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Lumping three nominal species into one: taxonomic revision of amphibian parasitic leeches of Torix in Far East Asia (Hirudinea: Glossiphoniidae)

Kambayashi, Chiaki Nakano, Takafumi 京都大学 DOI:10.1071/IS23042

2023.12

概要

Although most members of the freshwater leech family Glossiphoniidae have mid-body somites divided into three annuli, the genus Torix Blanchard, 1893 is distinguished by two-annuli somites. Torix has high species richness in Far East Asia, and three nominal species have been recognised in the Japanese Archipelago and adjacent regions that can be distinguished by a combination of both internal and external morphological characteristics. However, recent studies have shown that these diagnostic features are ontogenetically variable and this has resulted in taxonomic confusion among Torix species endemic to the Japanese Archipelago. In this study, we revisit the taxonomic accounts of T. orientalis (Oka, 1925) and T. tagoi (Oka, 1925), in addition to that of the recently redescribed T. tukubana (Oka, 1935) to clarify the diagnostic characteristics for each of the three species. Our morphological and molecular phylogenetic analyses demonstrated that the three Torix species in Japan are indistinguishable. We therefore conclude that these species should be synonymised and treated as a single species. The specific names orientalis and tagoi were simultaneously established under the genus Oligobdella Moore, 1918, therefore we acted as First Reviser and gave precedence to the name O. tagoi, thus the valid name for the Far East Asian Torix species is T. tagoi unless T. orientalis and T. tagoi are treated as distinct species. There are several type localities for T. tagoi and the name-bearing types have been lost, therefore we designate a neotype for this species to obviate zoological and nomenclatural issues.

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参考文献

COI

18S

ITS

KUZ Z4975

LC776047*

LC776025*

LC776026*

This study

KUZ Z4976

LC776048*

This study

KUZ Z4977

LC776049*

This study

KUZ Z4978

LC776050*

This study

Ingroup (= Torix tagoi)

(Topotype of T. orientalis)

Nemuro City, Hokkaido

Akkeshi-cho, Hokkaido

KUZ Z4979

LC776051*

LC776027*

This study

Samani-gun, Hokkaido

KUZ Z4980

LC776052*

LC776028*

This study

Yubari City, Hokkaido

KUZ Z4981

LC776053*

LC776029*

This study

Tomakomai City, Hokkaido

KUZ Z4982

LC776054*

LC776030*

This study

KUZ Z4983

LC776055*

This study

Oshamanbe-cho, Hokkaido

KUZ Z4984

LC776056*

LC776031*

This study

Ajigasawa-machi, Aomori Pref.

KUZ Z4985

LC776057*

This study

Sendai City, Miyagi Pref.

KUZ Z4986

Nagai City, Yamagata Pref.

KUZ Z4987

Tamura City, Fukushima Pref.

KUZ Z4988

LC776058*

LC776032*

This study

(Topotype of T. tukubana)

(Topotype of T. orientalis)

(Neotype of T. tagoi)

Tsukuba City, Ibaraki Pref.

Yumoto, Nikko City, Tochigi Pref.

LC538265

LC538285

LC776033*

Kambayashi et al. (2020); This study

KUZ Z4989

LC776059*

This study

KUZ Z4990

LC776060*

This study

KUZ Z4991

LC776061*

This study

KUZ Z4992

LC776062*

LC776034*

This study

KUZ Z4993

LC776063*

This study

Nagano City, Nagano Pref.

KUZ Z4994

Itoigawa City, Niigata Pref.

KUZ Z4995

LC776064*

LC776035*

This study

Nakatsugawa City, Gifu Pref.

KUZ Z4996

LC776065*

LC776036*

This study

Takayama City, Gifu Pref.

KUZ Z4997

LC776066*

LC776024*

LC776037*

This study

KUZ Z4998

LC776067*

This study

KUZ Z4999

LC776068*

This study

KUZ Z5000

LC776069*

This study

(Topotype of T. tagoi)

(Topotype of T. orientalis)

KUZ Z2972

Taga-cho, Shiga Pref.

KUZ Z5001

LC776070*

LC776038*

This study

Akame, Nabari City, Mie Pref.

KUZ Z5002

LC776071*

LC776022*

LC776039*

This study

KUZ Z5003

LC776072*

This study

KUZ Z5004

LC776073*

This study

KUZ Z5005

LC776074*

This study

Nantan City, Kyoto Pref.

