with two areoles; R₅ from distal areole, stalked with R₂-R₄. Opening on male metathorax dorsal sclerite absent. Outer margin hair pencil of male hindleg tibia short; inner absent; tarsomere segments 1–5 fused, claws absent. Female hind tibia with 2 + 2 spurs.

Male abdomen and genitalia:

anterolateral extensions on male 2nd sternite absent. One membranous pouch between male 7th and 8th sternite. Anterior margin of male 8th sternite elongated medially; cerata symmetrical; mappa constricted at base; membranous, posterolateral appendices absent. Posterior margin of male 8th tergite concave. Socii rudimentary, short. Ventral margin of tegumen straight. Valvae symmetrical, soft, setose, valvula and sacculus separate. Anterior margin of sacculus medially upturned, covered with pointing setae. Transtilla present. Juxta weakly sclerotized, narrow at base, but with long wing-like processes. Vinculum enlarged, dorsally narrow, v-shaped(Fig. 33). Aedeagus curved ventrally, with long caecum; without sclerotizations.

Female genitalia:

unknown.

Distribution and species diversity:

known from Madagascar only. One species.

Immature stages:

unknown.

Biology:

unknown.

Comments on systematics:

the genus is retained as valid but only tentatively (see comments on Dithalama). Prout speculated that Isoplenodia might be of a close relative of the African genera Epicosymbia and Isoplenia, on the basis of two forewing areoles, and long ciliated male antenna. This was not supported in this analysis and the condition of two forewing areoles (48: 1) was shown to be a very homoplastic character. I. arrogans was found to have cerata-like structures laterally on the male 8th sternite, but these are immobile and without apical setae, contrary to other studied material. Examination of female structures could give further clues about the systematic position of this species.

SOMATINA GUENÉE, [1858] (FIGS2, 27, 31, 56, 81, 120, APPENDIX)

Diagnosis:

monophyly supported by three synapomorphies: sacculi of valva are asymmetrical (105: 1, seven occurrences); valvuli of valva are asymmetrical (108: 1, seven occurrences); anterior margin of juxta is with wing-like processes, apex is fused to sacculus of valva (116: 2, seven occurrences).

Imago:

wingspan 23-33 mm. Male antenna fasciculate; proximal row of sensilla sometimes weakly elongated, those sensilla in single rows. Female antenna fasciculate, simple. Pilifer present. Wing patterns variable, colour ranging from white to reddish-yellow, greyish-brown in many species; often irrorated with dark scales; dark band on forewing termen between transverse anterior and median lines in many species; transverse lines dentate, straight or sometimes modified, taking unusual forms; discal spot weak, dark; iridescent scales usually absent, glossy in few species; terminal line usually discontinuous, equally wide; patterns often repeated on underside, fuscous, sometimes absent. Forewing with two areoles; R₅ from distal areole, stalked with R₂–R₄ or straight from cell. Outer and inner margin hair pencil of male hindleg tibia usually present, outer concealed in a furrow formed by long scales in many species; apical spurs usually absent, sometimes two; tarsomeres normally developed, 5-segmented; two claws. Female hind tibia with 2 + 2 spurs.

Male abdomen and genitalia:

pouch on male 2nd sternite round, posterior margin often invaginated; anterolateral extensions usually present. Small lateral pouches between sternites 3 and 4 rarely present. Usually one, sometimes two membranous pouches between male 7th and 8th sternite. Male 8th sternite variable, x-shaped and heavily sclerotized in many species; anterior margin elongated medially or bifurcate; cerata usually absent(Fig. 56), sometimes present as fully developed; mappa absent, setose or bare; posterolateral appendices usually present. Male 8th tergite posterior margin concave, with two round lobes or sharp projections(Fig. 120). Socii separate, covered with long setae. Ventral margin of tegumen unmodified, rarely with small round or elongated projections. Sacculus and valvula of valva separate, with pronounced asymmetry; valvula shape varies from wide, blunt-ended to narrow, long, with sharp apex; medial and lateral margin of sacculus often with upturned projections. Anterior margin of juxta with wing-like processes, apex rarely free, instead fused to sacculus in most species, and sometimes asymmetrical; sometimes juxta processes functionally replacing sacculus of valva. Transtilla narrow weakly sclerotized band, rarely wide. Vinculum enlarged, symmetrical and usually convex. Aedeagus bent, sometimes with small projections near apex; vesica complex, with several large, often twisted diverticula(Fig. 31); variably sclerotized but rarely with distinct cornuti.

Female genitalia:

lamella postvaginalis usually absent; lamella antevaginalis large, sclerotized, medially invaginated, flap-like structure in most species. Ostium bursae weakly sclerotized. Ductus bursae usually coiled, weakly sclerotized. Ductus seminalis opens from corpus bursae neck. Corpus bursae ovoid sac; signum elongated, made of separate spines, rather large compared to size of corpus bursae.

Distribution and species diversity:

from Africa to the Oriental region and Australia. Forty-four species and several subspecies, although see ‘Comments on systematics’.

Immature stages:

I have been unable to find any references. Nakamura (1994) described the pupa of Somatina indicataria(Walker), but the taxon is considered to belong to Scopula in the present study.

Biology:

S. anthophilata has been recorded from Gardenia jasminoides and Randia dumetorum(Rubiaceae) and Rosa(Rosaceae). Somatina omicraria and S. virginalis have been recorded from the Oleaceae genus Jasminum(Robinson et al., 2002).

Comments on systematics:

Sterrhinae species with two areoles on the forewings have traditionally been considered to belong to Somatina, in contrast to Problepsis and Scopula, which have one areole. Since this character was shown to be homoplastic in the present study (48: 1), and as both states occur in many lineages, Somatina was considered to be a polyphyletic assemblage of species, as noted by Inoue (1992), with many species formerly ascribed to it shown to belong to either Problepsis or Scopula.

While the remaining species should be revised against the generic descriptions given here and, if necessary, transferred to the appropriate genera (see Appendix for examples), Somatina is, however, likely to be a monophyletic group of species. It has a number of characteristic, although not unique, features. One such feature is the anterior margin of the juxta that is fused to sacculus of the valva (116: 2). The male of S. anthophilata, the type species of the genus, has small lateral pouches between sternites 3 and 4 (Holloway, 1997), although as these were not seen in other studied taxa, this autapomorphy was not coded. Traditionally, Somatina has been considered a close relative of Scopula or Problepsis(e.g. Prout, 1920-41,,1934-39), but in the present study species of the redefined genus are associated with Zythos. This connection is supported by the sclerotized posterior part of male 8th sternite (85: 1, Fig. 59), which is found elsewhere only in Dithalama desueta, and by the presence of parallel folds on the vesica (132: 1, 129), found also in a few Scopula taxa. Several characters, e.g. the structure of the socii and signum, indicate a close relationship with Scopula.

Somatina irregularis and S. eurymitra were considered incertae sedis, but were retained in their traditional generic combinations, as they did not group together with the recognized genera. Somatina irregularis grouped together with Scopula haemaleata, sharing with it a number of unique features, such as a bifurcated cerata (82: 1) and a juxta with long processes laterally (117: 1). Somatina eurymitra, meanwhile, showed affinities with Somatina and Problepsis; for example, the male 8th tergite is concave with two sharp lateral projections (88: 3), a feature typical of Somatina, while the cerata fused with the mappa (80: 1) is a common condition in Problepsis. Females of S. eurymitra were not available for study. It remains to be tested whether these taxa actually represent larger monophyletic groups.

