Fishes of Mongolia

Environment and Social Development
East Asia and Pacific Region
THE WORLD BANK

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Fishes of Mongolia
A check-list of the fishes
known to occur in Mongolia with
comments on systematics and nomenclature
MAURICE KOTTELAT
Fishes of Mongolia
A check-list of the fishes known to occur in Mongolia
with comments on systematics and nomenclature
Maurice Kottelat
September 2006
©2006 The International Bank for Reconstruction and Development/THE WORLD BANK
1818 H Street, NW
Washington, DC 20433 USA
September 2006
All rights reserved.
This report has been funded by The World Bank’s Netherlands-Mongolia Trust Fund for Environmental Reform (NEMO).
Some photographs were obtained during different activities and the author retains all rights over all photographs included in this
report.
Environment and Social Development Unit

East Asia and Pacific Region
World Bank
Washington D.C.
Contact details for author:
Maurice Kottelat
Route de la Baroche 12, Case Postale 57, CH-2952 Cornol, Switzerland.
Email: mkottelat@dplanet.ch
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findings, interpretations, and conclusions expressed in this paper do not necessarily reflect the views of the Executive Directors
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Contents
Contents
Foreword v
Acronyms and Abbreviations vii
Acknowledgments ix
Executive Summary xi
Introduction 1
Methods 3
Species Lists 7
Accounts of Species Recorded from Mongolia 13
Family Petromyzontidae 13
Family Acipenseridae 14
Family Salmonidae 15
Family Coregonidae 18
Family Thymallidae 23
Family Esocidae 26
Family Cyprinidae 27
Family Nemacheilidae 50
Family Cobitidae 56
Family Siluridae 58
Family Lotidae 58
Family Percidae 59
Family Cottidae 59
Family Odontobutidae 61
Unidentifiable Records 61
Accounts of Species Recorded from Adjacent Areas 63
Recommendations 65
Bibliography 67
Appendix 1. Nomenclatural Information 85
Appendix 2. Figures 93
Addendum 103
iii
Foreword
Foreword
ater is an essential element in Mongolia’s development plans; initiatives currently being
W considered include projects such as hydroelectric dams, water transfers, irrigation schemes,
and aquaculture. Assessing the impacts of these projects on freshwater biodiversity is hindered
by inadequate knowledge of the fauna.
The World Bank is supporting Mongolia in its efforts to ensure sustainable exploitation of its considerable
natural resources. This report is the latest product of the Environment and Social Development Department
in the East Asia and Pacific Region of the World Bank, and has been produced within the framework of
the Netherlands-Mongolia Trust Fund for Environmental Reform (NEMO). This wide-reaching initiative
has touched almost all aspects of environmental management in Mongolia in 2005–06. In addition to
this study, we have supported several other studies, such as assessing threats and devising management
needs for a number of animal groups, analyzing the illegal wildlife trade and the illegal timber trade, and
evaluating the success of tree planting projects.
This report establishes a reliable and comprehensive list of Mongolian fish. The work is based on a review
of the existing literature; interviews with local and international experts; examination of material preserved
in natural history museums and research institutes in Beijing, Wuhan, St. Petersburg, Berlin, Stockholm
and Paris; and supplementary fieldwork. Maurice Kottelat has applied his unparalleled knowledge of
the fish of the region to write a critical analysis of the fish fauna of Mongolia. This report represents an
essential foundation of knowledge. Together with the recently published Red List of Mongolian Fish and
the Action Plans for Mongolian Fish (which we also supported), it should provide indispensable material
to support environmental impact assessments for any development project affecting water resources in
Mongolia.
This is the second time such a report has been published by the World Bank. We do so again in recognition
of the foundational role of taxonomy in sustainable development, of the importance of freshwater
biodiversity in the lives of subsistence and commercial fishers, and of the important role that biological
knowledge plays in natural resource planning.
