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1 August 2002 A Preliminary Study on Relationships among Selected Australian Members of the Tribe Spilomelini (Lepidoptera: Crambidae: Pyraustinae)
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Abstract

A preliminary study was conducted on phylogenetic relationships among some selected genera of the Australian Spilomelini, focusing on relationships among the Australian Glyphodes group (Glyphodes Guenée, 1854, Dysallacta Lederer, 1863, Talanga Moore, 1885 and Agrioglypta Meyrick, 1932) and the 17 genera which are morphologically similar to it. Representatives of three genera of the Pyarustini were used as outgroups. Cladistic analysis of morphological data from the adult moths produced 10 equally MP trees (length = 221, CI=0.294, and RI=0.608). The clade formed by the 21 selected genera of the Australian Spilomelini had low bootstrap support even though a good apomorphy supported the monophyly of this group, namely, a strong, bilobed praecinctorium of abdominal tympanal organs. The analysis showed that the Glyphodes group is not monophyletic because the genus Chrysothyridia Snellen appears within it in the 10 MP trees. The concept of the Glyphodes group should be expanded to include Chrysothyridia and also the Synclera Zeller and Didymostoma (Walker) since the Synclera Didymostoma clade, as the hypothesised sister group of the Glyphodes group, is not sufficiently supported as a separate monophyletic group. The analysis also showed that genus Glyphodes is not a monophyletic group, while Metallarcha Meyrick is a monophyletic group.

INTRODUCTION

The Spilomelini was erected by Guenée(1854) as the family Spilomelidae, based on the type genus Spilomela Guenée. Munroe (1976) treated the group at tribal level and put it under the subfamily Pyraustinae. This concept was followed by Common (1990), Scoble (1992), Shaffer et al. (1996) and Munroe & Solis (1999).

As for most other crambid genera, the genera of the Australian Spilomelini are poorly defined and the relationships among them have never been examined. The current classification of the Australian Spilomelini is based on external similarity only, which has proved to be taxonomically unreliable for crambids in comparison to genitalic structures, and the monophyly of each genus of the Spilomelini needs to be examined phylogenetically.

The Spilomelini is a large tribe, with more than half of the species of pyraustines belonging to it. In Australia, there are about 360 species represented in about 125 genera (Shaffer et al. 1996). Therefore, only some genera of the Spilomelini were chosen and included in this study.

The aim of this preliminary study was to reconstruct the phylogenetic relationships among some selected genera of the Australian Spilomelini with focus on the relationships among 17 genera which are similar morphologically to the Australian Glyphodes group. The four genera, Glyphodes, Talanga, Dysallacta and Agrioglypta were defined as the Glyphodes group by Hampson (1896) based on external features such as palpi upturned, the 2nd segment broadly scaled in front, the 3rd segment porrect, maxillary palpi triangularly dilated with scales, antennae of male near simple, tibia with the outer spurs less than half the length of inner, and male with the anal tuft large. A cladistic study was undertaken using morphological characters of the adult moths, including many features from the male and female genitalia.

MATERIALS AND METHODS

Materials

Most specimens used in this study belong to the Australian National Insect Collection (ANIC), CSIRO Entomology, Canberra, or were borrowed from the National History Museum, London (BMNH). Both the outgroup and ingroup taxa included in the analysis are listed in Appendix 1.

Preparation of slide-mounts

Head structure preparation was made of each species used in the analysis. The labial palpi, the maxillary palpi and the antennae were dissected and mounted into euparal. The wing preparation was made following Common (1990) whereas the genitalic preparation following Robinson and (1976) and Maes (1985).

Morphological terms

The morphological terms used for adult features are as given in Zimmerman (1958), Munroe (1976), and Maes (1995). Definitions of certain terms which have been modified are given below:

The terms “strongly sclerotized” and “weakly sclerotized” refer to degree of sclerotization of various structures. A strongly sclerotized structure is darkened and totally opaque, and weakly sclerotized one is yellow brown, readily visible but translucent.

