京都大学 / 0048 / 新制・課程博士 / 博士(人間・環境学) / 甲第18358号 / 人博第671号 / 新制||人||161(附属図書館) / 25||人博||671(吉田南総合図書館) / 31216 / 京都大学大学院人間・環境学研究科相関環境学専攻 / (主査)教授 松井 正文, 教授 加藤 眞, 教授 市岡 孝朗 / 学位規則第4条第1項該当 / Doctor of Human and Environmental Studies / Kyoto University / DFAM

Most of the species of the family Rubiaceae with flowers arranged in head inflorescences are currently classified in three distantly related tribes, Naucleeae (subfamily Cinchonoideae) and Morindeae and Schradereae (subfamily Rubioideae). Within Morindeae the type genus Morinda is traditionally and currently circumscribed based on its head inflorescences and syncarpous fruits (syncarps). These characters are also present in some members of its allied genera, raising doubts about the monophyly of Morinda. We perform Bayesian phylogenetic analyses using combined nrETS/nrITS/trnT-F data for 67 Morindeae taxa and five outgroups from the closely related tribes Mitchelleae and Gaertnereae to rigorously test the monophyly of Morinda as currently delimited and assess the phylogenetic value of head inflorescences and syncarps in Morinda and Morindeae and to evaluate generic relationships and limits in Morindeae. Our analyses demonstrate that head inflorescences and syncarps in Morinda and Morindeae are evolutionarily labile. Morinda is highly paraphyletic, unless the genera Coelospermum, Gynochthodes, Pogonolobus, and Sarcopygme are also included. Morindeae comprises four well-supported and morphologically distinct major lineages: Appunia clade, Morinda clade (including Sarcopygme and the lectotype M. royoc), Coelospermum clade (containing Pogonolobus and Morinda reticulata), and Gynochthodes-Morinda clade. Four possible alternatives for revising generic boundaries are presented to establish monophyletic units. We favor the recognition of the four major lineages of Morindeae as separate genera, because this classification reflects the occurrence of a considerable morphological diversity in the tribe and the phylogenetic and taxonomic distinctness of its newly delimited genera.

Vespidae possui aproximadamente 5000 espécies, sendo 3500 dessas são eumeníneos. Zethus é o maior gênero de Eumeninae, com mais de 250 espécies no mundo. Atualmente, encontra-se dividido em quatro subgêneros: Z. (Zethus), Z. (Zethusculus), Z. (Zethoides) e Z. (Madecazethus). Z. (Zethoides), com 42 espécies, se destaca por construir ninhos expostos e apresentar comportamento subsocial. Essas espécies encontram-se divididas em oito grupos, cada qual uma unidade filogenética natural que, porém, foram criados sem qualquer análise filogenética. Exemplares de 18 espécies de Z. (Zethoides), correspondentes aos grupos foram examinadas junto com terminais de outras linhagens de Zethus, Zethini e Eumeninae para elaboração de uma análise cladística afim de averiguar as divisões elaboradas. Todos os grupos de espécies foram recuperados como monofiléticos, exceto Z. bilgumis, que é parafilético em relação ao grupo Z. clypearis. Z. (Zethoides), apesar de monofilético, junto com Z. (Madecazethus), situa-se dentro de Zethus s. s. Z. (Zethusculus) também não foi recuperado como monofilético. Zethus é parafilético em relação aCtenochilus. Assim, o grupo Z. clypearisnão foi recuperado e seus integrantes pertencem agora ao grupo Z. biglumis, e as seguinte sinonímias serão propostas: Ctenochilus (=Zethus); Z. (Madecazethus) (=Z. (Zethus)); Z. (Zethoides) (=Z. (Zethus)). / Vespidae has approximately 5000 species, of which 3500 are eumenines. Zethus is the largest genus in Eumeninae, with over 250 species. Currently, it is divided in four subgenera: Z. (Zethus), Z. (Zethusculus), Z. (Zethoides) and Z. (Madecazethus). Z. (Zethoides), with 42 species, stands out for building exposed nests and presenting subsocial behavior. These species are further divided in eight groups, each considered a phylogenetic unit that were created without any phylogenetic analysis. Eighteen species of Z. (Zethoides) corresponding to different groups were examined, altogether with terminals fromdistict lineages of Zethus, Zethini and Eumenine, in order to elaborate a cladistics analysis to verify the proposed divisions. All species groups were monophyletic except for the Z. biglumis group, that is paraphyletic in relation to the Z. clypearis group. Z. (Zethoides), although monophyletic as Z. (Madecazethus), is placed within Zethus s. s. Z. (Zethusculus) was also retrieved paraphyletic. Thus, the Z. clypearis group is dismantled, and its integrates are incorporated in the Z. biglumis group. Also, the following synonymies will be proposed: Ctenochilus (=Zethus); Z. (Madecazethus) (=Z. (Zethus)); Z. (Zethoides) (=Z. (Zethus)).