Magda Lovei Arshad M. Sayed

Environment Sector Manager Mongolia Country Manager
East Asia and Pacific Region The World Bank
The World Bank
v
Acronyms and Abbreviations
Acronyms and Abbreviations
CD Coefficient of difference
Code International Code on Zoological Nomenclature
EIA Environmental Impact Assessment
ESC Evolutionary Species Concept
IHB Institute of Hydrobiology, Chinese Academy of Sciences, Wuhan
IZCAS Institute of Zoology, Chinese Academy of Sciences, Beijing
MNHN Muséum National d’Histoire Naturelle, Paris
mtDNA Mitochondrial DNA (Deoxyribonucleic acid)
NEMO Netherlands-Mongolia Trust Fund for Environmental Reform
PCA Principle Component Analysis
SL Standard length
USSR Union of Soviet Socialist Republics
WWF World Wide Fund for Nature
ZISP Zoological Institute, Russian Academy of Sciences, St. Petersburg
ZMB Museum für Naturkunde, Berlin
vii
Acknowledgements
Acknowledgements
am pleased to thank the following persons for their help at different stages of this work. M.
I Erdenebat (Institute of Geo-Ecology, Mongolian Academy of Sciences) assisted in the field and
provided literature, information, and translations. A. Dulmaa (Institute of Biology, MAS) provided
literature and a visit to material under her care. Nina Bogutskaya translated some Russian texts.
E. Zhang translated some Chinese texts. B. Mendsaikhan (Institute of Geo-Ecology, MAS) provided
information and literature. Nina Bogutskaya, Alexander Naseka (Institute of Zoology, St Petersburg),
E. Zhang (Institute of Hydrobiology, Wuhan), Zhang Chun-Guang, Zhao Yahui, Li Gaoyang (Institute
of Zoology, Beijing), Peter Bartsch, and Christa Lamour (Museum für Naturkunde, Berlin) provided
access to material under their care and various museum and library facilities. Nina Bogutskaya read and
commented on the text.
Editorial assistance was provided by Bob Livernash, and the publication process was handled by Bryony
Morgan, who spent many long hours checking and re-checking the text and compiling photographs. I
am pleased to thank M. Erdenebat, Zeb Hogan and Johannes Schöffmann for permission to use their
photographs.
This publication would never have come to light without the efforts, help, tenacity and friendship of Tony
Whitten of the World Bank who planned and organised the work, and made enormous efforts to get all
possible benefit from it.
The study was funded by the Government of the Netherlands through the Netherlands-Mongolia Trust
Fund for Environmental Reform (NEMO).
ix
Executive Summary
Executive Summary
total of 76 native fish species are reliably recorded in Mongolia’s waters. Five of them are possibly
A new to science and unnamed. Five additional species are often reported as being present in
Mongolia but are in fact only presumed to exist and should be deleted from the Mongolian
faunal lists. Four other species are introduced species that have not been sighted for years and
presumably did not become established, and a further two are introduced species which have become
established. Nine species are known from immediately adjacent waters in China and Russia and might be
present, either as permanent inhabitants or vagrant individuals.
The systematic status and nomenclature of all species have been reevaluated. Compared to the last synthesis
of the fishes known from the same area (Baasanjav & Tsendayush, 2001), 11 (15 percent) of the 72
formerly recognized species are invalid, and the names of 28 (39 percent) of the then-known species were
incorrect (either because of misidentifications, or for various nomenclatural reasons). Therefore, in total
more than half (39 out of 72) of the species in this synthesis were incorrectly listed, to which a further
15 species not previously recognized should be added. This clearly demonstrates that present knowledge
of fish diversity in Mongolia is far from adequate, that the number of species is underestimated, and that
more species probably still await discovery. Survey work is needed in addition to an approach of taxonomy
incorporating modern standards, concepts, and procedures.
The fish fauna of the Chinese provinces of Xinjiang and Nei Mongol and of the Russian Tuva and Buryatia
Republics have been compared with the Mongolian fish fauna, where relevant, in an attempt to make the
nomenclatures used in the three countries compatible and in agreement with the International Code of
Zoological Nomenclature. Some of the systematic findings and nomenclatural changes are summarized in
the section containing species lists.
