ヨツバコガネ属の系統分類学的研究（鞘翅目，コガネムシ科）

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九州大学学術情報リポジトリ

Kyushu University Institutional Repository

ヨツバコガネ属の系統分類学的研究（鞘翅目，コガネムシ科）

和田, 薫

https://doi.org/10.15017/1654970

出版情報：Kyushu University, 2015, 博士（理学）, 論文博士バージョン：

権利関係：Fulltext available.

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Systematic study of the genus Parastasia Westwood, 1842 (Coleoptera, Scarabaeidae)

ヨツバコガネ属の系統分類学的研究

（鞘翅目, コガネムシ科）

Kaoru WADA

和田薫

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i

9a. Parastasia nigriceps nigriceps Westwood, 1842 9b. Parastasia nigriceps inconstans Fairmaire, 1879 10. Parastasia nigromaculata Blanchard, 1850 11. Parastasia nigroscutellata Ohaus, 1901 12. Parastasia novoguineensis Ohaus, 1898 13a. Parastasia oberthueri oberthueri Ohaus, 1900 13b. Parastasia oberthueri ishigakiana Nomura, 1964 14. Parastasia percheroni Montrouzier, 1860

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iii 15. Parastasia sakaii Wada, 2004

16. Parastasia stella Kuijten, 1992 17. Parastasia sulcata Ohaus, 1911 18. Parastasia takeshii Wada, 1997 19. Parastasia weberi Ohaus, 1898 Parastasia discolor –group

1. Parastasia aberrans Kuijten, 1992 2. Parastasia alternate Arrow, 1899

3. Parastasia bicolor Westwood, 1842 4. Parastasia carsteni Wada, 2004 5. Parastasia dempuensis Wada, 2008

6a. Parastasia discolor discolor Westwood, 1842 6b. Parastasia discolor mirabilis Arrow, 1899 6c. Parastasia discolor scutellaris Erichson, 1845 7. Parastasia diversipennis Ohaus, 1911

8. Parastasia glottidion Kuijten, 1992 9. Parastasia hainanensis Wada, 2009 10. Parastasia helleri Ohaus, 1898 11. Parastasia isidai Wada, 1989

12. Parastasia kinibalensis OHAUS, 1901 13. Parastasia kolakana Wada, 1996 14 Parastasia kuijteni Wada, 1996 15. Parastasia laratina Ohaus, 1903 16. Parastasia lobata Kuijten, 1992

17. Parastasia masumotoi Wada & Muramoto, 1999 18. Parastasia melanocephala Burmeister, 1844 19. Parastasia melanocephaloides Ohaus, 1900 20. Parastasia pascoei Waterhouse, 1895 21. Parastasia rufolimbata Blanchard, 1850 22. Parastasia rufonigra Ohaus, 1911 23. Parastasia selangorica Kuijten, 1992 24. Parastasia sulcipennis Gestro, 1888

25. Parastasia vittata Snellen van Vollenhoven, 1864 26. Parastasia wallacea Kuijten, 1992

27. Parastasia xanthopyga Kuijten, 1992

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iv Parastasia canaliculata –group

1. Parastasia andamanica Ohaus, 1898 2. Parastasia basalis Candeze, 1869 3. Parastasia binotata Westwood, 1842 4. Parastasia birmana Arrow, 1899

5. Parastasia canaliculata Westwood, 1842 6. Parastasia cingala Arrow, 1899

7. Parastasia circumferens Arrow, 1899 8. Parastasia coquerelii Fairmaire, 1868 9. Parastasia dalatina Kuijten, 1992 10. Parastasia dimidiata Erichson, 1845

11. Parastasia ephippium Snellen van Vollenhoven, 1864 12. Parastasia jamsonae Wada, 2008

13. Parastasia kangeanensis Wada, 2007 14. Parastasia klossi Ohaus, 1926 15. Parastasia mitsumata Wada, 2007 16. Parastasia moseri Ohaus, 1903 17. Parastasia polita Ohaus, 1911

18. Parastasia rufopicta Westwood, 1842 19. Parastasia sumbawana Ohaus, 1898 20. Parastasia takahikoi Wada, 2007 21. Parastasia timorensis Arrow, 1899 22. Parastasia yukioi Wada, 1989

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Parastasia westwoodi –group

1. Parastasia anomala Arrow, 1899 2. Parastasia asahi Wada, 2008 3. Parastasia burmeisteri Ohaus, 1898 4. Parastasia duchoni Ohaus, 1898 5. Parastasia fakfakensis Wada, 2008 6. Parastasia femorata Burmeister, 1844 7. Parastasia gestroi Ohaus, 1900 8. Parastasia incurva Ohaus, 1923 9. Parastasia indica Ohaus, 1898 10. Parastasia intermedia Ohaus, 1938 11. Parastasia lombokensis Wada, 2008 12. Parastasia marginata Boisduval, 1835

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v 13. Parastasia moultoni Ohaus, 1911

14 Parastasia nigripennis Sharp, 1888 15. Parastasia punctulata Ohaus, 1900 16. Parastasia vietnumensis Wada, 2008 17. Parastasia westwoodii Westwood, 1842 18. Parastasia yasutoshii Wada, 2008

Ⅶ. Acknowledgements 255

Ⅷ. References 256

Figures

Fig.I-1. The adults of Parastasia nigripennis Sharp, 1888 are observed feeding and mating on flowers of Homalomena propinqua Schott

Fig.III-1. Character positions of general morphology: the adult of Parastasia ferrieri Nonfried, 1895; male; dorsal and ventral view

Fig. III-2. Character positions of general morphology: the adult of Parastasia ferrieri Nonfried, 1895; male; lateral view

Fig. III-3. Character positions of general morphology: the head Fig. III-4. Dorsal view of labrum

Fig. III-5. Dorsal view of mentum Fig. III-6. Dorsal view of maxilla Fig. III-7. Dorsal view of mandible

Fig. III-8. Dorsal view of mandible showing apical teeth and mandibular plate Fig .III-9. Parastasia canaliculata Westwood, 1842; male; hind wing

Fig. III-10. Mesotarsi showing unguitractor plate

Fig. III-11. Prosternal projection showing the apical portion Fig. III-12. Preprosternum showing the ridge in the middle Fig. III-13. Dorsal view of male genitalia

Fig. III-14. Inner sac of male genitalia showing membranous structure, spine and sclerotized appendages

Fig. III-15. Inner sac of male genitalia showing membranous structure of the genus Parastasia spp.

Fig.IV-1. Strict consensus tree of the 567 MP trees (out-group: Melolonthinae, genera Polyphylla, Dasylepida, Helotrichia). Bootstrap values (over 50%; 1000 replications) were shown above each branch

Fig.IV-2. Strict consensus tree of the 20581 MP trees (out-group: genus Desmonyx and

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vi subfamily Dynastinae)

Fig.IV-3. Strict consensus tree of 278 MP trees based on the complete data set analysis (out-group : Melolonthinae, genera Polyphylla, Dasylepida, Holotrichia)

Fig. IV-4. Strict consensus tree of 18 MP trees based on the complete data set analysis of typical species (out-group: Melolonthinae, genera Polyphylla, Dasylepida, Helotrichia) Fig.IV-5. Strict consensus tree of 278 MP trees based on the confluens group and the

bimaculata group species data set analysis (out-group: genus Desmonyx)

Fig.IV-6. Strict consensus tree of 18 MP trees based on the discolor group species data set analysis (out-group: genus Desmonyx)

Fig.IV-7. Strict consensus tree of 13 MP trees based on the canaliculata group species data set analysis (out-group: genus Desmonyx)

Fig.IV-8. Strict consensus tree of 43 MP trees based on the westwoodi group species data set analysis (out-group: genus Desmonyx)

Fig.V-1. Distribution map of Parastasia bimaculata spp.