Choice of Outgroup

The most appropriate outgroup taxa for this analysis were genera from the sister group of the Spilomelini, which is the Pyraustini (Common, 1990; Munroe, 1976). Pyrausta testalis (Fabricius) was chosen as the prime outgroup (= Pyrausta 1), and P. panopealis (Walker) (=Pyrausta 2), Isocentris filalis Guenée and Hyalobathra archeleuca Meyrick were used as the comparison outgroups.

Choice of Ingroup

In this study, the ingroup taxa was chosen for those genera which share certain morphological characters with the Glyphodes group (Hampson 1896), such as shape of head, structures of maxillary and labial palpi, wing venation and abdomen features. I assumed that they were closely-related genera even though this criteria was assessed only based on an external examination which does not guarantee a close relationship. All the selected taxa are the type species of the genus concerned, unless otherwise stated.

In this analysis, I used three species of Glyphodes to represent the range of variation of the genus. The selected species were G. stolalis Guenée (the type species of the genus, referred to as Glyphodes 1), G. bicolor (Swainson) (referred to as Glyphodes 2) and G. bivitralis Guenée (referred to as Glyphodes 3). I used two species to represent Metallarcha because it is a large genus for which the current classification is based on external characters only (Shaffer et al., 1996). Metallarcha was represented by M. eurychrysa Meyrick (the type species of the genus, referred to as Metallarcha 1) and M. diplochrysa Meyrick (referred to as Metallarcha 2) in the subsequent analysis. All other genera were represented by just their type species.

Characters used

A total of 42 characters from the head, thorax, abdomen and genitalia structure were chosen. All characters were treated as ordered (Wagner Parsimony) with the exception of character 16 which was treated as unordered. The data matrix for morphological characters used in the analysis is presented in Appendix 2.

Head structure:

  1. Shape of frons of head: (0) rounded; (1) flat. The character state of rounded is plesiomorphic condition.

  2. Antennal ciliation on male: (0) normal; (1) reduced. The character state of normal (0) is plesiomorphic condition.

  3. Antennal ciliation of male and female: (0) of equal length; (1) of different length. The character state of equal length is plesiomorphic condition.

  4. Labial palpus: (0) ascending or upturned; (1) forward directed. I assumed that the character state of ascending or upturned is plesiomorphy.

  5. Third segment of labial palpus: (0) ovate or rounded; (1) cylindrical or elongate. I assumed that the character state of ovate or rounded (0) is plesiomorphy.

  6. Length of second segment of labial palpus: (0) very long, more than 2x length of first segment; (1) short, longer than length of first segment but less than 2x length of first segment; (2) very short, less than length of first segment. I assumed that the character state of very long more than length of first segment (0) is plesiomorphy.

  7. Length of last segment of maxillary palpus: (0) longer than length of 3rd segment; (1) as long as or shorter than length of 3rd segment. The character state of longer than length of 3rd segment is plesiomorphic condition.

    Thorax:

  8. Base of costa of forewing of male: (0) a strap-like frenulum hook present (Fig. 2.C); (1) a strap-like frenulum hook absent. This structure is very difficult to observe on the specimens and very often is lost during slide preparation. The best way to observe it is by removing the scales at the base of the costa with a fine brush. I assumed that the character state of a strap-like frenulum hook absent (1) is apomorphy for Spilomelini.

  9. Frenulum of female: (0) >2 bristles; (1) only 1 bristle. I assumed that the character state of >2 bristles (0) is plesiomorphy.

  10. Veins 1A+2A of forewing of male: (0) forming a loop (Figs. 2.C, 2.E); (1) not forming a loop (Figs. 2.A, 2.D); If vein 1A+2A are very slightly stained but form a loop, even if is not complete, they were scored as state 1. I assumed that the character state of not forming a loop (1) is apomorphy for Spilomelini.

  11. Wing venation of forewing of male: (0) R 2 distinctly separate from R 3+R 4 (Fig. 2.D); (1) R 2 close at basal of with R 3+R 4 (Fig. 2C); (2) R 2 fused to middle of R 3+R 4 (Fig. 2.D). The characters state of R 2 distinctly separate from R 3+R 4 is plesiomorphy.

  12. Base of R 5 of male's forewing: (0) from discal cell (Figs. 2.C, 2.D); (1) stalked with R 3+R 4 (Fig. 2.A). The character state from discal cell (0) is plesiomorphic condition.