Vespidae possui aproximadamente 5000 espécies, sendo 3500 dessas são eumeníneos. Zethus é o maior gênero de Eumeninae, com mais de 250 espécies no mundo. Atualmente, encontra-se dividido em quatro subgêneros: Z. (Zethus), Z. (Zethusculus), Z. (Zethoides) e Z. (Madecazethus). Z. (Zethoides), com 42 espécies, se destaca por construir ninhos expostos e apresentar comportamento subsocial. Essas espécies encontram-se divididas em oito grupos, cada qual uma unidade filogenética natural que, porém, foram criados sem qualquer análise filogenética. Exemplares de 18 espécies de Z. (Zethoides), correspondentes aos grupos foram examinadas junto com terminais de outras linhagens de Zethus, Zethini e Eumeninae para elaboração de uma análise cladística afim de averiguar as divisões elaboradas. Todos os grupos de espécies foram recuperados como monofiléticos, exceto Z. bilgumis, que é parafilético em relação ao grupo Z. clypearis. Z. (Zethoides), apesar de monofilético, junto com Z. (Madecazethus), situa-se dentro de Zethus s. s. Z. (Zethusculus) também não foi recuperado como monofilético. Zethus é parafilético em relação aCtenochilus. Assim, o grupo Z. clypearisnão foi recuperado e seus integrantes pertencem agora ao grupo Z. biglumis, e as seguinte sinonímias serão propostas: Ctenochilus (=Zethus); Z. (Madecazethus) (=Z. (Zethus)); Z. (Zethoides) (=Z. (Zethus)). / Vespidae has approximately 5000 species, of which 3500 are eumenines. Zethus is the largest genus in Eumeninae, with over 250 species. Currently, it is divided in four subgenera: Z. (Zethus), Z. (Zethusculus), Z. (Zethoides) and Z. (Madecazethus). Z. (Zethoides), with 42 species, stands out for building exposed nests and presenting subsocial behavior. These species are further divided in eight groups, each considered a phylogenetic unit that were created without any phylogenetic analysis. Eighteen species of Z. (Zethoides) corresponding to different groups were examined, altogether with terminals fromdistict lineages of Zethus, Zethini and Eumenine, in order to elaborate a cladistics analysis to verify the proposed divisions. All species groups were monophyletic except for the Z. biglumis group, that is paraphyletic in relation to the Z. clypearis group. Z. (Zethoides), although monophyletic as Z. (Madecazethus), is placed within Zethus s. s. Z. (Zethusculus) was also retrieved paraphyletic. Thus, the Z. clypearis group is dismantled, and its integrates are incorporated in the Z. biglumis group. Also, the following synonymies will be proposed: Ctenochilus (=Zethus); Z. (Madecazethus) (=Z. (Zethus)); Z. (Zethoides) (=Z. (Zethus)).