Fig.V-2. Distribution schematic diagram of the Parastasia confluens-group Fig.V-3. Distribution map and Phylogenetic relationships of the ferrieri subclade Fig.V-4. Distribution schematic diagram of the Parastasia bimaculata-group

Fig.V-5. Distribution map and Phylogenetic relationships of the bimaculata+marmorata subclade

Fig.V-6. Distribution schematic diagram of the Parastasia discolor-group Fig.V-7. Distribution schematic diagram of the Parastasia canaliculata-group

Fig.V-8. Distribution map and Phylogenetic relationships of the basalis+cingala+dalatina subclade

Fig.V-9. Distribution schematic diagram of the Parastasia westwoodi-group Fig.V-10. The center of distribution of Parastasia groups

Fig.V-11. Postulated distribution of land and sea in SE Asia at 10 Ma. (R. Hall 1998)

Fig.V-12. A paleogeographical reconstruction of the Sunda-Sahul region during one of the Quaternary glacical maxima

Fig.VI-1. General morphology of the adult: Parastasia kuijteni K. Wada, 1996; male; dorsal and ventral view

Fig.VI-2. General morphology of the adult: Parastasia glottidion Kuijten, 1992; male; lateral view

Fig. VI-3. Parastasia westwoodii Westwood,1842; male; elytron Plates

Plate. I: 1-14, Parastasia confluens-group

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vii Plate. II: 1-19, Parastasia bimaculata-group Plate. III: 1-27, Parastasia discolor-group Plate. IV: 1-22, Parastasia canaliculata-group Plate. V: 1-18, Parastasia westwoodi-group Plate.VI:1-36, Out-group

Maps

Map I: 1-14, Parastasia confluens-group Map II: 1-19, Parastasia bimaculata-group Map III: 1-27, Parastasia discolor-group Map IV: 1-22, Parastasia canaliculata-group Map V: 1-18, Parastasia westwoodi-group Tables

Table II-1. The species-groups of the genus Parastasia

Table III-1. List of the species of genus Parastasia studied for the analysis.

Table III-2. List of the species of out-groupｓ studied for the analysis.

Table III-3. Character matrix Appendix

Appendix 1. Check list of the genus Parastasia Westwood, 1842 by Wada 2015

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Abstract

This work provides the phylogeny and classification of the genera Parastasia and Desmonyx and revise the taxonomy of the genus Parastasia Westwood, 1842 (Coleoptera, Scarabaeidae).

Analyses were conducted using 152 morphological characters of 143 taxa as follows: 108 within the genus Parastasia and 35 of the Rutelinae (Rutelini, Anomalini and Adoretini), Dynastinae and Melolonthinae as the outgroups.

Resuls of the analysis demonstrates the following: (1) the genus Parastasia is included in the subfamily Dynastinae, (2) the genus Parastasia is monophyletic and includes four monophyletic groups, (3) the genera Chyphelytra, Ceroplophana, Dicaulocephalus, Fruhstorferia, Kibakoganea, Lutera, Masumotokoganea, Peperonota and Rutelarcha of Asian species are included in the tribe Rutelini and each genus is confirmed its phylogenetic position and (4) Desmonyx is confirmed the phylogenetic position and is included in the subfamily Dynastinae.

Based on the results of phylogenetic analyses, new classifications of the genus Parastasia with five species groups are proposed: (1) paraphyletic basal taxa as the confluens –group, (2) monophyletic bimaculata clade as the bimaculata-group, (3) monophyletic discolor clade as the discolor-group, (4) monophyletic canaliculata clade as the canaliculata-group and (5) monophyletic westwoodi clade as the westwoodi-group.

Description and keys to all species-groups, 100 species and 7 subspecies of the genus Parastasia are indicated. The biogeography and speciation of each species group are also discussed.

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1 I. Introduction

Approximately 94 species and 9 subspecies of the genus Parastasia Westwood, 1842 (Coleoptera, Scarabaeidae, Rutelinae, Rutelini, Parastasiina) have hitherto been known from the Palearctic and the Oriental Regions (Krajcik, 2008).

The genus Parastasia is widely distributed from the Andaman Islands and Indo-Himalaya area, through the Indochina Peninsula, to Sundaland and Oceania.

Moreover, endemic species are also distributed on the Ogasawara and Fiji Islands (islands on the Pacific Ocean), and the Seychelles (in the Indian Ocean). One species is distributed eastern North America (Machatschke, 1972).

Some species of Parastasia are widely distributed, while other species are indigenous to regions. Evolution and speciation in the genus Parastasia has been strongly influenced by not only ecological factors but also historical factors connected deeply with the complicated geological history of their areas of distribution.

The species of Parastasia are moderately-sized (1-4 cm in body length) and dark colored beetles having the following characters: clypeus with two or four denticles, fronto-clypeal suture obsolete in lateral 1/3 to 2/5, inner apices of the fourth meso- and metatarsomeres each with one prolonged triangular process in ventral view.

Fundamentally, the coloration of the body is red to black with yellow to dark brown maculations on the dorsal surface.

Some species exhibit unusual character traits such as a prolonged clypeal apex (P.

dimidiata Erichson, 1845), forwardly produced metasternal process (P. kinibalensis Ohaus, 1901) or unusual form of the mesotarsal claw with the outer claw split and greatly widened in the male (P. westwoodi Westwood, 1842).

Detailed ecology of adults is almost unknown, except that they are collected by light traps. From the dark coloration and the habit of flying to light, it seems that the adults are nocturnal. In some species of Parastasia, the adults are observed feeding on flowers of Homalomena propinqua Schott (P. gestoroi Ohaus, 1900) (Fig. I-1）and fermenting fruits (P. canaliculata Westwood, 1842) (Yumoto, 1999; Wada, 1984).

Larvae feed on decaying wood. Some species have been collected from reddish rotten wood (P. ferierri Nonfried, 1895) or white rotten wood (Parastasia nigriceps Westwood, 1842), and from the detritus around the termite colony (e.g. P. nigriceps inconstans Fairmaire, 1879) (Suzuki, 2000).

Considering the morphological variety mentioned above, the genus Parastasia may have interesting habits. Little is known, however, about life history, food habits, mating behavior, and so on in adults of this genus. And for many species, the larvae and

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2 pupae are still unknown.

A new classification of the tribe Rutelini including the genus Parastasia is proposed by Jameson (1998) as the result of phylogenetic analysis. She proposed the genera Acrobolbia and Peltonotus, both of which have been placed in the tribe Rutelini, should be placed in the subfamily Dynastinae. But the phylogenetic position of Parastasia has still been unclear.

In the first section of this study, I conduct phylogenetic analyses mainly based on morphological characters to reconfirm the monophyletic group “Parastasia lineage” and to make clear the relationships of the genus Parastasia. In the second section, I discuss the process and feature of the speciation based on the geological history relevant to the phylogenetic relationships of species-groups of Parastasia. Finally, I will propose a new taxonomic system based on the phylogenetic relationships of the species belonging to Parastasia.

II. Taxonomic history

The tribe Rutelini in the subfamily Rutelinae, consists of approximately 70 genera and is closely related to the subfamily Dynastinae (Machatschke, 1972). It has been pointed out that there are some systematic problems about the taxonomic position of the genera that belonged to the tribe Rutelini or the subfamily Dynastinae. Historically, the genus Parastasia Westwood, 1842 had been treated as the members of the subtribe Parastasiina of the tribe Rutelini that is closely related to subtribe Fruhstorferina and subtribe Rutelina (Arrow, 1917; Machatschke, 1972). However, the relationships of Parastasiina with subtribe Fruhstorferina, subtribe Rutelina and particularly subfamily Dynastinae were ambiguous.

A key to solve this problem was given by Jameson (1998) with the result of phylogenetic analysis. She showed phylogenetic relationships among 28 genera and 2 subgenera of the tribe Rutelini and pointed out invalidity of subtribal classification.