ZYTHOS FLETCHER, 1979 (FIGS5, 34, 59, 83, 101, 103, 114, 121, 124, 125, 129, 131, 132,APPENDIX)

Diagnosis:

monophyly supported by 20 synapomorphies: iridescent scales of wings are restricted to a few rows, excluding discal spots (16: 3, unique); iridescent scales of wings are glossy and longitudinally grooved (17: 1, unique, Fig. 101); discal spot of forewing is not unicolorous and dark (19: 1, six occurrences);discal spot of forewing is lunular (20: 2, two occurrences); presence of a spot at vein endings at forewing margin (40: 1, two occurrences); absence of hair pencil on inner margin of male hind tibia (61: 1, seven occurrences); male hindleg is with one claw only (67: 1, unique, Fig. 114); absence of pretarsus arolium on male hindleg (68: 0, six reversals); two pouches are present between male 7th and 8th sternites (76: 1, four occurrences, Fig. 59); absence of posterolateral appendices on male 8th sternite (86: 1, five occurrences, five reversals); posterior margin of male 8th tergite is convex (88: 4, unique, Fig. 121), posterolateral margin of male 8th tergite has two membranous appendices (90: 1, Fig. 121); absence of setae on socii (96: 0, unique); anterior margins of tegumen are dorsally fused (101: 1, unique, Fig. 125); absence of transtilla (102: 2, unique); valva have symmetrical sacculi (105: 0, two occurrences); valvula of valva lies ventral of sacculus (107: 1, unique, Fig. 124); valvuli of valvae are symmetrical (108: 0, two occurrences); anterior margin of juxta is without wing-like processes (116: 0, nine occurrences); signum has a bare zone laterally on both sides of median ridge (135: 1, unique, Fig. 131).

Imago:

wingspan 32-40 mm. Male antennae fasciculate, sensilla in single rows; female antennae fasciculate, simple. Pilifer present. Sensilla styloconicae forming serrate lining at distal end of proboscis. Chaetosemata prominent. Wing pattern variable, often with fine striations on extensive areas of wings; costa and middle parts often olive grey; often with longitudinally grooved iridescent scales on outer half of forewing, excluding discal spots (Fig. 101); subterminal shaded line and terminal line touching each other at fore- and hindwings in many species; discal spots dark, often lunular on forewings; pale markings on anterior side of hindwing discal spots in many species; terminal line discontinuous, equally wide, dark spot at vein end at forewing margin; underside often unicolorous, ranging from light brown to light orange. Forewings with two areoles; R₅ from distal areole, either stalked with R₂–R₄ or straight from accessory cell. Hair pencil of outer margin of male hindleg tibia present, point of origin prominent; inner margin hair pencil absent; tarsomeres normally developed; one claw(Fig. 114); arolium absent; pulvillus present. Female hindleg tibia with 2 + 2 spurs.

Male abdomen and genitalia:

pouch on male 2nd sternite round, large; anterolateral extensions present. Two membranous pouches between male 7th and 8th sternite. Male 8th sternite heavily sclerotized, anterior margin bifurcate; posterior margin with blunt extensions or medial projections; cerata, mappa and membranous posterolateral appendices absent (Fig. 59). Male 8th tergite posterior margin convex, with two membranous appendices on posterolateral margin(Fig. 121). Genitalia capsule oval. Socii wide, separate, without setae. Ventral margin of tegumen slightly curved inwards. Valvae symmetrical, valvula and sacculus separate; anterior margin of sacculus medially upturned or levelled; valvula moved ventrad of sacculus(Fig. 124), often curved dorsally. Transtilla absent; instead, anterior margins of tegumen are fused dorsally(Fig. 125). Anterior margin of juxta without wing-like processes. Vinculum enlarged. Aedeagus almost straight, apex with small blunt extension; vesica prominent, long, often with long lateral diverticulum and sclerotized parallel folds.

Female genitalia:

lamella antevaginalis flap-like, flexible, medially either concave or bilobed. Ostium bursae heavily sclerotized. Ductus bursae membranous, coiled; ductus seminalis arises from ductus bursae. Corpus bursae an elongate sac; signum often with narrow medial, sclerotized strip followed by bare zone(Fig. 131) and granulate field of separate spines on both sides laterally.

Immature stages:

unknown.

Distribution and species diversity:

from India to Papua New Guinea. Morphologically, a compact genus with 11 species.

Biology:

adults occur predominantly in the understorey of lowland rain forests and have been collected from carrion-baited pitfall traps (Holloway, 1997). The serrated distal end of the proboscis indicates an adaptation to piercing. Frequently taken from secondary forests as well (Chey, 1994).

Comments on systematics:

Prout (1920–41), followed by Holloway (1997), considered Zythos to be a close relative of Ignobilia on the basis of striated facies and pale underside of wings. In the present study Zythos is associated with Somatina(see Discussion under that genus).

THELIPOMELIA, PROBLEPSISANDSCOPULAGROUP OF GENERA

This group of genera is characterized by one synapomorphy (Figs 133-135): the fully developed cerata (79: 2, six occurrences, six reversals). Within the group, the relationships are not resolved. Lipomelia and Problepsis form a compact group, Somatina irregularis and Scopula haemaleata are incertae sedis, and externally heterogeneous Scopula is characterized by a combination of features.

Comments on systematics

See the Dithalama, Isoplenodia, Somatina and Zythos group of genera.

LIPOMELIA WARREN, 1893 (FIGS6, 35, 60,APPENDIX)

Diagnosis:

monophyly supported by ten synapomorphies: forewing costa is wide and grey (11: 2, two occurrences); presence of pale markings on anterior side of hindwing discal spot (23: 1, two occurrences); subterminal shade line and terminal line are touching each other at fore- and hindwings (42: 1, two occurrences, Fig. 6); absence of male hindleg pretarsus arolium (68: 0, six reversals); absence of male hindleg pretarsus pulvillus (69: 1, three occurrences); sacculi of valva are asymmetrical (105: 1, seven occurrences); valvuli of valva are asymmetrical (108: 1, seven occurrences); vinculum is asymmetrical (113: 1, six occurrences); anterior margin of juxta is without wing-like processes (116: 0, nine occurrences); presence of parallel folds on vesica (132: 1, seven occurrences).

Imago:

wingspan c. 25 mm. Head smooth-scaled; male antennae fasciculate, sensilla in single rows. Pilifer present. Wings brown, base striated, dark area between transverse posterior line and subterminal shade line at forewings; transverse lines cryptic, subterminal shade line and terminal line touching each other at fore- and hindwings(Fig. 6); discal spots white, with iridescent scales, surface smooth; pale markings on anterior side of hindwing discal spots; terminal line discontinuous, equally wide; underside fuscous. Forewings with one areole; R₅ stalked with R₂–R₄. Outer and inner margin hair pencil of male hindleg tibia well developed; distal spurs of hindleg tibia absent; tarsomeres short, segments 4–5 fused; two claws; arolium and pulvillus absent. Female hindleg tibia with 2 + 2 spurs (Prout, 1920-41).