  13. Wing pattern of forewing on both sex: (0) median band normal, narrow; (1) median band wide; (2) median band absent, transparent dots present; (3) no median band, no transparent dots but other patterns. I assumed that the character state of median band normal, narrow (0) is plesiomorphic condition.

    Abdomen:

  14. Praecinctorium of tympanum: (0) weakly bilobed (Fig. 3.B); (1) strongly bilobed (Fig. 3.A). The character state of strongly bilobed (1) is apomorphic condition for Spilomelini.

  15. Hair pencil of S8: (0) absent; (1) present. The character state of present (1) is apomorphic condition.

  16. T8 sclerite (inverted): (0) not forming inverted Y-shaped sclerite (Fig. 4.A); (1) roughly inverted Y-shaped sclerite, lateral margin parallel only in posterior 1/3 (Fig. 4.B); (2) clearly inverted Y-shaped sclerite and parallel in posterior 2/3 (Fig. 4.C). This character was treated as unordered.

  17. Shape of 8th abdominal segment in male: (0) abdomen normal; (1) very long. The character state of abdomen normal (0) is plesiomorphic condition.

  18. Scaling of 8th abdominal segment in male: (0) dense bundle of scales present; (1) without dense bundle of scales. The character state of dense bundle of scales present is plesiomorphy.

    Male genitalia:

  19. Length of aedeagus: (0) short, less than length of abdomen (Figs. 5.A-5.D); (1) very long, narrow, more than length of abdomen (Figs. 8.B, 8.F). The character state of very long, narrow, more than length of abdomen is apomorphic condition.

  20. Cornuti: (0) present (Fig. 5.B); (1) absent. The character state of cornuti present (0) is plesiomorphic condition.

  21. Coecum of aedeagus: (0) absent; (1) short (Fig. 5.A); (2) long (Fig. 5.B); (3) modified (Fig. 5.C). The character state of absent (0) is plesiomorphic condition.

  22. Sclerotization ventrally along aedeagus: (0) absent; (1) present (Fig. 5.A). The character state of absent (0) is plesiomorphic condition.

  23. Uncus: (0) present with a long “neck” (Figs. 6.A, 6.B); (1) present with a short “neck” (Fig. 6. D); (2) reduced (Fig. 6.F). The character state of present with long “neck” (0) is plesiomorphic condition.

  24. Apex of uncus: (0) with strong, short spines (Fig. 6.D); (1) with hair (Fig. 6.B); (2) without spines or hair (Fig. 6.A). The character state of with strong, short spines (0) is plesiomorphic condition.

  25. Base of uncus: (0) united to tegument (Figs. 6.D, 6.E); (1) split from tegument (Fig. 6.A–6.C); For taxa without uncus this character was scored as ? (inapplicable). The character state of split from tegumen (0) is plesiomorphic condition.

  26. Sclerotization of base of tuba analis: (0) present, normal sclerotized (Fig. 6.B); (1) absent; (0) present, weakly sclerotized. The character state of present, normal sclerotized (0) is plesiomorphic condition.

  27. Sclerotization of transtilla: (0) normal, sclerotized; (1) strongly sclerotized (Fig. 6.B); (2) not sclerotized. The character state of normal sclerotized (0) is plesiomorphic condition.

  28. Costa: (0) inflated (Fig. 7.B); (1) not inflated. The character state inflated (0) is plesiomorphic condition.

  29. Valva: (0) no fibula but editum or sella present (Fig. 7.C); (1) fibula present, normal (Figs. 7.D); (2) fibula with tip recurved (Fig. 7.B); (3) more than 1 fibula present (Fig. 7.E); (4) without any processes (Fig. 7.A). The character state of no fibula but editum or sella present is plesiomorphy.

  30. Valva: (0) without any sclerotization along middle (Figs. 7.A-7.C); (1) with sclerotization along middle (Fig. 7.D). The character state of without any sclerotization along middle (0) is plesiomorphic condition.