KUZ Z5006

LC776075*

This study

Toyooka City, Hyogo Pref.

KUZ Z5007

LC776076*

LC776040*

This study

Katsuta-gun, Okayama Pref.

KUZ Z5008

LC776077*

LC776041*

This study

Saihaku-gun, Tottori Pref.

TRPM-AO-

LC776078*

LC776042*

This study

0000256

(Identified as T. tukubana)

Miyoshi City, Hiroshima Pref.

KUZ Z5010

LC776079*

LC776043*

This study

Hatsukaichi City, Hiroshima Pref.

KUZ Z5011

LC776080*

LC776023*

LC776044*

This study

KUZ Z5012

LC776081*

This study

KUZ Z5013

LC776082*

This study

KUZ Z5014

LC776083*

This study

Okinoshima-cho, Shimane Pref.

KUZ Z5015

LC776084*

LC776045*

This study

Mima City, Tokushima Pref.

KUZ Z5016

Kumakougen-cho, Ehime Pref.

KUZ Z5017

LC776085*

LC776046*

This study

Dandong City, Liaoning Province,

OL779256

Zhu et al. (2023)

South Korea

Hir103/4

MN295423

Bolotov et al. (2019)

Primorsky Krai, Russia

Hir101

MN295420

Bolotov et al. (2019)

France

AF003259

Siddall and Burreson (1998)

Myanmar

Hir48/1

MN295394

Bolotov et al. (2019)

China

Outgroup

Hemiclepsis kasmiana

Oka, 1910

Hemiclepsis khankiana

Bolotov, Klass,

Bespalaya, Konopleva,

Kondakov & Vikhrev,

2019

Hemiclepsis marginata

(O. F. Müller, 1773)

Hemiclepsis

myanmariana Bolotov,

Klass, Bespalaya,

Konopleva, Kondakov &

Vikhrev, 2019

Hemiclepsis schrencki

Primorsky Krai, Russia

Hir88/1

MN295415

Bolotov et al. (2019)

Khabarovsk Krai, Russia

Hem1/2

MN295371

Bolotov et al. (2019)

Jingzhou, Hubei Province, China

MN106285

Yi et al. (2022)

Dečani, Kosovo

Gbalc1

LCHME036

Jovanović et al. (2021)

Bolotov, Klass,

Bespalaya, Konopleva,

Kondakov & Vikhrev,

2019

Hemiclepsis tumniniana

Bolotov, Klass,

Bespalaya, Konopleva,

Kondakov & Vikhrev,

2019

Hemiclepsis

yangtzenensis Yang &

Bolotov, 2021

Glossiphonia balcanica

Grosser & Pešić, 2016

Glossiphonia baicalensis

-20

Lake Baikal, Russia

AY047329

Light and Siddall (1999)

Ohrid Lake, North Macedonia

MAC1_1

LCHME020

Jovanović et al. (2021)

(Stschegolew, 1922)

Glossiphonia complanata

(Linnaeus, 1758)

Glossiphonia complanata

-20

Podgorica, Montenegro

MN1_1

LCHME008

-20

Glossiphonia concolor

Mecklenburg-Vorpommern,

(Apathy, 1888)

Germany

Glossiphonia concolor

Ukraine

Glossiphonia elegans

Manitoba, Canada

ROMIZI115

Verrill, 1872

Glossiphonia nebulosa

Gcon1

LCHME041

Jovanović et al. (2021)

KM095097

Kaygorodova and Mandzyak (2014)

MK479253

Mack and Kvist (2019)

LCHME044

Jovanović et al. (2021)

-20

Berlin, Germany

Gnebu4

Kalbe, 1964

Glossiphonia cf. nebulosa

-20

Dečani, Kosovo

Gnebu1

LCHME042

-20

Glossiphonia verrucata

Sweden

AY962458

Siddall et al. (2005)

Croatia

ROMIZI117

MK479263

Mack and Kvist (2019)

MK479264

Mack and Kvist (2019)

MH670835

Kaygorodova et al. (2020)

(Fr. Müller, 1844)

Glossiphonia verrucata

Croatia

ROMIZI117

Glossiphonia sp.

Lake Izumrudnoe, Russia

H56

673

674

Table 2. Uncorrected p-distances for the mitochondrial COI sequences of representative Torix specimens. Location numbers in Table 1

675

are presented in parentheses.