Male abdomen and genitalia:

round pouch and anterolateral extensions on male 2nd sternite present. One membranous pouch between male 7th and 8th sternite. Male 8th sternite anterior margin elongated medially; cerata present, coalescent at base(Fig. 60); mappa absent; membranous posterolateral appendices present. Male 8th tergite posterior margin concave. Socii short, setose, separate. Ventral margin of tegumen straight. Valvae asymmetrical, valvula and sacculus separate, ventral margin of left sacculus bifurcate. Transtilla present. Anterior margin of juxta with wing-like processes absent. Aedeagus curved dorsally; plate-shaped sclerotization at proximal end of ductus ejaculatorius; cornutus absent; vesica simple, large lateral diverticulum at base with sclerotized parallel folds, small diverticulum dorsally.

Female genitalia:

unknown.

Distribution and species diversity:

from India to Taiwan. One species.

Immature stages:

unknown.

Biology:

unknown.

Comments on systematics:

when Warren (1893) described Lipomelia, he did not discuss its systematic position. Prout (1920-41) considered it a close relative of Scopula on the basis of one areole on the forewing and male genital characters, although he noted that the facies of L. subusta resembles that of Zythos spp. In the present study, Lipomelia is associated with Problepsis on the basis of forewing iridescent scale characters. One of these characters, i.e. the structure of iridescent scales being glossy and smooth (17: 2, Figs 102, 134) is a unique synapomorphy of this clade. I have chosen not to synonymize Lipomelia with Problepsis, however, because the former lacks several characteristic features of the latter, e.g. ocellate discal spots on both pairs of wings (cf. Figs 6, 7), fused socii lying ventral to tuba analis (cf. Figs 35, 36, 117), and male 8th sternite anterior margin not being trifurcate and blunt (cf. Figs 60, 61). Male genitalia of Lipomelia, including 8th sternite, were found to be structurally similar to those of Scopula.

PROBLEPSIS LEDERER, 1853 (FIGS7, 27, 36, 61, 84, 102, 105, 106, 108, 115, 117, 128,APPENDIX)

Diagnosis:

monophyly supported by five synapomorphies: discal spot of forewing is ocellate and round (20: 3, three occurrences, Fig. 7); discal spot of hindwing is ocellate (22: 1, three occurrences); socii are fused at apex, lying ventral of tuba analis (92: 2, two occurrences, Fig. 117); ventral margin of tegumen is dentate (100: 1, two occurrences); anterior margin of juxta is with wing-like processes, apex is fused to sacculus of valva (116: 2, five occurrences).

Imago:

wingspan 25-50 mm. Male antennae fasciculate; sensilla in single or multiple rows, proximal row elongated in many species. Female antennae fasciculate, simple; sensilla in single rows. Pilifer present. Wings pale to white, greyish in few species; ocellated discal spots(Fig. 7) distinct; smooth, glossy, iridescent scales often present in discal spots or scattered throughout wings(Figs 102, 105, 106); transverse median line often wide, indistinct; subterminal shade line discontinuous, spotted; terminal line continuous or discontinuous, equally wide or wider at vein ends. Patterns not repeated or weakly expressed on underside. Forewings often with one areole, sometimes two; R₅ from proximal or distal areole, stalked with R₂–R₄. Outer margin hair pencil of male hindleg tibia present or absent; inner margin hair pencil often moderate, sometimes massive or absent; tarsomeres often reduced in length, 5-segmented or segments 4–5 fused; claws either two in number or absent. Female hind tibia with 2 + 2 spurs.

Male abdomen and genitalia:

pouch on male 2nd sternite round, posterior margin often invaginated; anterolateral extensions present or absent. One or two membranous pouches between male 7th and 8th sternite. Male 8th sternite elongated; anterior margin often trifurcate, blunt or sometimes bifurcate; cerata absent, rudimentary or fully developed, often fused to mappa; mappa present, bare; membranous posterolateral appendices present or absent. Male 8th tergite posterior margin concave. Socii fused at apex, ventral of tuba analis(Fig. 117), setose, sometimes with two lateral appendices. Ventral margin of tegumen sometimes unmodified, dentate in most species. Valvula and sacculus of valva separate, symmetrical, long, tapering towards apex, acute; medial margin of sacculus rarely upturned. Transtilla variably sclerotized, often invaginated medially. Anterior margin of juxta with wing-like processes, often anterior margin sclerotized only, apex fused to sacculus of valva, sometimes free. Vinculum enlarged, symmetrical, anterior margin often concave. Aedeagus often slightly bent, apex with one or several teeth; vesica large, complex, variably sclerotized, rarely with distinct cornuti; several diverticula, sometimes long and prominent.

Female genitalia:

lamella postvaginalis sclerotized in many species; lamella antevaginalis often absent, sometimes weakly developed, rarely flap-like, flexible. Ostium bursae often sclerotized; ductus bursae sclerotized, straight, wide(Fig. 84) in most species. Ductus seminalis arises from ductus bursae or corpus bursae neck. Corpus bursae ovoid, membranous sac. Signum absent sometimes, usually an ovoid patch of separate or fused spines, small compared to size of corpus bursae.

Distribution and species diversity:

most species occur from sub-Saharan Africa to Indonesia and Australia. Only a few reach the Palaearctic region. Fifty-one species and several subspecies.

Immature stages:

the only larval description is based on a species from south India, whose identity is uncertain. It is possibly P. deliaria Guenée (Bell in Holloway, 1997). The Japanese species illustrated in Sugi (1987) rest at 45 ° from the substrate, usually in the mid-rib of a leaf. Based on limited Japanese material (Nakamura, 1994), the pupal cremaster has one pair of setae only. Pupation takes place in a loose cocoon incorporating particles of earth, usually on the soil surface.

Biology:

all host-plant records relate to Oleaceae (Holloway, 1997). These include Olea(Holloway, 1997), Ligustrum(Sugi, 1987) and Jasminum(Singh, 1957). Many species are from lowland forest, including cultivated areas and dry heath, but Holloway (1997) reports species taken at heights of almost 1800 m in Borneo. In north-east China, P. phoebearia Erschov and P. plagiata(Butler) were taken from a lamp on a warm, south-facing sandy grove (pers. observ.).

Comments on systematics:

traditionally, Problepsis has been delimited to include species that have one forewing areole, large ocellated discal spots with iridescent scales and no spurs on male hind legs. Its closest relatives were assumed to be Antitrygodes, Scopula and Somatina(Sterneck, 1941; Prout, 1934-39; Holloway, 1997). In the present study, the clade comprising Lipomelia+Problepsis was found to be the closest relative of Scopula, in addition to two taxa incertae sedis(Fig. 133). Ocellated discal spots, whether round or semiround (20: 3 or 20: 4), were found to be a unique synapomorphy of Problepsis. Number of forewing areoles was shown to be a homoplastic character (Ch. 48), and although the majority of Problepsis species have one, two are also found, e.g. in P. centrophora(Prout). Absence of male hindleg spurs was found to be a plesiomorphic feature in Scopulini (62: 0). Fused socii of male genitalia (92: 2), which lie ventral to tuba analis, was found to be a diagnostic character delimiting the genus; it is present in all studied Problepsis taxa, and found outside of it in Antitrygodes cuneilinea(Walker) only. It is possible that a few Problepsis species are misplaced in Somatina, see Appendix. See also Discussion under Lipomelia.