As for the genus Parastasia, many new genera had been described based on some species because of variability of the forms and few numbers of specimens collected. The genus Parastasia was described by Westwood (1842) based on Parastasia canaliculata as the type species. Then, Burmeister (1844) and Lacordaire (1856) described Parastasia in more detail. Although Burmeister (1844) mentioned the genus Caelidia Dejean, 1833, Dejean had not been described the genus Caelidia and only catalogued on Caelidia quinquemaculata Dejean, 1833 . Arrow (1899) considered the genus Caelidia as

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nomen nudum and treated as a synonym of Parastasia. The genus Barymorpha Guerin- Méneville, 1843 was described based on Barymorpha bimaculata Guerin- Méneville, 1843 as a type. This genus was divided from Parastasia based on the difference in the structure of claws. Hoverer, it was considered as a synonym of Parastasia by Burmeister (1844). The genus Polymoechus Leconte, 1856 is the only species which occurs in North America. This genus was described based on Polymoechus brevipes Leconte, 1856 as a type. Then, it was considered as a synonym of the genus Parastasia by Arrow (1917). Although the genus Urleta Westwood, 1875 was described based on Urleta ometoides Westwood, 1875 as a type, Urleta ometoides was treated as a junior synonym of P. dimidiata Erichson, 1845 and the genus Urleta was considered as a synonym of the genus Parastasia by Arrow (1899). The genus Echmatophorus Waterhouse, 1895 was described based on Echmatophorus pascoei as a type. Then, the genus Echmatophorus was considered as a synonym of Parastasia by Arrow (1899).

Sawada (1938) advocated the genus Ohkubonus with Ohkubonus quadridentatus Sawada, 1938 as a type, based on the difference in the structure of mouth parts and so on. Afterwards, Sawada (1950) treated Parastasia ferrieri Nonfried, 1895 as the genus Ohkubonus and considered O. quadridentata as a junior synonym of O. ferrieri. In his study, Sawada considered that the genus Ohkubonus would be classified to the subtribe Dynastini. Nomura (1964, 1965, 1966) treated the genus Ohkubonus as invalid and as the synonym of Parastasia. However, Kobayashi (1973, 1983) and Nakane (1983) treated the genus Ohkubonus as valid. The genera Cyphelytra Waterhouse, 1875, Lutera Westwood, 1875 and Rutelarcha Waterhouse, 1874 were considered as the synonyms of the genus Parastasia by Arrow (1917). Afterwards, these genera were treated as valid by Ohaus (1938b) and Kuijten (1988). Kuijten (1988) mentioned that the genera Rutelarcha, Cyphelytra and Lutera must be united, and transferred them to the subtribe Rutelina.

The total 150 species of the genus Parastasia were described by Westwood (1842), Guerin-Méneville (1843), Burmeister (1844), Erichson (1845), Blanchard (1850), Leconte (1856), Montrouzier (1860), Snellen van Vollenhoven (1864), Fairmaire (1868, 1879, 1883), Candeze (1869), Westwood (1842), Ohaus (1898, 1900, 1901, 1903, 1911, 1923, 1926, 1938), Arrow (1899), Gestro (1876, 1888), Sharp (1888), Schaufuss (1887), Nonfried (1892, 1895), Waterhouse (1895), Sawada (1938), Nomura (1964), Frey (1969), Nakane (1983), Kuijten (1992), Ghai et al. (1988) and Wada (1989, 1996, 1997, 1999, 2003, 2004, 2007, 2008, 2009). However, 45 species were treated as invalid as junior synonyms of other species for the reason that it is often confused that the color form and that male and female were separated as another species in the past classification. The

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classificatory arrangement concerning on the all known 72 species and 10 subspecies was made by Kuijuten (1992).

The revisions and catalogues on the genus Parastasia were published by Ohaus (1934b), Machatschke (1972), Kuijten (1992) and Kracjik (2008). In his revisional work, Ohaus (1934b) divided the genus Parastasia into 10 species groups: confluens group, dolens group, bimaculata group, nigromaculata group, discolor group, canaliculata group, anomala group, westwoodi group, vittata group and heterocera group (Table II-1).

Arrow (1917) considered that the genera Cyphelytra Waterhouse, 1875, Lutera Westwood, 1875 and Rutelarcha Waterhouse, 1874 were junior synonyms of the genus Parastasia. Machatschke (1972) added three species groups: ochracea group, luteola group and quadimaculata group that are included in species of the genera Cyphelytra Waterhouse, 1875, Lutera Westwood, 1875 and Rutelarcha Waterhouse, 1874 (Table II-1). Kuijten (1992) postponed dividing the species groups of the genus Parastasia until data and knowledge have much increased.

However, there were many discoveries of new species after Kuijuten (1992).

Moreover, the species groups proposed by Ohaus (1934b) were not fully examined until now, and their phylogenetic relationship is not clear.

All the nomenclatural acts in this doctoral thesis would be disclaimed based on the Article 8.3.

of ICZN.

III. Material and Methods

1. Type specimens and taxonomic materials

In the course of this study, I examined type series as far as possible. Specimens examined for this study, including type specimens, were loaned from 18 institutes and private collections. The collection managers and curators are listed in parenthesis. A total about 5000 specimens formed the basis of this study. Particularly, the materials were provided for analysis from Dr. Mary Liz Jameson of the University of Nebraska State Museum, Lincolon (Wichita state University in Wichita, Kansas), Dr. Pol Limbourg of the Institut royal des Sciences naturelles de Belgique, Mr. Takeshi Matsumoto, Hyogo, Mr. Masayuki Fujioka, Tokyo, Mr. Kaoru Sakai, Tokyo and Mr. Shinji Nagai, Nagano (all from Japan). Acronyms for loaning institutes and private collections are indicated below.

BMNH The Natural History Museum, London, UK.

(Malcolm D. Kerley)

DEI Deutsches Entomologisches Institut, Eberswalde, Germany.

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5 (Lother Zerche)

FREY George Frey Collection at ZSMC, München, Germany.

(Martin Baehr, Max Kuhbander)

HECO The Hope Entomological Collections of the University Museum, Oxford, UK.

(C. O'tool)

ISNB The Institut royal des Sciences naturelles de Belgique, Bruxelles, Belgium.

(Pol Limbourg)

ZMHB Museum für Naturkunde der Humboldt Universität zu Berlin, Berlin, Germany.

(Manfred Uhlig, Joachim Schulze, Johannes Frisch, Joachim Willers) MNHN Muséum National d'Histoire Naturelle, Paris, France.

(Roger-Paul Dechambre, Thierry Deuve, Azadeh Taghavian) NSMT National Science Museum (Nat. Hist.), Tsukuba, Japan.

(Shuhei Nomura)

HNHM Magyar Természettudományi Múzeum, Budapest, Hungary.

(Ottó Merkl)

RIEB The Reserch Institute of Evolutionary Biology, Tokyo, Japan.

(Shuhei Yamaguchi, Toshiaki Aoki)

RMNH The Nationaal Natuurhistorisch Museum, Leiden, Netherlands.

(J. Krikken, Ma Eulàlia Gassó Miracle)

SMTD Staatliches Museum für Tierkunde, Dresden, Germany.

(Dirk Ahrens)

UNSM The University of Nebraska State Museum, Lincolon, USA.

(Mary Liz Jameson)

ZSMC Zoologische Staatssammlung des Bayerischen Staates, München, Germany.

(Martin Baehr)

PCF Private collection of Masayuki Fujioka, Tokyo, Japan.

PCN Private collection of Shinji Nagai, Nagano, Japan.

PCZ Private collection of Carsten Zorn, Gnoien, Germany.

PCW Private collection of Kaoru Wada, Tokyo, Japan.