Die zu den Sirenia, oder Seekühen, zählende Gattung Halitherium ist mit Arten aus dem Obereozän bis Untermiozän bekannt. Obwohl Halitherium als monophyletisch angesehen wird, bestätigen alle bisherigen phylogenetischen Analysen die Paraphylie dieser Gruppe. Auch die auf Halitherium basierende nur fossil bekannte Unterfamilie Halitheriinae ist paraphyletisch und umfasst wiederum fast ausnahmslos paraphyletische Gattungen. Der Fokus liegt auf der Typusart H. schinzii. Deren Holotyp, ein Premolar, wird als undiagnostisch definiert und infolgedessen H. schinzii als nomen dubium eingestuft. Die Neubeschreibung sämtlicher dieser Art zugeordneter Skelettreste liefert neue morphologische Daten. So kann die Hypothese von zwei sympatrisch vorkommenden Morphospezies im Unteroligozän Zentraleuropas auf Basis mehrerer unterscheidender Merkmale gestützt werden. Für die Verwandtschaftsanalyse der „Halitherium“ traditionell zugeordneten Arten und die Ermittlung ihrer phylogenetischen Stellung innerhalb der Ordnung Sirenia finden strenge kladistische Prinzipien Berücksichtigung. Eine revidierte, ergänzte und erweiterte Merkmalsmatrix stellt dabei den bisher größten morphologischen Datensatz über Sirenia dar. Die phylogenetischen Analysen zeigen, dass die „Halitherium“ Arten keine monophyletische Gruppe bilden. Im Zuge dieser systematisch-taxonomischen Revision werden die „Halitheriinae“ eingezogen und vier neue Gattungen aufgestellt. Des Weiteren wird eine neue Klassifikation der Sirenia vorgeschlagen, in der eine konsequente Unterscheidung zwischen einer paraphyletischen Stammgruppe und einer monophyletischen Kronengruppe Anwendung findet. Diese Studie liefert neue Daten über die Diversität und Biogeographie von Sirenen. Die herausragendsten Ergebnisse sind zum einen die Revision einer der zweifelhaftesten Sirenia Gruppen, die „Halitheriinae“. Zum anderen wird für den Ursprung der Kronengruppensirenen ein eher unteroligozäner statt eozäner Zeitpunkt postuliert. / The genus Halitherium includes a number of fossil sirenian species, or sea cows, ranging from the late Eocene to early Miocene. Although Halitherium is assumed to be monophyletic, all previous phylogenetic analyses reveal this group to be paraphyletic. As such, the exclusively extinct subfamily Halitheriinae based on Halitherium is paraphyletic comprising mainly genera that are invariably paraphyletic as well. The focus lies on the type species H. schinzii and the morphological basis for its establishment. The holotype, a single premolar, is considered non-diagnostic, which resulted in the recognition of this taxon name as a nomen dubium. Abundant skeletal material originally assigned to “H. schinzii” is re-described providing new data on the morphology of this sirenian. In this process, the hypothesis of two sympatric morphospecies in the lower Oligocene of Central Europe is corroborated by a suite of distinguishing characters. For the analysis of the interrelationships of the species traditionally assigned to “Halitherium”, and the identification of their phylogenetic position within the order Sirenia, robust cladistic principles are applied. A revised, supplemented and extended data matrix represents the hitherto largest data set on Sirenia based on morphological characters. The phylogenetic analyses show that the “Halitherium” species do not form a monophyletic group. In the course of this systematic and taxonomic revision the “Halitheriinae” are refuted and four new genera are established. Furthermore, a new systematic framework is introduced for Sirenia primarily distinguishing between a paraphyletic stem group and a monophyletic crown group. This study provides new data on the past sirenian diversity and biogeography. The most important results are that one of the most disputed sirenian groups, the “Halitheriinae”, is revised, and that the divergence time of crown group sirenians is estimated as early Oligocene rather than Eocene.

The evolutionary origin and the phylogeny of higher metazoan taxa is still under debate although considerable progress has been made in the past 20 years. Metazoa represents a monophyletic group of highly diverse animals including Bilateria, Cnidaria, Porifera, Ctenophores, and Placozoa. Bilateria comprises the majority of metazoans and consists of three major clades: Deuterostomia, Spiralia (= Lophotrochozoa sensu lato), and Ecdysozoa, whereas the sister group taxa Spiralia and Ecdyzozoa form the monophyletic clade Protostomia. Molecular data have profoundly changed the view of the bilaterian tree of life. One of the main questions concerning bilaterian phylogeny is the on-going debate about the evolution of complexity in Bilateria. It was assumed that the last common ancestor of Deuterostomia, Ecdysozoa and Spiralia had a segmented and coelomate body organization resembling that of an annelid. On the contrary, the traditional view is the evolution of Bilateria from a simple body organization towards more complex forms, assuming that the last common ancestor of Bilateria resembles a platyhelminth-like animal without coelomic cavities and segmentation. To resolve this question, it is necessary to unravel the phylogenetic relationships within Bilateria. By using mitogenomic and phylogenomic data, this thesis had a major contribution to clarify phylogenetic relationships within problematic metazoan taxa: (1) the phylogeny of Deuterostomia, (2) the questionable monophyly of Platyzoa, and first assumptions concerning the phylogeny of Gnathostomulida, Gastrotricha and Polycladida, (3) phylogenetic relationships within annelid taxa, especially Terebelliformia, Diurodrilidae, and Syllidae, with new insights into the evolution of mitochondrial gene order, and (4) new insights into the evolution of annelids, especially the interstitial ones, as well as the colonization of the interstitial realm.