  31. Ventral angle of vinculum: (0) simple, strongly curved, ovate, triangular, ventrally not expanded (Fig. 5.A); (1) ventral portion of vinculum not expanded, hardly curved (Fig. 5.F); (2) ventral angle with keel-shaped sclerotized extension (Fig. 5.H); (3) ventral angle with sclerotized extension not keel-shaped but invaginated (Fig. 5.G). The character state of simple, strongly curved, ovate, triangular, ventrally not expanded (0) is plesiomorphic condition.

  32. Base of coremata: (0) absent; (1) present but simple; (2) complex structures without extension (Fig. 8.C); (3) complex structure with a lamellate extension (Fig. 8.E). The character state of absent (0) is plesiomorphic condition.

  33. Shape of coremata scales: (0) long, hair-like smooth (Fig. 8.A); (1) long, large, paddle-shaped scales; (2) long, hair-like with sculptured surface (Fig. 8.C); (3) strong widened, lamellate scales (Fig. 8.E). For taxa without base of coremata this character was scored as ? (inapplicable). The character state of long, hair-like smooth (0) is plesiomorphic condition.

    Female genitalia:

  34. Corpus bursae: (0) with a smooth membranous wall (Figs. 9.A, 9.B); (1) with spines or sclerotization on wall (Figs. 9.C, 9.D). The charter state of with a smooth membranous wall (0) is plesiomorphic condition.

  35. Signum: (0) present, only one, rhomboidal-shaped (Fig. 9.A); (1) present, only one, not rhomboidal-shaped (Fig. 9.E); (2) present, a pair (Fig. 9.B); (3) absent. The bipartite signum of G. bicolor was treated as state 2. The character state of present one, romboidal-shaped (0) is plesiomorphic condition.

  36. Length of ductus bursae: (0) very short, antrum attached directly to corpus bursae (Fig. 9.D); (1) long, antrum separated from corpus bursae by distinct ductus bursae (10.A–10.C). The character state very short, antrum attached directly to corpus bursae (0) is plesiomorphic condition.

  37. Accessory bursae: (0) absent; (1) present (Fig. 9.D). The character state of accessory bursae absent (0) is plesiomorpic condition.

  38. Ductus seminalis: (0) originating from corpus bursae (Figs. A, 9.C, 9.D); (1) originating from ductus bursae (Figs. 10.A-10.D). The character state of originating from corpus bursae (0) is plesiomorphic condition.

  39. Antrum: (0) absent; (1) reduce to weakly sclerotized; (2) well-sclerotized (Fig. 9.D). The character state of absent is plesiomorphy.

  40. Sclerotization of anterior most part of ductus bursae, posterior to antrum: (0) present (Fig. 10.A); (1) absent. The character state of present (0) is plesiomorphic condition.

  41. Sclerotization of lamella postvaginalis: (0) present (Fig. 10.A); (1) absent. I assumed that the character state of present (0) is plesiomorphic condition.

  42. Length of anterior apophyses: (0) longer than posterior apophyses; (1) shorter than posterior apophyses. The character state of longer than posterior apophyses (0) is plesiomorphic condition.

Fig. 1

A strict consensus of the 10 MP trees, numbers on the node are bootstrap values (from 100 bootstrap replicates × 10 random addition sequences).

i0289-0003-19-8-915-f01.tif

Fig. 2

Wing venation of male: A. Forewing of Aboetheta pteridonoma; B. Hindwing of A. pteridonoma; C. Forewing of Pyrausta panopealis; D. Forewing of Tatobotys janapalis; E. Forewing of Niphograpta albiguttalis. slfr.-strap-like frenulum hook; fr.-frenulum.

i0289-0003-19-8-915-f02.tif

Fig. 3

Praecinctorium of tympanum: A. Glyphodes bivitralis; B. Pyrausta panopealis. prae.-praecinctorium.

i0289-0003-19-8-915-f03.tif

Fig. 4

T 8 of male: A. Pyrausta testalis; B. Talanga sexpunctalis, C. Glyphodes stolalis.

i0289-0003-19-8-915-f04.tif

Fig. 5

A–D, Aedeagi and E–H Vinculum of male genitalia: A. Glyphodes stolalis; B. Diasemiopsis ramburialis; C. Diplopseustis perieresalis; D. Tatabotys janapalis; E. Aboetheta pteridonoma; F. Hyalobathra archeleuca; G. Glyphodes stolalis; H. Hymenia perspectalis. cr.-cornuti; coe.-coecum; vnc.-vinculum.