2. Dissection

The technique of dissection was important for scrutinizing and examination of characters. The following parts can be observed only by dissection; mouth parts (galea, labrum, mentum, maxilla, mandibles), hind wings, abdomen, spiracles, genitalia and internal sac.

Dried specimens were softened by soaking boiling water and separated into 3 parts, i.e. head, thorax and abdomen. Mouth parts were extracted from the head under microscope vision using

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tweezers and insect pins. In most cases, the left mandible, left maxilla, labrum and mentum were extracted and card-mounted using aqueous glue. The left hind wing, pterothorax and abdominal sternites were extracted from the abdomen using tweezers and insect pins. Furthermore, the genitalia and spiculum gastrale were extracted from inside of the abdomen.

The male genitalia were entirely softened by putting in 5% KOH at least overnight. After wards, internal sac was turned out by applying water pressure from the base of genitalia using small-sized hypodermic syringe under the microscope. The old and non-reversible internal sac was extended with tweezers as reverse-side. The internal sac was kept in the tube pipe with glycerin.

The genitalia and spiculum gastrale were card-mounted using aqueous glue. The left hind wing was extracted from the body. The wing was dried and put between slide glasses in an extended position. The slides were fastened with a wide clip and scotch tape. The wing was mounted on the plastic plate using aqueous glue. The card-mounted mouth-parts, genitalic parts and the abdomen were pinned. The head, thorax and abdomen were replaced after dissection and pinned. The data labels were attached to the materials.

3. Phylogenetic Analysis

3-1. Examination taxa for phylogenetic analysis

Jameson (1990, 1994, 1996, 1997) analyzed of the tribe Rutelini based on many morphological characters. In this study, the characters used by Jameson are re-examined. Characters available for the phylogenetic analysis of the genera and species from Southeast Asian are added to those employed by Jameson. Phylogenetic analysis was based on 152 characters. Characters used in this analysis partly followed those of the preceding study by Jameson (1998), with the addition of structures of the epipharynx, shape of the male genitalia and inner sac structure of male genitalia. All the 100 species and 8 subspecies of the genus Parastasia were analyzed as in-group (Table III-1). Of these 100 species, some characters of 25 species were not able to observe because one of the sexes was collected. Analyzed all species were illustrated in Plate I-V.

In order to determine the taxonomic status of the genus Parastasia, 22 genera of out-group taxa were selected from the Rutelinae (Rutelini, Anomalini and Adoretini), Dynastinae and Melolonthinae, that is, the genera Ceroplophana, Cyphelytra, Desmonyx, Dicaulocephalus, Fruhstorferia, Kibakoganea, Lutera, Masumotokoganea, Peperonota, Rutelarcha (Rutelini), Anomala, Mimela (Anomalini), Adoretus (Adoretini), Blabephorus, Eophileurus, Oryctomorphus, Peltonotus, Pseudohomox, Rutelisca (Dynastinae), Dasylepida, Holotrichia and Polyphylla (Melolonthinae) (Table III-2). Thirty-five species of out-group taxa listed above were used to analyze the relationships of Parastasia. Especially the out-group genera Lutera, Chyphelytra, Rutelarcha of the Rutelini (Rutelinae), which were once included in the genus Parastasia (Arrow,

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1917; Machatschke, 1972), were analyzed for all the species. To confirm the relationship between Parastasia and subfamily Dynastinae, the species of the following genera of Dynastinae were used to analyze: Oryctmorphus, Eophileurus, Pseudohomox, Blabephorus and Peltonotus. The genera Polyphylla, Dasylepida and Holotrichia of Melolonthinae were used for out-group to analyze the relationship between Rutelinae and Dynastinae. Analyzed all the species were illustrated in Plate VI.

1-36.

3-2. Characters and character states

Character matrix is shown in Table III-3. All the characters were discrete rather than continuous, and were treated as unweighted. Characters were coded as either binary or multistate (0-6). Multistate characters were treated as unordered because transformation series could not be determined a priori. Unknown data that were not observed were coded as “?”. Characters used for phylogenetic analysis are indicated below and shown in Figs. III-1-15.

Head

1. Antenna 10-segmented (0), or 9-segmented (1).

2. Lamellar antennal club 3-segmented (0), 4-segmented or more than 4-segmented (1).

3. Antennal club almost the same length or shorter than the rest segment of antenna (0), longer than the rest segment of antenna but under twice length of the segment (l), longer or prolonged over twice length of the segment (2).

4. Clypeus semicircular (0), rounded (1), truncate (2), trapezoidal (3), or quadrate (4).

5. Clypeus almost glabrous (0), furnished with spine-like setae (1), or with long, slender setae (2).

6. Apical margin of clypeus not prolonged (0), or prolonged (1).

7. Clypeal apex plane or weakly reflexed (0), reflexed and emarginate in middle (1), with one denticle (2), with two denticles (3), with four denticles (4), or prolonged with two horns (5).

8. Clypeus without horn or tubercle (0), with a horn or tubercle in apical to medial portions in male (1), or with two horns or tubercles in apical to medial portions in male (2).

9. Baso-medial portion of the clypeus plane (0), or raised (1).

10. Base of the clypeus not reflexed (0), or reflexed (1).

11. Fronto-clypeal suture obsolete or lacking in the middle (0), or complete (1).

12. Fronto-clypeal suture laterally plane (0), or raised above the plane of the frons (1). This character is obscure in the enlarged mandibles of the males of the genera Dicaulocephalus and Peperonota, though the females apparently show state (1), so that the state (1) might be the ground plan for the genera.

13. Eye canthus without ridge or thickening (0), with a ridge or thickening (1), or strongly protruded (2).

14. Eye small, post-occipital region large (0), or eye large, size of post-occipital region reduced (1).

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15. Head without depressions between eyes (0), with shallow depressions (1), or with swelling (2).

16. Frons without horn or tubercle (0), or with horn or tubercle (1).

17. Eye without a circumocular depression in ventral view (0), or with a circumocular depression (1).

18. Head densely furnished with hair-like setae (0), densely furnished with spinose setae (1), or partly furnished with hair-like setae in lateral portions or almost glabrous (2).

19. Vertex without horn or tubercle (0), or with a horn or tubercle in medial portion (1).

Mouth parts

20. Apex of the labrum produced beyond the clypeal apex (0), or hidden and not obviously produced (1).

21. Labrum vertically produced (0), or horizontally produced (1).

22. Labrum lacking median and apical process (0), with weakly produced median and apical process (1), or with a produced, tooth-like median and apical process (2).

23. Labrum inflated at apex (0), or flattened at apex (1).

24. Apex of labrum forked (0), rounded (1), narrowly truncate for less than 1/3 the width (2), broadly truncate for more than 1/3 the width (3), triangularly produced (4), or concave medially (5).

25. Base of submentum level to the mentum (0), or perpendicularly produced (1).

26. The mentum almost as long as width (0), or longer than width (1).

27. Mentum with apex not recurved (0), or apex protruded posteriad and recurved (1).

28. Mentum with apex protruded (0), or apex not protruded (1).

29. Apical margin of mentum slightly rounded or straight (0), emarginate (1), notched (2), or with tooth-like projection (3).

30. Mentum almost plane in baso-medial portion (0), or with a large hole (1).

31. Stipes not flange-like (produced apically and laterally) (0), or flange-like (1).

32. Lacinia with apically-produced tooth on the inner margin (0), or lacking lacinial tooth (1).

33. Maxilla with more than three obvious teeth (0), with teeth reduced to one or two (1), or with bristles or pegs (2).

34. Maxilla with medial tooth fused (0), or medial tooth hinged (1).

35. The length of the 2nd maxillary palpal segment shorter than the 4th (0), or almost equal or longer than the 4th (1).

36. Mandible not produced beyond the labrum (0), or produced beyond the labrum (1).

37. Mandible at apex without reflexed teeth (0), with one reflexed tooth (1).

38. Mandible reflexed at apex (0), or not reflexed (1).

39. Mandible not produced in male (0), distinctly protruded and curved backwards in male (1), or distinctly protruded forwards in male (2).