SCOPULA SCHRANK, 1802 (FIGS8–26, 29, 37–54, 62–79, 85–100, 104, 107, 110–113, 116, 118, 122, 123, 126, 127, 130,APPENDIX)

Diagnosis:

monophyly supported by one synapomorphy: absence of posterolateral appendices on male 8th sternite (86: 1, five occurrences).

Imago:

wingspan 11–70 mm, usually 25-35 mm. Male antennae fasciculate; sensilla in single or multiple rows, proximal row sometimes elongated markedly; rarely bipectinate. Female antennae usually fasciculate, simple, occasionally sensilla arranged in single or even multiple rows. Pilifer present. Wings very variable, no single character can be used to define this group; almost all colours are found, but usually straw or grey-brown are dominant; often irrorated, sometimes unicolorous; iridescent scales absent; three wavy transverse lines running from forewing to hindwing are common, dentate in many species, reduced to series of spots in other or absent; sometimes two dark marks on forewings medial to transverse posterior line and another dark mark at forewing termen; few species with reticulate pattern; few with wide dark areas on outer third of wings; terminal line at forewings usually discontinuous, equally wide; smoothly curved around forewing apex occasionally or absent. Discal spot usually dark, small; cryptic in some, rarely absent; fringe usually unicolorous, occasionally chequered. Patterns usually repeated on underside, weak, sometimes absent. Forewings usually with one areole, sometimes two; R₅ usually from proximal areole, either stalked with R₂–R₄ or free, sometimes from distal areole, stalked with R₂–R₄ or free or even from M₁. Hindwing Sc + R₁ and Rs usually stalked for short distance, then divergent or sometimes connected by R₁. Outer and inner margin hair pencil of male hindleg tibia either present or absent; sometimes pronounced; if absent, replaced with two spurs at distal end in many species, occasionally one or absent; tarsomeres usually normally developed, sometimes reduced in length, 5-segmented; rarely segments 1–5 or 2–5 fused; two claws, absent in few groups. Female hindleg tibia with 2 + 2 spurs, sometimes only two distally.

Male abdomen and genitalia:

pouch on male 2nd sternite round, posterior margin occasionally invaginated or rarely elongated, surface smooth or with fine decorations, one- or occasionally two-chambered; occasionally laterally elongated; with or without pointing setae, sometimes absent; anterolateral extensions present or absent. Sometimes tufts of setae on male 6th and 7th segments laterally. Usually one membranous pouch between male 7th and 8th sternite, rarely two or absent. Male 8th sternite anterior margin often medially elongated, sometimes cup-like or bifurcate, rarely trifurcate; cerata usually present, asymmetrical, free(Fig. 62), with apical setae, or rarely fused to mappa, sometimes seen as bifurcate, rudimentary or absent; mappa usually present, bare(Fig. 76), sometimes setose or posterior margin separate, rarely absent; posterolateral appendices usually absent, pronounced occasionally; posterior margin of male 8th sternite not sclerotized. Male 8th tergite posterior margin usually concave with two round lobes(Figs 118, 122) or rarely with sharp projections; laterally constricted in few species. Socii size and shape variable, typically separate at apex, setose, well developed, slightly upturned, occasionally reduced to tuft of setae, rarely massive, or in few instances fused at apex, dorsal of tuba analis. Tegumen ventral margin usually unmodified, rarely with lateral nondentate projections. Transtilla present, usually U-shaped, occasionally W-shaped; weakly sclerotized. Sacculus and valvula of valva separate, partially fused in few instances; size and shape very variable; symmetrical in most cases but pronounced asymmetry not unusual; valvula usually soft, blunt-ended(e.g. Fig. 48), bent ventrally, setose, blunt-ended, often with apical spines; sacculus usually more sclerotized, tapering towards apex, sometimes acute or dark; bifurcate in some species; occasionally rudimentary; various projections may arise from lateral and ventral margins. Anterior margin of juxta typically with wing-like projections(Fig. 52), sometimes absent; apex usually free, but both or one extensions fused to sacculus in few species; often asymmetrical, size and shape variable; juxta often tube-shaped, well sclerotized with cup-like extension ventrally. Vinculum enlarged, margin typically convex, rarely concave; symmetrical, sometimes asymmetrical; cup-shaped and turned ventrally in few species. Aedeagus size and shape variable, in many instances narrow and straight or wide and slightly bent; caecum usually a continuation of aedeagus, enlarged occasionally; sharp projections at apex (carina) sometimes; apex sharp or blunt; vesica size and shape variable; often simple sac, with small diverticula and plate-shaped sclerotization at proximal end of ductus ejaculatorius; or large, complex, with elongated, twisted diverticula, variably sclerotized; distinct cornuti usual but more often sclerotizations are less definitive; pronounced plate-shaped sclerotizations at distal end of vesica occasionally; all instances between extremities occur.

Female genitalia:

papillae anales often round, ventral margin upturned occasionally. Lamella postvaginalis usually weakly developed, irregular plate or made of parallel folds or absent. Lamella antevaginalis usually sclerotized, flap-like, semiround(Fig. 99), posterior margin invaginated in few species; horseshoe-shaped in others, laterally wider; asymmetry usual; normally bare but occasionally heavily setose. Ostium bursae usually sclerotized, with lateral extensions, reaching proximal part of ductus bursae, dorsally membranous. Ductus bursae variable in length, width, shape and amount of sclerotization; membranous and coiled in many cases, or short and wide, or indistinct. Ductus seminalis arises from ductus bursae, corpus bursae neck or corpus bursae. Corpus bursae elongated sac, usually smooth but heavily wrinkled in others. Signum usually present as an elongated patch of separate spines(Fig. 130), sometimes two patches, which may be fused medially along axis, sometimes spines fused forming long and narrow band. Usually corpus bursae with spines on inner surface, length and density variable.

Distribution and species diversity:

virtually ubiquitous. The genus has successfully colonized even the remotest islands in Polynesia and New Zealand (Holloway, 1997; Dugdale, 1988). About 800 described species and numerous subspecies. See ‘Comments on systematics’ under Somatina.

Immature stages:

material is scarce in museums and pertinent literature is scattered, often dealing with single species. Too little is known to make comprehensive conclusions. The most detailed descriptions can be found in McGuffin (1967: 12), who dealt with Canadian species. Their application to the world-fauna may be questionable, but they provide a good starting point: ‘Egg round, somewhat longer than wide, usually larger at one end than at other (figs 77, 84c) with longitudinal ridges and cross striae; white, cream coloured, or light green; turning pink, scarlet, brown, or black; attached to leaves of food plant or laid loosely on ground near food plant.’ The illustrations of eggs of Palaearctic Scopula species fit the description well (Sannino & Balbiani, 1984; Sannino & Espinosa, 1999). The caterpillar is usually long, slender, green or brown in colour, with longitudinal patterns. The resting posture is either stick-like, at an angle of 45 °, or ventrally bent to form a complete loop (Sugi, 1987; Murase, 1998; Ohbayashi, 2000; Tominaga, 2000; Ebert, 2001). According to McGuffin (1967: 12): ‘20–30 annulets on each anterior abdominal segment; cuticle smooth at 50×; body setae short, with blunt tips; setae on legs relatively long, pointed; prolegs of mature larva with 6–10 crochets in two incompletely separated groups (fig. 82g).’ If disturbed, the larvae of many Palaearctic species drop down and make several powerful side-to-side movements (pers. observ.).