i0289-0003-19-8-915-f05.tif

Fig. 6

Uncus and tegument of male genitalia: A. Hyalobathra archeleuca; B. Glyphodes stolalis; C. G. bivitralis; D. Hymenoptychis sordida; E. Synclera traducalis; F. Sameodes cancelalis. trs.-transtilla; tgm.-tegumen; unc.-uncus.

i0289-0003-19-8-915-f06.tif

Fig. 7

Valvae of male genitalia (ventral view): A. Myrmidonistis hoplora; B. Sameodes cancellalis; C. Pyrausta panopealis; D. Glyphodes bivitralis; E. Hymenoptychis sordida. cst.-costa; edt.-editum; fbl.-fibula.

i0289-0003-19-8-915-f07.tif

Fig. 8

Male genitalia: valvae and aedagi: A, B. Glyphodes bivitralis; C, D.G. stolalis; E, F. Talanga sexpunctalis. crm.-coremata; fbl.-fibula; trs.-transtila; unc.-uncus; vnc.-vinculum.

i0289-0003-19-8-915-f08.tif

Fig. 9

Corpus bursae: A. Hyalobathra archeleuca; B. Talanga sexpunctalis; C. Spoladea recurvalis; D. Otiophora leucura; E. Metallarcha eurychrysa. ac.br.-accessory bursae; ant.-antrum; db.-ductus bursae; sg.-signum.

i0289-0003-19-8-915-f09.tif

Fig. 10

Female genitalia: A. Glyphodes bicolor; B. Glyphodes stolalis; C Agrioglypta excelsalis; D. Talanga sexpunctalis. lm.pv.-lamella post vaginalis; ap.-apophysis.

i0289-0003-19-8-915-f10.tif

Phylogenetic Inference

The cladistic analysis was undertaken using PAUP*4.0 beta version (Swofford 1998) on a Macintosh computer, and involved 42 characters and 28 taxa. All searches of the data matrix were performed with the Heuristic option. Random addition with 10 replicates was used to generate initial trees for TBR branch-swapping routine.

The MP trees that resulted from this analysis were summarized in a strict consensus tree. Bootstrap tests (Felsenstein 1985) and T-PTP (Topology-dependent cladistic permutation Tail Probability) tests (Faith 1991) were applied to estimate support for clades in the tree. The MacClade version 3 (Maddison & Maddison 1992) was used to evaluate the changes of character states.

RESULTS

A heuristic search in PAUP with all taxa included produced 10 equally MP trees (length=221, CI=0.294, and RI=0.608). The genus Chrysothyridia consistently appears within the Glyphodes group.

A strict consensus tree, with bootstrap support indicated, is presented in Fig. 1. Only a few nodes have bootstrap support. Support for the monophyly of the 24 taxa of the Spilomelini is very low. The monophyly of the group ((Didymostoma + Synclera) + (Glyphodes 2 + (Chrysothyridia + (Glyphodes 1 + (Glyphodes 3 + (Dysallacta + (Agrioglypta + Talanga))))))) and the group (Glyphodes 1 + (Glyphodes 3 + (Dysallacta + (Agrioglypta + (Talanga))))) also are weakly supported.

A T-PTP test was conducted only for the monophyly of the group formed by Didymostoma + Synclera + Glyphodes 2 + Chrysothyridia + Glyphodes 1 + Glyphodes 3 + Dysallacta + Agrioglypta + Talanga. There were significant support for the monophyly for the group, with P (normal) 0.01 and P (reverse) 1.0.

The list of apomorphies demonstrating the monophyly of the groups is presented here only for those groups that comprise more than two taxa and which have bootstrap support. Only the apomorphies that are regarded as unambiguous or least controversial are listed for each monophyletic group considered.

The monophyly of the 24 taxa of the Spilomelini is supported by the following possible apomorphies:

  1. A strong, bilobed praecinctorium (a median extension of the intersegmental thoracho-abdominal membrane) of abdominal tympanal organs. This state is assumed to be the ancestral condition for the Spilomelini following Munroe (1976) and Minet (1981, 1991).