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40. Lower apex of mandibular plate without denticles (0), with one denticle (1), with two denticles (2), or with three denticles (3).

41. Mandibular crown prolonged (0), or not prolonged (1).

42. Basal margin of mola almost straight or slightly curved (0), or concave (1).

43. Mola with rows of denticulation almost parallel (0), or those of the posterior portion perpendicular to those of the anterior portion (1).

44. Mola with rows of denticulation partly vestigial (0), with shallow elongate longitudinal grooves or clathrate shallow grooves (1), with deep grooves (2), or with large depressions on the mola (3).

45. Mola without protrusion (0), or with a protrusion in apical portion (1).

46. Mola without denticles (0), or with denticles in lateral portion (1).

47. Epipharynx dorsally flattened and without setae (0), vaulted in apical to medial portions and furnished with setae, the remaining part is flattened and without setae (1), or entirely vaulted and with setae (2).

48. Epitorma not fused and obviously defined (0), or entirely fused and not defined (1).

49. Epitorma plane with glabrous part in medial portion (0), lacking the glabrous part in medial portion (1), or with broadly plane area in medial portion and confused (2).

50. Torma weakly chitinous and membranous (0), or strongly chitinous (1).

51. Epipharynx with one simple lateral torma (0), or with two lateral tormae (1).

Pronotum

52. Basal bead of pronotum complete (0), incomplete (1), or lacking (2).

53. Pronotum without depressions (0), with a vague depression in apico-medial portion (1), or with an obvious depression in apico-medial portion (2).

54. Lateral region of pronotum without fovea (0), or with fovea (1).

55. Pronotum without depressions (0), or with a pair of vague depressions in about lateral 1/3 (1).

56. Anterior margin of pronotum with apical bead complete (0), or apical bead incomplete (1).

57. Pronotum densely furnished with hair-like setae (0), partly furnished with hair-like setae inmarginal portion (1), with spinose setae (2), or almost glabrous (3).

Scutellum

58. Base of the scutellum obliquely angled (0), depressed below the plane of the elytra (1), or with a medial, plane extension and depressed sides (2).

59. Width of the scutellum greater than length (0), almost equal to length (1).

Width of the scutellum was measured from the meeting point of elytral base and the base of scutellum. Length was measured from the base of the scutellum to its apex.

Elytra

60. Basal portion of elytra along the scutellum plane (0), or depressed (1).

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If this character appears in one sex only, it was regarded as state (1) in Parastasia canaliculata.

61. Rows of punctures on elytra shallow with the intervals almost plane (0), or deep with the intervals obviously convex (1).

If this character appears in one sex only, it was regarded as state (1) in Parastasia sulcipennis.

62. Rows of punctures on elytra almost parallel (0), or contorted in middle and connect with one another (1).

63. Lateral margin of elytra in female plane (0), or sinuate and raised at apical 1/3 (1).

64. Elytral margin without a membranous border (0), or with a membranous border (1).

65. Elytra entirely furnished with hair-like setae (0), furnished with hair-like setae only in lateral portions (1), almost glabrous (2), or furnished with spinose setae (3).

Hind wing

66. AP3+4 of hind wing simple at base (0), or with a bulbous, setaceous, enlarged vein at base (1).

67. Basal half of ScA of hind wing without pegs (0), or with pegs (1).

68. Apical half of ScA of hind wing lacking pegs and row of setae (0), with pegs but lacking row of setae (1), with both pegs and row of setae (2), or with row of setae but lacking pegs (3).

69. RA3+4 of hind wing without row of setae (0), or with row of setae (1).

70. Vein AA1+2 of hind wing lacking (0), protruded (1), connected with CuA (2), or bent downwards and subequal to AA3+4 (3).

71. Basal region of vein AA3+4 of hind wing almost straight (0), or AA3+4 intercepted (1).

Spiracles and Tergites

72. Pleural suture between sternite 7 and tergite 7 distinct (0), or indistinct (1).

73. Tergite 2nd to 4th furnished with setae in medial portions (0), only 2nd tergite with long setae (1), or glabrous (2).

74. Tergites on lateral edge unicolored (0), or bicolored (1).

Propygidium

75. Propygidium long (more than 2/3 of pygidium and longer than each tergite) (0), or short (almost equal to or slightly longer than each tergite) (1).

76. Surface of propygidium with setigerous punctures (0), or punctate but without setae (1).

Pygidium

77. Pygidium of male furnished with setae (0), or glabrous (1).

78. Pygidium of female furnished with setae (0), or glabrous (1).

Appendages: Coxae

79. Mesocoxae more or less contiguous, not widely separated (0), or widely separated (1).

80. Metacoxae with medial region contiguous with metatrochanter and not produced posteriorly (0), with medial region produced posteriorly but not beyond the posterior border of the metatrochanter (1), or with medial region distinctly produced posteriorly beyond the posterior

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Appendages

81. Anterior marginal portions of metacoxa with spur and setae (0), with setae only (1), or with spur only (2).

Appendages: Tibiae

82. Base of the inner protibia notched (0), or simple (1).

83. Protibia with external edge tridentate (0), bidentate (1), or quadridentate (2).

84. Mesotibial apex without spinose process at outer margin (0), with one spinose process at outer margin (1), with two spinose processes at external edge (2), or with three spinose processes at external edge (3).

85. Apex of mesotibia simple at middle (0), with one spinose process (1), or with many spines (2).

86. Mesotibia of male without produced corbel (0), or with produced corbel (1).

87. Apex of mesotibia with many spinose setae (0), with sparse stout spinose setae (1), with sparse hair-like setae (2), or lacking setae (3).

Character state (0) is defined as spinose setae separated by about one seta-width. Character state (1) is defined as spinose setae separated by more than one seta-width.

88. Apex of metatibia without tooth at outer margin (0), with a tooth at apex of outer margin (1), bidentate at external edge (2), or tridentate at external edge (3).

89. Apex of metatibia with many spinose setae (0), sparse stout spinose setae (1), sparse hair-like setae (2), or lacking setae (3).

Character state (0) and (1) is defined as spinose setae more than 0.075 mm in width. Character state (2) is defined as hair-like setae less than 0.05 mm in width.

90. Fifth mesotarsomere in male shorter than the 1st-4th ones (0), or equal or longer than the 1st-4th ones (1).

91. Fifth metatarsomere in male equal or shorter than the 1st one (0), longer than the 1st one, but shorter than the 1st-4th ones (1), or equal to or longer than the 1st-4th ones (2).

92. Outer edge of first metatarsomere produced (0), not produced (1), or with a spinous protrusion (2).

93. Inner apex of the fourth metatarsomere lacking attenuation (0), with a triangular process in lateral view (1), or with three prolonged triangular process in lateral view (2).

94. Ventrolateral apex of the fourth metatarsomere in male with two long, hair-like setae (0), with one outer hair-like seta and one inner stout, spinose seta (1), with one inner and one outer stout and spinose setae (2), with several spinose setae (3), or with several spines, spinose setae and hair-like setae (4).

95. Ventrolateral apex of the fourth metatarsomere in female with two long, hair-like setae (0), with one outer hair-like seta and one inner stout, spinose seta (1), with one inner and one outer stout

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and spinose setae (2), with several spinose setae (3), or with several spines, spinose setae and hair-like setae (4).

Appendages: Tarsus

96. Apex of fifth protarsomere with entire membrane encasing ungues (0), or with internal, longitudinal slit (1).

97. Apex of fifth meso- and metatarsomere with entire membrane encasing ungues (0), or with medial, longitudinal slit (1).

98. Inner, median surface of the fifth protarsomere in male lacking median projection (0), with one median projection (1), or with one anterior and one posterior projections (2).