Many species are polyphagous, and recorded host-plants range from woody to herbaceous taxa. They include Scrophulariaceae (Striga), Sterculiaceae (Theobroma), Asteraceae (Lactuca), Polygonaceae (Rumex), Lamiaceae(Thymus), Poaceae (Zea), Moraceae (Ficus) Rosaceae (Filipendula, Sorbus), Liliaceae (Litanthus) and Ericaceae (Vaccinium) for Scopula; Rubiaceae (Adina, Breonia, Mitragyna, Hymenodictyon and Mussaenda) and Marantaceae (Maranta) for the Antitrygodes-group; Rubiaceae (Oxyanthus, Randia) and Sapindaceae (Blighia) for the Aletis-group (Staude, 1999; Tanahara, 1999; Robinson et al., 2002); Lamiaceae (Thymus), Caryophyllaceae (Dianthus, Gypsophila) and Brassicaceae (Alyssum) for the Glossotrophia-group (King, 2000; Hausmann & Dötterl, 2003) and Rubiaceae (Pygmaeothamnus) for the Epicosymbia-group (Staude, 1999). A few species are known to be of minor pest status, attacking tobacco (Sannino & Balbiani, 1984;,Sannino & Espinosa, 1999) and groundnut (Satpathi, 1995).

Pupal cremaster has usually four pairs of setae, terminal pair usually much enlarged (Khotko, 1977; Patocka, 1994), reduced to two in S. indicataria(Nakamura, 1994). In the Glossotrophia-group the proboscis case extends beyond the cremaster, curving inwards (King, 2000; Hausmann, 2001).

Biology:

Scopula includes species of both forested and open habitats as shown in the host-plant list. A number of species have been recorded feeding as adults on the tears, sweat and blood seeping from wounds of large mammals in South-east Asia (Bänziger & Fletcher, 1985). Many species are day-active, and the species of the African Aletis and Cartaletis groups are exclusively diurnal.

Parasitoids:

Homolobus truncator(Say) (Braconidae: Homolobinae) has been reported to parasitize the larva of S. annae(Mentzer) of the Glossotrophia group (King, 2000).

Distinct species groups can be defined within Scopula. Their description is, however, beyond the scope of this paper. They include the Antitrygodes, Epicosymbia, Glossotrophia and Aletis groups.

Comments on systematics:

Scopula is the largest and externally most heterogeneous genus of the tribe. Although only one character was found to support the monophyly of this genus, overall support is nevertheless not as weak as it may appear from the cladogram. Having studied a large number of species across different biogeographical regions, I have seen that the moths of this genus have many characteristic features such as separate sacculus and valvula of the valva, urceolate shaped juxta and spined signum, in addition to diagnostic structures of the male 8th sternite and tergite. All were found to be homoplastic, and accordingly, many are present in the majority of species, although one or a few may be absent in any given species. For example, the absence of ceras in many apical species such as Scopula sentinaria, Stigma kuldschaensis and Dasybela achroa was found to be due to secondary losses (79: 0, CI = 0.09, RI = 0.51). It is thus difficult to define this genus in terms of unique features; instead, Scopula can be characterized by a combination of characters, as already noted by Covell (1970). Many distinctive species groups within Scopula were found, but their relationships remain unresolved, probably due to the high degree of homoplasy that the characters display (Fig. 134).

All the synonymized genera were found to have diagnostic structural characters of Scopula, although some of the taxa were found to have very divergent external appearance from the type species of the genus, S. ornata(Appendix).

ACKNOWLEDGEMENTS

I am greatly indebted to the financial supporters of this study: The Finnish Cultural Foundation, Department of Population Biology, University of Helsinki, Department of Entomology, Finnish Museum of Natural History, Finnish Lepidopterists' Society, and Otto A. Malm's Donationfund, Societas pro Fauna et Flora Fennica, and my wife Karoliina Reunanen after funding from other sources had ceased. A SYS-resource travel grant to BMNH allowed me to study their important collections. The following gave me access to collections under their care and/or loaned material for study: D. Goodger, M. Honey and G. Martin (BMNH), A. Hausmann (ZSBS), M. Horak and T. Rangsi (ANIC). L. Kaila (ZMH) is thanked for supervising the study, H. Sihvonen, for photographing the slide mounts, J. Holloway, for helpful discussions and G. Martin (BMNH) for checking old literature references. B. Landry (Muséum d'histoire naturelle, Genève), J. Hyvönen (University of Helsinki) and two anonymous referees for valuable comments on an earlier version of the manuscript, A. Albrecht for programming the macros and D. Alaruikka (ZMH) for revising my English. The following people are thanked for advice, discussions and support: O. Biström, K. Mikkola, J. Muona, N. Laurenne and S-L. Nyéki (ZMH).

REFERENCES

Appendices

Appendix

PRELIMINARY REVISED CHECKLIST OF THE GENERA AND SPECIES OF THE SCOPULINI

The definition of the tribe Scopulini follows Sihvonen & Kaila (2004). The concept also includes Aletini and Problepsini (Holloway, 1996,,1997). The checklist is largely based on Scoble (1999); this publication is recommended for readers who require further details. Species and subspecies (the latter indented) are listed in alphabetical order under their revised generic names. An asterisk indicates that the species has been studied by the author (external and genital characters). Synonyms are also indented. Comments are printed in a smaller font within square parentheses. While the need for replacement names as a result of primary or secondary homonymy is clear, they have not been proposed here due to the fact that in many cases type specimens were not studied. As not all putatively valid species were included in the study, several monophyletic groups may eventually be recovered that do not appear as such in the list. This is especially true of the Somatina species assemblage, where numerous species await combination with the appropriate genera; instances are mentioned in the comments.

Scopulini

Scopulini Duponchel, 1845 [1844-46](as Scopulites). Type species: Phalaena paludata Linnaeus, 1767, a junior synonym of Phalaena ornata Scopoli, 1763 [Portugal].

Aletini Hampson, 1918 (as Aletinae). Type species: Papilio helcita Linnaeus, 1763. Indiis [incorrect locality, probably tropical Africa].

Problepsini Wiltshire, 1990. Type species: Caloptera ocellata Frivaldszky, 1845. [Greece].