  2. Base of costa of male's forewing without a strap-like frenulum hook. It is possible that the loss of a strap-like frenulum hook has occurred several times in other sub-families of the Crambidae.

The possible apomorphies of the group ((Didymostoma + Synclera) + (Glyphodes 1 + (Glyphodes 2 + (Glyphodes 3 + (Chrysothyridia + (Dysallacta + Agrioglypta + (Talanga))))))) are:

  1. Signum paired. This state is assumed to be apomorphic condition of this group. All the members of this group have paired signa except Glyphodes 1 where the signum is reduced to a thickening of the bursa wall. I assumed that the signum of this species is reduced into a thickening of the bursa wall.

  2. T8 (inverted) forming a roughly Y-shaped sclerite, with its lateral margin parallel only in posterior 1/3. This state is known to have evolved several times within the Spilomelini. T8 (inverted) of Pyrausta also forms a roughly Y-shaped sclerite, with the lateral margin parallel only in posterior 1/3.

The monophyly of the group ((Glyphodes 1+ (Glyphodes 3 + (Dysallacta + (Agrioglypta + (Talanga)))))) is supported by the following possible apomorphies:

  1. Apex of uncus with hairs. This state is known to have evolved twice during its evolution within the Spilomelini. In Otiopora, apex of the uncus also has hairs.

  2. Cornutii absent. The loss of cornuti occurs several times within the pyraustines and frequently in all cram-bids group where cornuti are present in the ground plan.

DISCUSSION

In the 10 equally MP trees, the genus Chrysothyridia appears consistently within the Glyphodes group. The four genera, Glyphodes, Talanga, Dysallacta and Agrioglypta, that were believed to form a monophyletic group (Hampson, 1896; Shaffer in pers. comm), but do not do so unless the generic concept is expanded. Species of Chrysothyridia share at least one good apomorphy with the Glyphodes group, namely paired signa. The analysis also shows that the genus Glyphodes is not a monophyletic group, whereas the two species of Metallarcha do form a monophyletic group.

Well-supported relationships occur only for the following taxon pairs: Talanga and Agrioglypta, Hymenia and Spoladea, Diplopseustis and Aboetheta, Tatabotys and Hymenoptychis, and Otiopora and Eurryparodes.

The bootstrap analysis showed that the group formed by the 24 taxa of the Spilomelini is poorly supported. It is important to note that the low bootstrap support for monophyly does not amount to evidence of non-monophyly. A node supported by relatively few characters will be recovered infrequently in bootstrap replicates, even if there is evidence for their existence through apomorphies (Trueman 1993). According to Trueman (1993) and Bremer (1994) it is possible that in the bootstrap test some of the original characters may not be sampled and are thus omitted, whereas other characters are sampled more than once. At least one good apomorphy, namely, a strong, bilobed praecinctorium of abdominal tympanal organs supports the Spilomelini.

The group formed by Didymostoma + Synclera + Glyphodes 2 + Chrysothyridia + Glyphodes 1 + Glyphodes 3 + Dysallacta + Agrioglypta + Talanga also receives low bootstrap support (57%) but the T-PTP test gives significant support. The group formed by Didymostoma + Synclera and the group formed by Glyphodes 2 + Chrysothyridia + Glyphodes 1 + Glyphodes 3 + Dysallacta + Agrioglypta + Talanga did not receive bootstrap support. The Didymostoma + Synclera clade as the hipothesised sister group of the Glyphodes group, is not sufficiently supported as a separate monophyletic group. Therefore the definition of the Glyphodes group should be expanded to include also these two genera. There could be more genera of the Glyphodes group since only 17 genera apart of the Glyphodes have been sampled in this study.

This current analysis should be regarded as preliminary because it was based only on the adults of selected genera of Spilomelini. More species of the Spilomelini need to be examined, and further analysis should be based not only on adult characters but also on other characters sources such as the immature stages, molecular data and biological information in order to test the validity of the relationships proposed here.