Voucher

1. KUZ Z4976; topotype of T. orientalis (1)

2. KUZ Z4982 (5)

0.026

3. KUZ Z2972; topotype of T. tukubana (11)

0.034

0.042

4. KUZ Z4992; topotype of T. orientalis (12)

0.035

0.043

0.0039

5. KUZ Z4997; neotype of T. tagoi (16)

0.035

0.043

0.030

0.031

6. KUZ Z5002; topotype of T. orientalis (18)

0.034

0.043

0.032

0.035

0.024

7. KUZ Z5011 (24)

0.037

0.052

0.039

0.040

0.035

0.041

8. KUZ Z5017 (27)

0.035

0.043

0.034

0.039

0.019

9. China; identified as T. tukubana (28)

0.048

0.057

0.051

0.042

0.044

676

0.040

677

Table 3. Comparisons of morphological characters between Torix tagoi and three

678

congeneric species.

Character

Torix tagoi

Torix cotylifer

Torix mirus

Torix novaezealandiae

Male gonopore

XI/XII

XI (a1 + a2)/a3

Crop caeca

7 pairs: 1st–6th

7 pairs: 1st pair

6 pairs: 1st pair being very

pairs being

extending to

equal-sized

lateral margin of

small

body

Ovisacs

tubular

globular

679

tubular

680

681

Fig. 1. Map showing the sampling localities of Torix specimens used in this study. Open

682

circles, the square, and triangles indicate the type localities of T. orientalis, T. tukubana,

683

and T. tagoi, respectively. The numbers of localities are shown in Table 1.

684

685

686

Fig. 2. Maximum likelihood tree for 1,267 bp of the mitochondrial COI gene, along with

687

the results of five species-delimitation analyses (ABGD, ASAP, GMYC, bPTP, and

688

mPTP) as indicated by vertical bars. Voucher and location number corresponding to Table

689

1 are presented for Torix. The blue and red shading indicate the eastern (locations #1–19,

690

#21 in Table 1) and western (#20, #22–27) lineages of Japanese Torix, respectively. Open

691

circles, the square, and triangles indicate the topotypes of T. orientalis, T. tukubana, and

692

T. tagoi, respectively. Numbers on nodes represent bootstrap values for maximum

693

likelihood and Bayesian posterior probabilities.

694

695

696

Fig. 3. Statistical parsimony network of nuclear ITS haplotypes. The blue and red circles

697

indicate haplogroups of the eastern and western lineages of Japanese Torix shown in Fig.

698

2, respectively. Black circles indicate the missing haplotypes. The number on each circle

699

corresponds to the location number in Table 1.

700

701

702

Fig. 4. Torix specimens used in this study: (a) dorsal and (b) ventral views of the neotype

703

of T. tagoi (KUZ Z4997); (c) dorsal and (d) ventral views of the topotype of T. orientalis

704

(KUZ Z4975); (e) dorsal and (f) ventral views of a juvenile specimen (KUZ Z4980); (g)

705

dorsal view of the live neotype of T. tagoi; (h) ventral view of live adult individual (KUZ

706

Z5006). Scale bars: 10 mm (a–d, g–h), 1 mm (e–f).

707

708

709

Fig. 5. Torix tagoi, neotype (KUZ Z4997): (a) dorsal view of the digestive tract and male

710

genital organs; (b) dorsal view of the reproductive system and positions of ganglia XI–

711

XVIII. Abbreviations: ac, atrial cornu; agm, anterior ganglionic mass; cc, crop cecum; fg,

712

female gonopore; ic, intestinal cecum; mg, male gonopore; oes, oesophagus; ov, ovisac;

713

pb, proboscis; pcc, post-crop cecum; rt, rectum; scm, salivary cells mass; sd, sperm duct;

714

ts, testisac. Scale bar: 5 mm (a), 1 mm (b).

...

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概要

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Lumping three nominal species into one: taxonomic revision of amphibian parasitic leeches of Torix in Far East Asia (Hirudinea: Glossiphoniidae)

概要

この論文で使われている画像

関連論文

Rediscovery of the presumably extinct fairy lantern Thismia kobensis (Thismiaceae) in Hyogo Prefecture, Japan, with discussions on its taxonomy, evolutionary history, and conservation

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