Dithalama

Dithalama Meyrick, 1888. Type species: Dithalama cosmospila Meyrick, 1888. Australia.

cosmospila Meyrick, 1888*

desueta(Warren, 1902)*

Dithalama is possibly a non-monophyletic assemblage of species (see Diagnosis and Isoplenodia below). The following unstudied Dithalama taxa are left tentatively under their current generic combination, waiting for their status to be revised against the diagnosis provided for Dithalama.

persalsa(Warren, 1902)

ioparia(Swinhoe, 1902)

punctilinea Swinhoe, (1902)

tetrasticha(Lower, 1902

Isoplenodia

Isoplenodia Prout, 1932. Type species: Isoplenodia arrogans Prout, 1932. Madagascar.

The generic status of Isoplenodia Prout and especially its relationship to Dithalama warrants further investigation (see Dithalama). It is retained tentatively as a valid genus because females, which often are phylogenetically informative, were not available for study. Isoplenodia may turn out to be a junior synonym of Dithalama.

arrogans Prout, 1932*

Somatina

SomatinaGuenée, [1858]. Type species: Somatina anthophilata Guenée, [1858]. [India].

Nebessa Walker, 1869. Type species: Nebessa chalyboeta Walker, 1869. Congo.

Somatinopsis Warren, 1896. Type species: Somatinopsis nigridiscata Warren, 1896. Indonesia.

Somatodes Guenée, [1858][misspelling of Somatina]

Somatina is possibly a non-monophyletic assemblage of species. The following taxa have been checked and agree with the diagnosis provided.

anthophilata Guenée, [1858]*

chalyboeata(Walker, 1869)*

densifasciaria Inoue, 1992*

[S. densifasciaria Inoue shares the diagnostic conditions of Somatina, for example the posterior margin on the male 8th tergite that is concave with two sharp projections (character 88: 3) (Inoue, 1992). It is externally similar to S. rosacea Swinhoe.]

discata Warren, 1909*

[S. discata Warren, 1909 is possibly a junior synonym of Somatina nigridiscata Warren, 1896, based on examination of the genitalia of the type material from The Natural History Museum, London: Somatina discata holotype (slide BMNH GEO 19799) and Somatinopsis nigridiscata holotype (slide BMNH GEO 19798)].

lia Prout, 1915*

nigridiscata(Warren, 1896)*

[See S. discata Warren, 1909.]

sedata Prout, 1922*

The following taxa are left tentatively within their current generic combination, awaiting comprehensive revision.

accraria Swinhoe, 1904

[Somatina accraria Swinhoe, 1904 should probably be combined with the genus Problepsis. It is for the time being retained in Somatina, however, as I have not yet prepared type specimens of this species, deposited at BMNH. S. accraria is closely related to S. figurata Warren (Prout, 1929−35; Janse, 1933−35); see latter for further comments.]

apicipuncta Prout, 1915

centrofasciaria(Leech, 1897)

[Only a female type specimen of this taxon is known, deposited at BMNH. Prout (1920−41, 1934−39) speculated that it should perhaps be combined with Discoglypha Warren 1896.]

ctenophora Prout, 1915*

[S. ctenophora Prout, 1915 should probably be combined with Problepsis. Janse (1933−35) illustrated the male genitalia, which share several features of Problepsis, e.g. socii fused at apex, lying ventral to the tuba analis. S. ctenophora is for the time being retained in Somatina, however, as I have not yet prepared the type specimen of this species, deposited at BMNH.]

eurymitra Turner, 1926 incerta sedis*

[Somatina eurymitra Turner, 1926 is incertae sedis, but retained in Somatina until a revision is done. It may not be congeneric with Somatina, as my cladistic results indicate that it is the sister taxon to the Dithalama+Isoplenodia clade (Figs 133, 134).]

figurataWarren, 1897*

rufitacta Warren, 1905

ssp. candida Prout, 1932

ssp. transfigurata Prout, 1922

[Somatina figurata Warren, 1897 should probably be combined with Problepsis. Janse (1933−35) illustrates the male genitalia, which share several features of Problepsis, e.g. socii fused at apex, lying ventral to the tuba analis, and distal part of aedeagus that has a distinct sclerotized spine (character 122: 1). S. figurata is for the time being retained in Somatina, however, as I have not yet prepared the type specimen of this species, deposited at BMNH.]

fletcheri Herbulot, 1958

fraus Prout, 1916

fungifera Warren, 1909

hombergi Herbulot, 1967

impunctulata(Warren, 1901)

[Only the male type specimen of this taxon is known, deposited at BMNH. Prout (1929−35) speculated that it should perhaps be combined with Scopula Schrank, 1802.]

ioscia Prout, 1932

irregularis(Warren, 1898) incerta sedis*

[Cosymbia? irregularis(Warren, 1898) is retained in Somatina awaiting a revision of the genus. I have found that it shares two unique synapomorphies with Scopula haemaleata(Warren, 1898). It is unknown whether this clade represents a larger species group. The synapomorphies may prove to be homoplastic if more taxa are included in a cladistic analysis.]

lemairei Herbulot, 1978

macroanthophilata Xue, 1992

maeandrata Prout, 1925

mozambica(Thierry-Mieg, 1905)

[Somatina mozambica(Thierry-Mieg), which was originally combined with Problepsis but later transferred to Somatina, based on its two-areole forewing (Prout, 1929−35), may eventually be recombined with Problepsis. I have shown this character to be homoplastic and the genus Problepsis, among others, has one or two areoles. Characters mentioned in the description of the species, e.g. weakly developed ocellated discal spots that have metallic (= iridescent) scales suggest affinity with Problepsis(Thierry-Mieg, 1905). I have not dissected the genitalia, and the depository of the type specimen is unknown.]

obscuriciliata Wehrli, 1924

omicraria(Fabricius, 1798)

[Somatina omicraria(Fabricius) may eventually be combined with Problepsis. Characters such as weakly developed ocellated discal spots that have silvery (= iridescent) scales and forewing venation (Prout, 1920−41) suggest affinity with Problepsis. I have not dissected the genitalia; type specimen is at ZMUC.]

cana Hampson, 1895

extrusata(Walker, 1861)

ossicolor Warren, 1898

plynusaria(Walker, [1863])

congruaria(Walker, 1869)

postlineata(Warren, 1899)

[Prout (1920−41) speculated that this taxon should perhaps be combined with Scopula Schrank, 1802. I have not dissected the genitalia; type specimens are at BMNH.]

probleptica Prout, 1917

[Somatina probleptica Prout may eventually be combined with Problepsis. Characters such as well developed ocellated discal spots that have silvery (= iridescent) scales and forewing venation suggest affinity with Problepsis, as already indicated by Prout (1929–35). I have not dissected the genitalia; type specimen is at BMNH.]

prouti Janse, 1934*

[Somatina prouti Janse, 1934 should probably be combined with Problepsis. Janse (1933–35) illustrated the male genitalia, which shares features of Problepsis, in addition to wings that have weakly ocellated discal spots with iridescent scales not seen in Somatina. Somatina prouti has two areoles on the forewing and this has been considered a Somatina feature. My cladistic analysis shows that this character is homoplastic and that Problepsis, among others, has one or two areoles. Female is unknown. Somatina prouti is for the time being retained in Somatina, however, awaiting a comprehensive revision.]

purpurascens Moore, [1887]

pythiaria(Guenée, [1858])