99. Inner, median surface of the fifth mesotarsomere in male lacking median projection (0), or with one median projection (1).

100. Inner surface of the fifth metatarsomere in male lacking median projection (0), with one median projection (1), or with one anterior and one posterior projection (2).

Appendages: Claws

101. Inner claw of the protarsus in male simple (0), weakly and narrowly split at apex (1), or widely and deeply split (2).

Claws are defined as “simple” if the claw lacks a split apex (although it may be thickened) (0);

weakly and narrowly split if the split is narrower than the bisected portions (1); or widely and deeply split if the split is wider or subequal to the bisected portions (2).

102. Outer claw of mesotarsus in male simple (0), weakly and narrowly split (1), widely and deeply split (2), or split and the lower branch particularly widened (3).

103. Outer claw of metatarsus in male simple (0), weakly and narrowly split (1), or widely and deeply split (2).

104. Outer claw of protarsus, and inner claws of meso- and metatarsi in male simple (0), or split (1).

105. Inner and outer claws of the protarsus in male equal in size (0), inner claws larger than outer claws (1), or inner claws smaller than outer claws (2).

106. Inner and outer claws of the meso- and metatarsus in male equal in size (0), outer claws larger than inner claws (1), or outer claws smaller than inner claws (2).

107. Inner claw of protarsus without basal tooth (0), or with basal tooth (1).

108. Inner claw of protarsus in female simple (0), weakly and narrowly split at apex (1), or widely and deeply split (2).

109. Outer claw of mesotarsus in female simple (0), weakly and narrowly split (1), or widely and deeply split (2).

110. Outer claw of Metatarsus in the female simple (0), weakly, narrowly split (1), or widely and deeply split (2).

111. Outer claw of protarsus and inner claws of meso- and metatarsi in female simple (0), or split

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112. Inner and outer claws of protarsus in female equal in size (0), inner claws larger than outer claws (1), or inner claws smaller than outer claws (2).

113. Inner and outer claws of meso- and metatarsus in female equal in size (0), outer claws larger than inner claws (1), or outer claws smaller than inner claws (2).

Appendages: Unguitractor Plate

114. Unguitractor plate of the protarsus cylindrical (0), plate-like at base and cylindrical at apex (1), or plate-like and triangular (2).

115. Unguitractor plate of the protarsus exposed beyond base of claws (0), or hidden beyond base (1).

116. Unguitractor plate of the meso- and metatarsi cylindrical (0), plate-like at base and cylindrical at apex (1), or plate-like and triangular (2).

117. Empodium of meso- and metatarsi with two setae (0), with three setae (1), with many setae (2), or with a seta (3).

Venter: Prosternum

118. Prosternal projection produced to trochanter (0), produced half-way to trochanter (1), or not appreciably produced (2).

119. Prosternal projection in the ventral area not developed (0), weakly developed (1), roundly swollen (2), or triangularly and entirely flat (3).

120. Prosternal projection in the apical triangular (0), rounded (1), bi-dentate (2), or entirely flat (3).

121. Prosternal projection in the ventral area smooth (0), furnished with setae (1).

122. Preprosternum wide (0), or narrow (1).

123. Preprosternum with one stout ridge in the middle (0), or concave and with two ridges in the middle (1).

124. Prosternum not appreciably produced at the middle (0), or protruded anteriad (1).

Venter: Proepisternum

125. Anterior angle of proepisternum not produced (0), or produced (1).

126. Suture between the proepisternum and anterio-medial prosternum poorly defined (indicated by a line) (0), or well defined and ridge-like (1).

Venter: Meso- and Metasternum

127. Mesosternum not produced (0), weakly produced, not surpassing base of mesocoxae (1), or produced anteriorly beyond base of mesocoxae (2).

128. Mesosternum not appreciably produced beyond mesometasternal suture (0), or appreciably produced (1).

129. Metasternum not produced (0), produced to the apex of mesotrochanter (1), produced to the apex of mesocoxa (2), or produced to the base of prosternal projection (3).

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130. Metasternum in male in lateral view flat or weakly recurved (0), or recurved (1).

131. Mesosternal suture well delineated (0), or poorly delineated (1).

Venter: Abdominal sternites

132. Sternite 2 flattened (0), weakly cariniform anteriad and flattened posteriad (1), or strongly cariniform at the middle (2).

133. Sternites 2-5 subequal in length at the middle and at the sides (0), or shorter in length at the middle than at the sides, thus causing the sternites to be greatly concave (1).

Metanotum and Metendosternite

134. Metendosternite T-shaped (0), or Y-shaped (1).

Male Genitalia

135. Phallobase in lateral to ventral portions entirely sclerotized (0), phallobase in lateral to ventral portions membranous (1), or phallobase entirely membranous (2).

136. Parameres symmetrical (0), or asymmetrical (1).

137. Parameres hinged dorso-ventrally (0), laterally (1), or fused (2).

138. Ventral piece of parameres membranous and poorly defined (0), or well-sclerotized and well-defined (1).

139. Basal portion of parameres entirely fused (0), or weakly fused and defined by suture (1).

140. Apical portion of parameres simple (0), or depressed or cleft (1).

141. Ventral portion of parameres glabrous (0), or furnished with setae (1).

142. Parameres not prolonged and less than 1/2 of genitalia in length (0), or prolonged and more than 1/2 of genitalia in length (1).

143. Apical to medial portions of internal sac without setae (0), or with setae (1).

144. Medial to basal portions of internal sac without extremely small spine (0), scarcely with extremely small spines (1), or with many hardened small spine (2), with hardened small and large spines (3), with large spines (4).

145. Basal portion of internal sac without saw-formed plates (0), or with long saw-formed plates (1).

146. Medial portion of internal sac without large spines (0), with a large spine (1), with two large spines (2), or with many large spines (3).

147. Apical portion of internal sac without spines (0), with small spines (1), with medium-sized spines (2), or with large and small spines (3).

148. Medial portion of Internal sac without appendages (0), with a membranous appendage (1), with a thin semicircular appendage (2), with a chitinous fin-shaped appendage (3), with a chitinous, thick fin-shaped appendage bearing a spine at apex (4), or with a chitinous, thick plate-shaped appendages bearing many spines (5).

149. Medial portion of internal sac without spinous plate (0), or with a spine on the plate (1).

150. Basal portion of internal sac without plate-shaped appendages (0), with plate-formed

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appendages (1), or with plate-shaped appendages bearing many spines (2).

151. Spiculum gastrale with branches and associated sclerites separated (0), branches and associated sclerites fused (1), or sclerites entirely lacking (2).

Color

152. Dorsal surface with vitreous lustre (0), or with metallic lustre (1).

3-3. Phylogenetic analysis using Maximum-parsimony method

Maximum-parsimony (MP) analysis was conducted with PAUP 4.0b10 (Swofford 2002) under heuristic searches using a tree-bisection-reconnection (TBR) branch -swapping algorithm, and 10 replicates of a random-addition sequence.

The confidence levels of each branch were estimated by 1000 bootstrap replications (Felsenstein 1985) using 1000 bootstrap replications using heuristic searches with simple addition sequence and nearest-neighbor interchange (NNI) branch-swapping for the MP analysis.

Phylogenetic analysis was conducted the five stages of analysis which is described below.

(1) Phylogenetic position and phylogeny of the genus Parastasia based on all taxa analysis.

In this phylogenetic analysis, all taxa including all species of Parastasia are used and the Holotrichia was used as outgroup (Fig.IV-1).

(2) Phylogeny of the genus Parastasia based on second analysis using Desmonyx and Dynastinae as the out-group.

This phylogenetic analysis, all species of Parastasia are used and treated the Desmonyx and Dynastinae as outgroup (Fig.IV-2).

