Taxonomy and Phylogeny of Cotylea (Platyhelminthes: Polycladida) from Japan

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Taxonomy and Phylogeny of Cotylea (Platyhelminthes: Polycladida) from Japan

露木, 葵唯

北海道大学. 博士(理学) 甲第15286号

2023-03-23

10.14943/doctoral.k15286

http://hdl.handle.net/2115/89609

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theses (doctoral)

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Hokkaido University Collection of Scholarly and Academic Papers : HUSCAP

Taxonomy and Phylogeny of Cotylea
(Platyhelminthes: Polycladida) from Japan
日本産吸盤亜目ヒラムシ類（扁形動物門：多岐腸目）の系統分類学的研究

Aoi Tsuyuki

A Ph.D. dissertation presented

to

Department of Natural History Sciences,
Graduate School of Science,
Hokkaido University,
Sapporo
060-0810

in
March 2023

This dissertation is based on the following papers:
Chapter II-1
1. Tsuyuki A, Oya Y, Kajihara H. 2022b. Reversible shifts between interstitial and
epibenthic habitats in evolutionary history: molecular phylogeny of the marine
flatworm family Boniniidae (Platyhelminthes: Polycladida: Cotylea) with
descriptions of two new species. PLoS ONE 17: e0276847.
Chapter II-3
2. Tsuyuki A, Oya Y, Jimi N, Kajihara H. 2020a. Description of Pericelis
flavomarginata sp. nov. (Polycladida: Cotylea) and its predatory behavior on a
scaleworm. Zootaxa 4894(3): 403–412.
3. Tsuyuki A, Oya Y, Kajihara H. 2022c. Two new species of the marine flatworm
Pericelis (Platyhelminthes: Polycladida) from southwestern Japan, with an
amendment of the generic diagnosis based on phylogenetic inference. Marine
Biology Research 17(9–10): 946–959.
Chapter II-4
4. Tsuyuki A, Oya Y, Kajihara H. 2020b. First record of Stylostomum ellipse (Dalyell,
1853) (Platyhelminthes, Polycladida) from the Pacific Ocean. Check List 16(3):
773–779.
Chapter II-5
5. Tsuyuki A, Oya Y, Kajihara H. 2019. A new species of Prosthiostomum
(Platyhelminthes: Polycladida) from Shirahama, Japan. Species Diversity 24(2):
137–143.
6. Tsuyuki A, Kajihara H. 2020. A giant new species of Enchiridium (Polycladida,
Prosthiostomidae) from southwestern Japan. ZooKeys 918: 15–28.
ii

7. Tsuyuki A, Kohtsuka H, Kajihara H. 2021. Description of a new species of the
marine flatworm Prosthiostomum (Platyhelminthes: Polycladida) and its three
known congeners from Misaki, Japan, with inference of their phylogenetic
positions within Prosthiostomidae. Zoological Studies 60(29): 1–20.
Chapter II-6
8. Tsuyuki A, Kohtsuka H, Hookabe N, Kajihara H. 2022a. First record of Bulaceros
porcellanus Newman & Cannon, 1996 (Platyhelminthes, Polycladida, Cotylea)
from Japanese waters, with a revision of the generic diagnosis based on
morphology and molecular phylogeny. Plankton and Benthos Research 17(2): 147–
155.

iii

ACKNOWLEDGEMENTS
I am deeply grateful to my supervisor Prof. Hiroshi Kajihara for his help and guidance
during my master’s and doctoral research. Dr. Keiichi Kakui is acknowledged for his
useful advice in my research. I also thank Prof. Kazuhiro Kogame, Prof. Hiroshi
Kajihara, and Dr. Keiichi Kakui for reviewing this thesis and providing valuable
comments.
I deeply thank Dr. Yuki Oya for the helpful guidance and supports of polyclad
taxonomy in Japan.
I have been helped by innumerable people in sampling and collecting
specimens. I am thankful to Dr. Naoto Jimi, Dr. Yuki Oya, Mr. Hisanori Kohtsuka, Dr.
Daisuke Uyeno, Dr. Shinta Fujimoto, Ms. Natsumi Hookabe, Mr. Naohiro Hasegawa,
Prof. Toru Miura, Dr. Kohei Ogushi, Dr. Hiroki Kise, Ms. Midori Matsuoka, Dr. Seiji
Arakaki, Prof. Mutsunori Tokeshi, Dr. James Reimer, Dr. Gaëlle Quéré, Dr. Angelo
Poliseno, Dr. Yuka Kushida, Dr. Guin Yee Soong, Dr. Takuma Fujii, Dr. Shinri Tomioka,
Mr. Haruki Ishiyama, Mr. Shoki Shiraki, Prof. Masato Kiyomoto, Dr. Hiroaki Nakano,
Ms. Mayuko Nakamura and the staffs at Akkeshi Marine Station, Misaki Marine
Biological Station, Noto Marine Laboratory, Oshoro Marine Station, Seto Marine
Biological Laboratory, Kuroshio Biology Research Institute, Ishigaki Sensui Do for
assistance in sampling.
I am also grateful to Prof. Marco Curini-Galletti, Prof. Sigmer Quiroga, and Dr.
Christopher Edward Laumer for providing helpful information. Dr. Carolina Noreña,
Dr. Daniel Marquina, Mr. Jorge Rodriguez Monter, Dr. Stephen Keable, Dr. Geoff
Thompson, Dr. Mal Bryant, Dr. Sue-Ann Watson, Dr. Robyn Cumming, and Dr. Terry