[Argyris pythiaria Guenée may eventually be combined with Problepsis. Characters illustrated and mentioned in the description of species, for example weakly developed ocellated discal spots that have lead metallic (= iridescent) scales suggest affinity with Problepsis(Guenée, 1858). I have not dissected the genitalia; lectotype female is deposited at MNHN.]

rhodochila Prout, 1932

rosacea Swinhoe, 1894*

ssp. anaemica Prout, 1914

[Somatina rosacea Swinhoe has several features, including genitalia, that it shares with either Somatina or Scopula, thus making its generic combination difficult to judge. It also demonstrates that the generic association of Somatina species needs to be studied further.]

rufifascia Warren, 1896

maculata Warren, 1898

rubridisca Swhinhoe, 1900

sordida Warren, 1898

sanctithomae Herbulot, 1958

sublucens(Warren, 1907)*

[Only the female of S. sublucens(Warren) was available for study (BMNH 20648). As features like spinose signum show affinity with Scopula, the generic combination of this species remains to be tested.]

subviridata(Warren, 1901)

syneorus Prout, 1915

transvehens Prout, 1918

[Somatina transvehens Prout may eventually be combined with Problepsis. Characters such as weakly developed ocellated discal spots that have silvery (= iridescent) scales and forewing venation suggest affinity with Problepsis, as already indicated by Prout (1920–41). I have not dissected the genitalia; type specimen is in BMNH.]

vestalis(Butler, 1875)*

[Somatina vestalis(Butler, 1875) should probably be combined with Problepsis. Janse (1933–35) illustrates the male genitalia, which share several features of Problepsis, e.g. socii fused at apex, lying ventral to the tuba analis. In addition, the wings have weakly ocellated discal spots with iridescent scales not seen in Somatina. S. vestalis has two areoles on the forewing and this has been considered a feature of the genus. My cladistic analysis shows that this character is homoplastic and that Problepsis, among others, has one or two areoles. Female genitalia (slide BMNH GEO 4274) also suggest affinity with Problepsis. S. vestalis is for the time being retained in Somatina, however, as I have not yet prepared the type specimen of this species, deposited at BMNH.]

virginalis Prout, 1917*

[Somatina virginalis Prout, 1917 should probably be combined with Problepsis. Wings have weakly ocellated discal spots with iridescent scales not seen in Somatina. S. virginalis has two areoles on its forewing and this has been considered a feature of the genus. My cladistic analysis shows that this character is homoplastic and that Problepsis, among others, has one or two areoles. Female genitalia (slide BMNH GEO 4275) also suggest affinity with Problepsis. Somatina virginalis is for the time being retained in Somatina, however, as I have not yet prepared type specimen of this species, deposited at BMNH.]

wiltshirei Prout, 1938

Zythos

Zythos Fletcher, 1979. Type species: Nobilia turbata Walker, 1862. Malaysia. [Replacement name for Nobilia Walker, 1861].

Nobilia Walker, 1861. Type species: Nobilia turbata Walker, 1862. Malaysia. [Junior homonym of Nobilia Gray, 1855 (Mollusca).]

aphrodite(Prout, 1932)*

ssp. rooki(Prout, 1938)

avellanea(Prout, 1932)*

clypeata Yazaki, 1996*

cupreata(Pagenstecher, 1888)

nebulosa(Warren, 1897)

erotica(Prout, 1932)*

fastigata(Prout, 1938)*

modesta Yazaki, 1996*

molybdina(Prout, 1938)

ssp. tombarensis(Prout, 1938)

obliterata(Warren, 1897)*

simplaria(Snellen, 1899)

strigata(Warren, 1896)*

ssp. rubescens(Prout, 1938)

turbata(Walker, 1862)*

strigularia(Snellen, [1886])

Lipomelia

Lipomelia Warren, 1893. Type species: Lipomelia subusta Warren, 1893. India.

Defoa Swinhoe, 1893. Type species: Defoa ustata, a junior synonym of Lipomelia subusta Warren, 1893. [India].

subusta Warren, 1893*

ustata(Swinhoe, 1893)

Problepsis

Problepsis Lederer, 1853. Type species: Caloptera ocellata Frivaldszky, 1845. [Greece]. [Replacement name for Caloptera Frivaldszky, 1845.]

Argyris Guenée, [1858]. Type species: Argyris ommatophoraria Guenée, [1858], a junior synonym of Problepsis ocellata Frivaldszky, 1845. [Lebanon.]

Caloptera Frivaldszky, 1845. Type species: Caloptera ocellata Frivaldszky, 1845. [Greece]. [Junior homonym of Caloptera Gistl, 1834 (Coleoptera).]

Problepsiodes Warren, 1899. Type species: Problepsis conjunctiva Warren, 1893. India.

achlyobathra Prout, 1928*

ssp. emphyla Prout, 1938*

ssp. violescens Prout, 1934

aegretta Felder & Rogenhofer, 1875*

egretta Hale-Carpenter, 1932 [misspelling of aegretta]

ssp. insculpta Prout, 1917

albidior Warren, 1899*

ssp. matsumurai Prout, 1938

apollinaria(Guenée, [1858])*

ssp. aphylacta Prout, 1938

ssp. candidior Prout, 1917

ssp. deparcata Prout, 1925

ssp. hemicyclata Warren, 1897

ssp. wilemani West, 1930

argentea Warren, 1900

asira Wiltshire, 1982*

borneamagna Holloway, 1997*

centrophora(Prout, 1915) comb. nov.*

[Somatina centrophora Prout, 1915 is transferred from Somatina to Problepsis.]

clemens Lucas, 1890*

margaritata Warren, 1896

conjunctiva Warren, 1893*

ssp. subjunctiva Warren, 1893

craspediata Warren, 1897

ssp. frosti Prout, 1938

ssp. rotifera Prout, 1916

crassinotata Prout, 1917*

deducta Herbulot, 1962

deliaria Guenée, [1858]*

delphiaria Guenée, ([1858])*

ssp. auriculifera Warren, 1897

argentisquama(Warren, 1899)*

diazoma Prout, 1938

digammata Kirby, 1896*

digammata Warren, 1897

[Junior primary homonym and a junior synonym of P. digammata Kirby, 1896. P. digammata Warren, 1897 was erroneously listed as a junior synonym of P. aegretta aegretta Felder & Rogenhofer, 1875 (Vári, Kroon & Krüger, 2002).]

digrammata Prout, 1933 [misspelling of digammata]

discophora Fixsen, 1887*

superans coreana Bryk, 1949

ssp. kardakoffi Prout, 1938*

erythra Wiltshire, 1982*

eucircota Prout, 1913*

evanida Prout, 1932

exanimata Prout, 1935

flavistigma Swinhoe, 1904*

ssp. dilatistigma Prout, 1917

herbuloti Viette, 1968

insignita Prout, 1938

korinchiana Rothschild, 1920*

latonaria Guenée, ([1858])*

longipannis Prout, 1917

lucifimbria(Warren, 1902)

magna Warren, 1906*

maxima Thierry-Mieg, 1905

meroearia Saalmüller, 1884*

mayottaria(Oberthür, 1923)

metallopictata(Pagenstecher, 1888)

venus Thierry-Mieg, 1905

minuta Inoue, 1958*

mitis Joannis, 1932

neumanni Prout, 1932

ocellata(Frivaldszky, 1845)*

ommatophoraria Guenée, ([1858])

ssp. cinerea(Butler, 1886)*

ochripicta Warren, 1901

paredra Prout, 1917*

phoebearia Erschov, 1870*

[Replacement name for Argyris deliaria Bremer, 1864.]