Acknowledgments

I am grateful Dr. M. Horak from CSIRO Entomology, Canberra and Dr. P. J. Gullan from Division of Entomology, University of California, Davis, USA (formerly of the Australian National University) for discussing and advising during my research and also for critical comments that improved the manuscript. My thanks go to the Director of Australian National Insect Collection, CSIRO Entomology and to the School of Botany and Zoology, the Australian National University, Canberra, for providing of all the facilities for my research studies. I thank Mr Kevin R. Tuck who kindly arranged the loan specimens from the Natural History Museum, London and Mr M. Shaffer (Natural History Museum, London) and Mr E. D. Edwards (CSIRO Entomology), who generously answered numerous queries. This study was supported by a grant from GEF-Indonesia Biodiversity Project.

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Appendices

Appendix 1

The list of the specimens examined

Outgroup:

Hyalobathra archeleuca Meyrick

Queensland: 1#, 15°08′S 145°15′E, 37 km W of Cooktown, 22.i.1976, V. J. Robinson & Pratt. New South Wales: 1$, 28°25′S 152°46′E, Richmond River, x.1923, V. J. Robinson.

Isocentris filalis Guenée

Western Australia: 1#, 15°42′S 128˚41′E, Ivanhoe, 7.iv.1962, IFBC. Northern Territory: 1$, 14°27′S 132°16′E, Katherine, 17.iv.1962, IFBC.

Pyrausta panopealis (Walker)

Northern Territory: 1#, 12°36′S 132°52′E, Mudginberri, 25.v.1973, EDE & MSU. Queensland: 1$, 19°00′S 146°12′E, Paluma, 7.iv.1979, D. W. Frith.

Pyrausta testalis (Fabricius)

Queensland: 1 #, 28°05′S 153°27′E, Burleigh, 17.vi.1963, MSU; 1∃, 7°34′S 151°57′E, Toowoomba, 9.iv.1973, IFBC.

Ingroup:

Aboetheta pteridonoma Turner

New South Wales: 2#, 30°20′S 152°43′E, Dorrigo NP, xi.1926, V. J. Robinson; 3$, 5#, 30°24′S 152°21′E, Ebor, 5.i.1916.

Agrioglypta enneactis Meyrick

Fiji: 3$, 1#, Fiji, 1966/1967, H. S. Robinson (British Museum, 1967.236).

Chrysothyridia invertalis (Snellen)

Queensland: 2$, 2#, 13°45′S 143°22′E, Upper Leo Ck, McIlwraith Ra, 11.vii.1989, E. S. Nielsen & EDE & M. Horak.

Diasemiopsis ramburialis (Duponchel)

Queensland: 1$, 28°10′S 153°32′E, Coolangatta, 8.ix.1913. New South Wales: 1#, 35˚00′S 150˚45′E, Jervis Bay, 17.v.1956, IFBC.

Didymostoma euphranoralis (Walker)

Celebes [Sulawesi]: Type. 1#. Pyralidae. Brit. Mus. Slide No. 20111.

Diplopseustis perieresalis (Walker)

New South Wales: 1#, 28°25′S 152°46′E, Richmond River, 23.ix.1934, V. J. Robinson; 1 $, 29˚01′S 151˚04′E, Como West, 20.xi.1970, L. Willan & V. J. Robinson.

Dysallacta negatalis (Walker)

New South Wales: 3$, 28°25′S 152°46′E, Richmond River, i.1936, V. J. Robinson; 1#, 28°52′S 153°24′E, 2 km NE by E of Rous Mill, 18.x.1976, IFBC & EDE.

Eurrhyparodes bracteolalis (Zeller)

Northern Territory: 1$, 12°35′S 132°52′E, Magela Ck, 2 km N of Mudginberri, HS, 25.v.1973, EDE & MSU. Queensland: 1#, 12°40′S 142°41′E, Batavia Downs, 25.v.1993, P. Zborowski & A. Roach.

Glyphodes actorionalis Walker

Queensland: 1#, 10°41′S 142°32′E, Cape York, 10.vi.1928, W. B. Barnard; 1$, 10°48′S 142°28′E, Lockerbie Area, Cape York, 13-27.iv.1973, S. R. Monteith.

Glyphodes bicolor (Swainson)

Northern Territory: 1$, 11°01′S 136°45′E, Rimbija Isl., 14.i.1977, EDE; Queensland: 1#, 15°29′S 145°16′E, 1 km SE of Mt. Cook, Cooktown, 13.x.1980, EDE.