iv

Miller are acknowledged for arrangement of the histological photomicrographs of the
type series.
This study was supported by the Research Institute of Marine Invertebrates (FY
2019), Kuroshio Biology Research Institute (FY 2018 for Aoi Tsuyuki), Japan Science
and Technology Agency SPRING (No. JPMJSP2119), and Japan Society for the
Promotion of Science (JSPS) under KAKENHI grant number 22J13246, and the
Japanese Association for Marine Biology (JAMBIO). I am deeply grateful to financial
support by JST and JSPS. Special thanks to my family, friends, and Biodiversity 1
members for kind supports.

v

TABLE OF CONTENTS
ACKNOWLEDGEMENTS ............................................................................................ iv
I. GENERAL INTRODUCTION .................................................................................... 1
II. TAXONOMIC AND PHYLOGENETIC STUDIES OF COTYLEA IN JAPAN ..... 13
II-1. BONINIIDAE BOCK, 1923................................................................................... 13
Introduction .................................................................................................................... 13
Material and methods ..................................................................................................... 15
Results ............................................................................................................................ 19
Taxonomy .................................................................................................................. 19
1. Genus Boninia Bock, 1923 ............................................................................... 19
1. Boninia uru Tsuyuki, Oya, and Kajihara, 2022b ......................................... 19
2. Boninia yambarensis Tsuyuki, Oya, and Kajihara, 2022b .......................... 24
Molecular phylogeny ................................................................................................. 27
Ancestral habitats ....................................................................................................... 28
Discussion....................................................................................................................... 28
II-2. CESTOPLANIDAE LANG, 1884 ......................................................................... 32
Introduction .................................................................................................................... 32
Material and methods ..................................................................................................... 33
Results ............................................................................................................................ 35
Taxonomy .................................................................................................................. 35
2. Genus Eucestoplana Lang, 1884....................................................................... 35
3. Eucestoplana cf. cuneata (Sopott-Ehlers and Schmidt, 1975).................... 35
4. Eucestoplana sp. .......................................................................................... 37
Molecular analyses .................................................................................................... 40
vi

Discussion....................................................................................................................... 41
II-3. DIPOSTHIDAE Woodworth, 1898 ........................................................................ 43
Introduction .................................................................................................................... 43
Material and methods ..................................................................................................... 45
Results ............................................................................................................................ 47
Taxonomy .................................................................................................................. 47
3. Genus Pericelis Laidlaw, 1902 ......................................................................... 47
5. Pericelis flavomarginata Tsuyuki, Oya, Jimi, and Kajihara, 2020a ............ 48
6. Pericelis lactea Tsuyuki, Oya, and Kajihara, 2022c .................................... 52
7. Pericelis maculosa Tsuyuki, Oya, and Kajihara, 2022c .............................. 55
Molecular phylogeny ................................................................................................. 58
Discussion....................................................................................................................... 59
II-4. EURYLEPTIDAE Stimpson, 1857 ........................................................................ 61
Introduction .................................................................................................................... 61
Material and methods ..................................................................................................... 62
Results ............................................................................................................................ 63
Taxonomy .................................................................................................................. 63
4. Genus Stylostomum Lang, 1884 ........................................................................ 63
8. Stylostomum ellipse (Dalyell, 1853) ............................................................ 63
Discussion....................................................................................................................... 67
II-5. PROSTHIOSTOMIDAE LANG, 1884 ................................................................ ...

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