deliaria(Bremer, 1864)

[Junior primary homonym of Argyris deliaria Guenée, [1858].]

plagiata(Butler, 1881)*

riminota Prout, 1938

plenorbis Prout, 1917*

rorida Prout, 1932*

sancta Meyrick, 1888

shirozui Inoue, 1986*

similinotata Prout, 1917*

subreferta Prout, 1935

superans Butler, 1885*

ssp. summa Prout, 1935

transposita Warren, 1903

triocellata Bastelberger, 1908. comb. rev.*

[Problepsis triocellata(Bastelberger, 1908) is transferred from Somatina to Problepsis.]

ssp. scenica(Prout, 1938)

vulgaris Butler, 1889*

attenuata Warren, 1909

Scopula

Scopula Schrank, 1802. Type species: Phalaena paludata Linnaeus, 1767, a junior synonym of Phalaena ornata Scopoli, 1763. [Portugal].

Acidalia Bruand, 1846. Type species: Geometra strigaria Hübner, [1799], a junior synonym of Geometra virgulata[Denis & Schiffermüller], 1775. [Austria]. [Junior homonym of Acidalia Hübner, [1819] 1816 (Lepidoptera: Nymphalidae).]

Acidalia Treitschke, 1825. Type species: Geometra strigaria Hübner, [1799], a junior synonym of Geometra virgulata[Denis & Schiffermüller], 1775. [Austria]. [Junior homonym of Acidalia Hübner, [1819] 1816 (Lepidoptera: Nymphalidae).]

Acidalina Staudinger, 1898. Type species: Acidalina decolor, Staudinger, 1898. Algeria.

Aletis Hübner, [1820]. Type species: Papilio helcita Linnaeus, 1763. Indiis [incorrect locality, probably tropical Africa]. syn. nov.

Anacosymbia Prout, 1913. Type species: Anacosymbia perstrigulata Prout, 1913. [South Africa]syn. nov.

Antanepsia Turner, 1908. Type species: Autanepsia poliodesma Turner, 1908. Australia [missspelling of Autanepsia]

Antilycauges Prout, 1913. Type species: Emmiltis pinguis Swinhoe, 1902. [Taiwan]. syn. nov.

Antitrygodes Warren, 1895. Type species: Macaria divisaria Walker, 1861. [India]. syn. nov.

Autanepsia Turner, 1908. Type species: Autanepsia poliodesma Turner, 1908. Australia. syn. nov.

Calothysanis Hübner, [1823] 1816. Type species: Geometra imitaria Hübner, [1799] 1796. Europe.

Cartaletis Warren, 1894. Type species: Aletis libyssa Hopffer, 1858. Mozambique. syn. nov.

Chlorocraspedia Warren, 1899. Type species: Chlorocraspedia ansorgei Warren, 1899. Uganda.

Cinglis Guenée, [1858]. Type species: Fidonia humifusaria Eversmann, 1837. [Russia]. syn. nov.

Craspedia Hübner, [1825] 1816: Type species: Phalaena ornata Scopoli, 1763. [Italy].

Csopula Fischer von Röslerstamm, 1842. [misspelling of Scopula].

Cymatida Sodoffsky, 1837. Type species: Geometra strigaria Hübner, [1799] 1796, a junior synonym of Geometra virgulata[Denis & Schiffermüller], 1775. [Austria]. [Unnecessary replacement name for Acidalia Treitschke, 1825].

Cymatoides Agassiz, 1847. [Emendation of Cymatida Sodoffsky, 1837].

Dasybela Turner, 1908. Type species: Emmiltis achroa Lower, 1902. Australia. syn. nov.

Dasybella Turner, 1908 [misspelling of Dasybela]

Dosithea Duponchel, 1829. Type species: Phalaena ornata Scopoli, 1763. [Portugal].

Dosithoea Rambur, 1833 [misspelling of Dosithea]

Epicosymbia Warren, 1897. Type species: Epicosymbia perrufa Warren, 1897, a junior synonym of Acidalia? dentisignata Walker, [1863]. [South Africa]. syn. nov.

Eucidalia Sterneck, 1941. Type species: Phalaena immorata Linnaeus, 1758. Europe.

Glossotrophia Prout, 1913. Type species: Acidalia confinaria Herrich-Schäffer, 1847. Hungary. syn. nov.

Holarctias Prout, 1913. Type species: Haematopis sentinaria Geyer, 1837. [Canada].

Ignobilia Prout, 1932. Type species: Ephyra urnaria Guenée, [1858]. Borneo. syn. nov.

Induna Warren, 1897. Type species: Induna rufisalsa Warren, 1897. [South Africa].

Isoplenia Warren, 1897. Type species: Isoplenia trisinuata Warren, 1897. [South Africa]. syn. nov.

Leucoxena Warren, 1900. Type species: Leucoxena lactea Warren, 1900. [Kenya]. syn. nov.

Leptaletis Warren, 1894. Type species: Aletis variabilis Butler, 1878. [Angola]. syn. nov.

Leptomeris Hübner, [1825] 1816. Type species: Geometra umbelaria Hübner, [1813] 1796. Europe.

Lipocentris Warren, 1905. Type species: Lipocentris rubriceps Warren, 1905. Angola.

Longula Staudinger, 1892. Type species: Longula extraordinaria Staudinger, 1892, a junior synonym of Lycauges donovani Distant, 1892. [Lebanon].

Lycauges Butler, 1879. Type species: Lycauges lactea Butler, 1879, a junior synonym of Acidalia emissaria Walker, 1861. Japan.

Oar Prout, 1913. Type species: Phalaena pratana Fabricius, 1794. [North Africa]. syn. nov.

Phyletis Guenée, [1858]. Type species: Phyletis silonaria Guenée, [1858]. [Ethiopia].

Pigia Guenée, [1858]. Type species: Pigia infantularia Guenée, [1858], a junior synonym of Phalaena opicata Fabricius, 1798. [Sri Lanka].

Pleionocentra Warren, 1898. Type species: Scopula plionocentra Prout, 1920. [Geometra minorata sensu Warren, nec Boisduval 1833]. Niger.

Prasonesis Meyrick, 1889. Type species: Prasonesis microphylla Meyrick, 1889. [Papua New Guinea]. syn. nov.

Pseudocinglis Hausmann, 1994. Type species: Glossotrophia eurata Prout, 1913. Turkmenistan. syn. nov.

Psilephyra Bastelberger, 1909. Type species: Psilephyra bilineata Bastelberger, 1909, a junior synonym of Induna curvimargo Warren, 1900. [Tanzania].

Pylarge Herrich-Schäffer, 1855. Type species: Idaea commutata Freyer, [1832] 1833, a junior synonym of Scopula ternata, Schrank, 1802. [Germany].

Runeca Moore, 1888. Type species: Runeca ferrilineata Moore, 1888. India.