Glyphodes stolalis Guenée

Western Australia: 2#, 14°33′S 126°57′E, Drysdale River, 18-21.viii.1975, IFBC & MSU. Queensland: 1$, 10°48′S 142°28′E, Locker-bie, Cape York, 3.iii.1964, IFBC & MSU.

Hydriris chalybitis Meyrick

Northern Territory: 1$,, 11°09′S 132°09′E, Black Point, Cobourg Pen., 19.v.1977, EDE; 1#, 12°35′S 132°52′E, Magela Ck, 2 km N of Mudginberri, HS, 25.v.1973, EDE & MSU.

Hymenia perspectalis (Hübner)

Queensland: 5$, 1#, 27°28′S 153°02′E, Brisbane, 3.vi.1906, F. P. Dodd.

Hymenoptychis sordida (Zeller)

Queensland: 10$, 2#, 12°46′S 143°17′E, Claudie River, 5-16.v.1961, J. Macqueen.

Metallarcha diplochrysa Meyrick

New South Wales: 3$, 3#, 31°18′S 149°02′E, Warrumbungles, 24.x.1967, V. J. Robinson.

Metallarcha eurychrysa Meyrick

Queensland: 2$, 1#, 28°21′S 145°45′E, 32 km S by E of Cunnamulla, 20.iii.1990, EDE & J. H. Fisk.

Myrmidonistis hoplora Meyrick

Northern Territory: 2$, 12°28′S 132°52′E, Jabiluka Lagoon, 14 km N of Mudginberri, 13.xi.1972, EDE & MSU. Queensland: 1∃∃∃∃, 1#, 11°45′S 142°35′E, Heathlands, 16-22.iii.1994, P. Zborowski.

Niphograpta albiguttalis (Warren)

Queensland: 3$, 3#, 27°28′S 153°02′E, Brisbane, 10.x.1978.

Otiophora leucura (Lower)

Queensland: 1$, 13°59′S 143°33′E, Silver Plains HS, Cape York, 7.v.1965, J. L. Wassel; 2#, 27°28′S 153°02′E, Brisbane, 21.ii.1906 ñ 15.x.1915.

Sameodes cancellalis (Zeller)

Queensland: 1$, 13°40′S 142°40′E, Rokeby, 26.v.1993, P. Zborowski & Shattuck; 1#, 14°54′S 142°50′E, Long Lagoon, Coleman River, 16.viii.1993, P. Zborowski & J. Balderson.

Spoladea recurvalis (Fabricius)

Western Australia: 1$, 21°35′S 117°04′E, 1 km NNE of Millstream, HS, 3.iv.1971, MSU & Mitchell. Northern Territory: 1#, 23°42′S 133°52′E, Alice Springs, 15.ii.1966, Britton & MSU & Mc Innes.

Synclera traducalis Zeller

Africa: Type. 1#. Pyralidae. Brit. Mus. Slide No. 60.92.

Talanga sexpunctalis Moore

Northern Territory: 1$, 11°01′S 136°45′E, Rimbija Isl., 16.i.1977, EDE. Queensland: 2#, 17°16′S 145°54′E, Base Cableway, Mt. Bellenden-Ker, 23.x.1981, EDE.

Tatobotys janapalis (Walker)

Northern Territory: 1$, 15°54′S 136°32′E, Batten Point, 30 km NE by E Borroloola, 30.x.1975, EDE. Queensland: 1#, 19°16′S 146°49′E, Townsville, 19.iii.1900, F. P. Dodd.

Abbreviation and Symbols used in Appendix 1:

#: male

$: female

EDE: ED. Edwards.

IFBC: IFB. Common.

MSU: MS. Upton.

Appendix 2

Data matrix

i0289-0003-19-8-915-ta01.gif
and Hari Sutrisno "A Preliminary Study on Relationships among Selected Australian Members of the Tribe Spilomelini (Lepidoptera: Crambidae: Pyraustinae)," Zoological Science 19(8), (1 August 2002). https://doi.org/10.2108/zsj.19.915
Received: 15 December 2001; Accepted: 1 May 2002; Published: 1 August 2002
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