(3) Phylogeny of the genus Parastasia based on the complete data set analysis

This analysis was performed to be limited the species of the genus Parastasia and the outer group that had the complete data set (Fig.IV-3).

(4) Phylogeny of the genus Parastasia based on Some typical species of the complete data set analysis using Desmonyx and Dynastinae as the outgroup.

This analysis was performed to select the representative species of each clade and glade of the genus Parastasia that had the complete data set (Fig.IV-4).

(5) Phylogeny of the genus Parastasia based on species-group taxa analysis using Desmonyx and Dynastinae as the outgroup

This analysis was conducted using each species groups of the genus Parastasia that was

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shown in the clade or the glade on the basis of the result of first analysis (Figs.IV-5-8).

IV. Results

1. Phylogenetic position and phylogeny of the genus Parastasia based on all taxa analysis The phylogeny of all species and subspecies of the genus Parastasia and closely related Asian genera of the same tribe Rutelini; Ceroplophana, Cyphelytra, Desmonyx, Dicaulocephalus, Fruhstorferia, Kibakoganea, Lutera, Masumotokoganea, Peperonota Rutelarcha, Asian genera of the tribe Anomalini; Anomala, Mimela; Asian genus of the tribe Adoretini; Adoretus and the subfamily Dynastinae consisting of six genera; Blabephorus, Eophileurus, Oryctomorphus, Peltonotus, Pseudohomox, Rutelisca was examined. The out-group used in analysis was the subfamily Melolonthinae consisting of three genera; Polyphylla, Dasylepida, Holotrichia.

Under this situation, the cladistic analysis resulted in 567 equally parsimonious trees (tree length = 1850, consistency index = 0.14, retention index = 0.60, rescaled consistency index = 0.08) and the strict consensus tree was obtained for these shortest trees. The consensus tree was shown in Figs. IV-1. The clade supported by a bootstrap value of ≧50% was indicated above each branch.

The monophyly of the Parastasia clade was supported by a bootstrap proportion (BP) value of 82%.

The result of phylogenetic analysis indicated that the lineage is divided into two branches at the base (Fig.IV-1). One lineage supported by a BP of 73% is the dynastine clade which includes the Parastasia clade, Desmonyx, Oryctomorphus, three genera of Dynastinae (Eophileurus, Blabephorus, Pseudohomonyx) and Peltonotus. The other clade supported by a BP of 70%, which I refer to as the

“rutelinae clade” includes Lutera-Chyphelytra-Rutelarcha clade, Ceroplophana-Dicaulocephalus- Peperonota-Masumotokoganea-Fruhstorferia-Kibakoganea clade, Rutelisca, the Adoretini and the Anomalini clades.

Within the dynastine clade, the genus Peltonotus clde is firstly branched from the other clades, and the remaining branches in order of the Dynastinae clade, the Oryctomorphus clade, the Desmonyx clade, and the Parastasia clade (BP70%).

The Parastasia clade is divided into the following groups; Basal glade, bimaculata clade and major Parastasia clade including canaliculata clade, westwoodi clade, discolor clade, small subclades and many miscellaneous species (Fig.IV-1).

The members of the basal glade are considered to firstly divide species group. However, relationships at the base of the cladogram are most poorly resolved. The grade consists of 19 species and 1 subspecies that include 4 small subclades. The basal members are: P. lutea, P. tenomensis, P.

peterzorni + P. quadrimaculata, P. montrouzieri, P. kraatzi, P. ferrieri (P. ferrieri ferrieri + P.

ferrieri formosana) + P. sawadai, P. dolens, P. pulupuluensis + P. terraereginae + P. confluens, P.

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brevipes, P. bigibbosa, P. oberthueri oberthueri + P. oberthueri ishigakiana, P. sulcata, P. hitomi, P.

christmasensis, and P. glottidion.

The bimaculata clade is a well-defined clade that is sister to the major Parastasia clade. Basal to the higher clade is a well-resolved clade and the members are divided into two subclade. Within the bimaculata clade, the basal members are P. novoguineensis and P. sakaii. The members of first subclade are P. assimilis, P. nigroscutellata, P. nigriceps inconstans, P. nigriceps nigriceps, P.

bimaculata bimaculata, P. bimaculata nicobarica, P. marmorata discophora, P. marmorata marmorata. The members of second subclade are P. exophthalma, P. nigromaculata, P. fujiokai, P.

maluku, P. stella, P. negrosensis, P. percheroni, P. takeshii.

The major Parastasia clade includes following 3 clades, 6 small subclades and miscellaneous species.

The canaliculata clade is a typical group representing the genus Parastasia and relationships are well resolved within the major Parastasia clade. Members of the canaliculata clade are: P.

bicolor, P. basalis, P. cingala, P. dalatina, P. yukioi, P. andamanica, P. polita, P. dimidiata, P.

jamsonae, P. mitsumata, P. takahikoi, P. kangeanensis, P. ephippium, P. rufopicta, P. binotata, P.

birmana, P. canaliculata, P. klossi, P. sumbawana, P. timorensis.

The westwoodi clade is a well-defined clade comprising: P. weberi, P. anomala, P. intermedia, P. indica, P. incurva, P. burmeisteri, P. duchoni, P. gestroi, P. nigripennis, P. marginata, P.

vietnumensis, P. femorata, P. moultoni, P. punctulata, P. westwoodii, P. asahi, P. yasutoshii, P.

fakfakensis, P. lombokensis. Relationships of the most derived members are resolved.

The discolor clade indicated as a resolved clade included small number of species within the major Parastasia clade comprising: P. discolor discolor, P. discolor scutellaris, P. rufolimbata, P.

discolor mirabilis.

Members of the small subclades and miscellaneous species within the major Parastasia clade are P. coquerelii + P. hainanensis, P. kinibalensis + P. kolakana, P. kuijteni + P. sulcipennis, P.

laratina + P. vittata, P. dempuensis + P. masumotoi, P. melanocephala + P. melanocephaloides, P.

aberrans, P. alternate, P. carsteni, P. circumferens, P. diversipennis, P. helleri, P. isidai, P. lobata, P. moseri, P. pascoei, P. rufonigra, P. selangorica, P. wallacea and P. xanthopyga.

2. Phylogeny of the genus Parastasia based on second analysis using Desmonyx, Oryctomorphus and Dynastinae as the outgroup

Secondary, all species were analyzed again to examine the phylogenetic relationships among 108 taxa in Parastasia clade by using the genus Desmonyx and the subfamily Dynastinae except genus Rutelisca as the out-group. Under this situation, the cladistic analysis resulted in 535 equally parsimonious trees (tree length = 1270, consistency index = 0.1614, retention index = 0.5586, rescaled consistency index = 0.0902) and the strict consensus tree was obtained for these shortest

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trees. The results of the phylogenetic analysis were given by the consensus of 535 trees was shown in Fig. IV-2. The clade supported by a bootstrap value of ≧50% was indicated above each branch.

The monophyly of the Parastasia clade was supported by a bootstrap proportion (BP) value of 72%.

The results indicated almost the similar clade among 108 taxa in Parastasia clade compaired with the phylogenetic position of the genus Parastasia based on all taxa analysis (Fig.IV-1). The different point of the results compared with the results of the previous analysis is the below.

The Parastasia clade is divided into the following groups: Basal glade, bimaculata clade, discolor clade and major Parastasia clade including westwoodi clade and canaliculata clade.

The basal glade consists of one small subclade and some miscellaneous species, and indicated clear unity compaired with the previous analysis.

The bimaculata clade is firstly branched from the basal grade and consists of two subclades and many miscellaneous species (Fig.IV-2). The first subclade such as bimaculata + marmorata subclade included in only 2 species: P. bimaculata bimaculata, P. bimaculata nicobarica, P. marmorata discophora, P. marmorata marmorata. The second subclade newly included some species: P. xanthopyga, P. christmasensis, P. hitomi, P. sulcata, P. oberthueri ishigakiana + P. oberthueri oberthueri and P. lobata.

The discolor clade is secondly branched from the bimaculata clade and formed clearly. The members of the discolor clade are P. carsteni, P. helleri, P. pascoei, P. wallacea, P. discolor discolor + P. discolor mirabilis + P. discolor scutellaris + P. rufolimbata, P. kinibalensis + P.

kolakana, P. isidai, P. diversipennis, P. laratina, P. vittata, P. hainanensis.Two subclades and one species of major Parastasia clade in the previous analysis were formed basal position in the discolor clade: P. selangorica, P. dempuensis + P. masumotoi and P. melanocephala + P.

melanocephaloides.

The major Parastasia clade formed from canaliculata clade and westwoodi clade.

The canaliculata clade indicated clear the relationships within the major Parastasia clade.

one species between the basal glade and the bimaculata clade in the previous analysis and four miscellaneous species of major Parastasia clade in the previous analysis were formed in the basal position of the canaliculata clade: P. glottidion, P. coquerelii, P. moseri, P. kuijteni + P.

sulcipennis.

The westwoodi clade indicated clear the relationships within the major Parastasia clade.

One species of the bimaculata clade in the previous analysis and three miscellaneous species of major Parastasia clade in the previous analysis were formed in the basal position of westwoodi clade: P. novoguineensis, P. alternate + P. rufonigra and P. aberrans.

3. Phylogeny of the genus Parastasia based on the complete data set analysis

A thirdly analysis was conducted to examine the many polytomies within the genus

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Parastasia and to clarify the divergence order in Parastasia clade. The phylogeny of 106 species consisted of 72 species in the Parastasia and 34 species in the out-group was examined after excluding 42 species that has unknown data. Under this situation, the cladistic analysis resulted in 278 equally parsimonious trees (tree length = 1410, consistency index = 0.1809, retention index = 0.6297, rescaled consistency index = 0.1139) and the strict consensus tree was obtained for these shortest trees. The results of the phylogenetic analysis are indicated by the strict consensus of 278 trees and shown in Fig. IV-3. The clade supported by a bootstrap value of ≧ 50% was indicated above each branch. The monophyly of the Parastasia clade was supported by a bootstrap proportion (BP) value of 82 %.

The results indicated almost the similar clade among 69 taxa in Parastasia clade compaired with the phylogenetic position of the genus Parastasia based on all taxa analysis.

The different point of the results compared with the results of the previous analysis is the below.

The Parastasia clade is divided into the following two groups: Basal glade and major Parastasia clade that were formed in westwoodi clade, canaliculata clade, discolor clade, bimaculata clade, two small subclades and 10 miscellaneous species. The phylogenetic position of the bimaculata clade was newly formed in the major Parastasia clade.

The basal glade is formed with P. dolens, P. montrouzieri, P. lutea, P. confluens, P.

kraatzi, P. ferrieri ferrieri + P. ferrieri formosana, P. quadrimaculata. P. brevipes that was included in the basal glade by previous analysis situated in the major Parastasia clade.

The bimaculata clade was consisted of one small subclade and 6 species; P. bimaculata bimaculata + P. bimaculata nicobarica + P. marmorata discophora + P. marmorata marmorata, P. nigriceps inconstans, P. nigriceps nigriceps, P. negrosensis, P. nigromaculata, P.

nigroscutellata and P. percheroni.

The discolor clade was consisted of one small subclade and 5 species; P. discolor discolor + P. discolor scutellaris + P. diversipennis + P. isidai + P. kolakana, P. helleri, P.

laratina, P. pascoi, P. vittata and P. wallacea.

The canaliculata clade was consisted of 12 species; P. basalis, P. cingala, P. binotata, P.

ephippium, P. rufopicta, P. birmana, P. canaliculata canaliculata, P. mitsumata, P. takahikoi, P.

klossi, P. circumferens and P. coquerelii.

The westwoodi clade was consisted of 14 species; P. anomala, P. asahi, P. marginata, P.

burmeisteri, P. lombokensis, P. yasutoshii, P. incurve, P. indica, P. duchoni, P. gestroi, P.

nigripennis, P. westwoodii, P. femorata, P. moultoni and P. punctulata.

The members of two small subclade and 10 miscellaneous species in the major Parastasia clade are the following species: P. melanocephala + P. melanocephaloides, P.

oberthueri oberthueri + P. oberthueri ishigakiana + P. hitomi, P. aberrans, P. alternate, P.

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andamanica, P. timorensis, P. moseri, P. moseri, P. sulcipennis, P. novoguineensis, P.

dimidiata, P. brevipes and P. fujiokai.

4. Phylogeny of the genus Parastasia based on some typical species of the complete data set analysis using Desmonyx, Dynastinae and Melolonthinae as the outgroup

A fourth analysis was conducted to examine the phylogenetic position within major Parastasia clade. Some typical species selected from each major Parastasia clade; P.

canaliculata, P. cingala, P. ephippium and P. circumferens from the canaliculata clade, P.

indica, P. moultoni, P. westwoodii and P. yasutoshii from the westwoodi clade, P. discolor discolor, P. diversipennis, P. wallacea and P. vittata from the discolor clade and P. nigriceps nigriceps, P. bimaculata bimaculata, P. nigromaculata and P. fujiokai from the bimaculata clade. The species of the basal glade selected the below species; P. quadrimaculata, P. ferrieri ferrieri, P. montrouzieri, P. lutea, P. kraatzi, P. confluens, P. oberthueri ishigakiana, P. dolens and P. brevipes. The species of the out-group was selected and consisted of 5 species;

Polyphylla albolineata, Dasylepida fissa, Holotrichia sp., Desmonyx humeralis and Oryctomorphus bimaculatus.

Under this situation, the cladistic analysis resulted in 18 equally parsimonious trees (tree length = 502, consistency index = 0.3984, retention index = 0.5459, rescaled consistency index

= 0.2175) and the strict consensus tree was obtained for these shortest trees. The strict consensus tree was shown in Fig. IV-4. The clade supported by a bootstrap value of ≧ 50%

was indicated above each branch. The monophyly of the Parastasia clade was supported by a bootstrap proportion (BP) value of 77%.

As a result of the analysis, the basal glade did not form a clade and was consisted of 6 species. In these species, two species were sister to Desmonyx humeralis; P. quadrimaculata, P.

ferrieri ferrieri. And four species were next to the two species, such as P. montrouzieri, P. lutea, P. kraatzi and P. confluens.

The major Parastasia clade was consisted of three clades and 7 miscellaneous species.

The three clades are canaliculata clade, westwoodi clade and discolor clade.

The members of the bimaculata clade were included in the major Parastasia clade, but were not formed the bimaculata clade. The westwoodi clade and the canaliculata clade form the sister group, but the discolor clade is formed independently. Four miscellaneous species of the major Parastasia clade are included in the bimaculata clade by the results of the previous analysis; P.

bimaculata bimaculata, P. nigriceps nigriceps, P. nigromaculata and P. fujiokai. The rest species within the major clade were boundary species between the bimaculata clade and the basal glade by the result of first analysis, such as P. oberthueri ishigakiana, P. dolens and P.

brevipes.

ヨツバコガネ属の系統分類学的研究（鞘翅目，コガ ネムシ科）

ヨツバコガネ属の系統分類学的研究（鞘翅目，コガ ネムシ科）

Systematic study of the genus Parastasia Westwood, 1842 (Coleoptera, Scarabaeidae)

ヨツバコガネ属の系統分類学的研究

（鞘翅目, コガネムシ科）

Kaoru WADA

和田 薫

CONTENTS

ヨツバコガネ属の系統分類学的研究（鞘翅目，コガネムシ科）

ヨツバコガネ属の系統分類学的研究（鞘翅目，コガネムシ科）

和田薫