This exercise has revealed a large number of nomenclatural inconsistencies across boundaries, and many
species in Mongolia, China, and Russia which are not yet properly named. There are indications (and in
many cases, firm evidence) that a number of widely distributed “species” in fact are artificial assemblages
of species restricted to a small geographic area. This has immediate implications for resource management
and conservation because species endemic to a small area, or to a single lake or stream, have greater
biodiversity value and thus require closer attention. Also, the transplantation across drainage boundaries
of fish stocks believed to be different populations of a single species may actually be the introduction of a
species into the range of another species, carrying all the risks associated with introductions, including the
risk of replacement of the original species with a hybrid complex.
xi
Introduction
Introduction
his publication results from work conducted in 2005 as part of a consultancy funded by the
T World Bank’s Netherlands-Mongolia Trust Fund for Environmental Reform (NEMO). This
particular project concerned freshwater fish biodiversity in the context of establishing reliable
lists of Mongolian animals, in order to prepare reliable lists of threatened species for which
conservation action is necessary.
This work is based on a review of the existing literature, interviews with local and international experts,
examination of material preserved in natural history museums and research institutes in Beijing, Wuhan,
St. Petersburg, and Berlin (plus others that I had examined before this project in Stockholm and Paris),
and a limited amount of field work (one weekend in central Mongolia).
Although there is a general assumption that the diversity of the Mongolian fish fauna is well known and
there is not much interest in investing time, money, and effort in further taxonomic work, my observations
show that much remains to be done. I found that a number of the Mongolian fish species are still unnamed,
many are misidentified, and that several earlier taxonomic works present problems. Besides, there are a vast
number of nomenclatural problems, most of which cannot be solved without additional extensive data on
the fauna of adjacent areas in Russia, Kazakhstan, China, and Korea.
The problems largely result from different conceptual approaches to taxonomy, the lack of communication
(an obvious result of international politics and linguistic problems, from which I also suffered), and in
part an ignorance or non-respect of international practices or of the International Code of Zoological
Nomenclature.
As an example of the serious limitation of our knowledge of the Mongolian fish fauna and of the extremely
serious need for careful attention, I will cite my limited experience in the field. My schedule and terms of
reference during the first research visit to Mongolia did not allow for actual field work, but I did manage
to spend a weekend in the field. Of course, this could not be far from Ulaanbaatar. One of the two places
I visited was Terelj Nature Reserve, close to the bridge on Tuul River, an area which had probably been
visited by all ichthyologists who have been in Mongolia in the last 50 years.
Field work was not planned when I left for Mongolia and I was therefore totally unequipped for efficient
sampling. Erdenebat M. and I had to resort to primitive tools almost constructed on the spot. Nevertheless,
we collected six species, of which there were identification problems for four. One is locally identified
under the name of a European species (Phoxinus phoxinus), but is clearly a distinct species; one is identified
1
Fishes of Mongolia—A check-list of the fishes known to occur in Mongolia with comments on systematics and nomenclature
as a very variable species (Barbatula toni) with a strong suspicion that several species are confused under
that name; two are species new to science (Barbatula sp. Tuul, Triplophysa sp. Tuul), of which the last one
is the first record of the genus in the Selenge (and Yenisei) drainage. The last two also belong to species
reported to be very variable (Rhynchocypris czekanowskii, Cobitis melanoleuca), which still deserve study
throughout their ranges (all of northern Asia and parts of Europe and northern China).
The purpose of the present check-list is to present an overview of our present knowledge of the diversity,
systematics, and nomenclature of the fishes of Mongolia. It also includes a selected bibliography of the
publications of greatest immediate concern when working on this fauna.
Although two books have already been published on the fishes of Mongolia (Sokolov, 1983; Baasanjav
& Tsendayush, 2001) they are outdated and somewhat misleading. The first task is to ensure that the
nomenclature used in Mongolia, China, Russia, Korea, and in the rest of the world are compatible
and conforms to the International Code of Zoological Nomenclature [Code hereafter] (International
Commission on Zoological Nomenclature, 1999).
Decisions of a strictly scientific nature can be influenced by non-scientific considerations, and this
negatively impacts the efficiency of field work and research. It also interferes with scientific exchanges.
Exchange of material, data and knowledge is necessary for good management of natural resources, and is
beneficial for the country in the long run; if one is able to benefit from the results of research conducted
abroad, there is no need to replicate it and this saves time, effort and money. The present work had to be
done with access to only a limited number of samples and this obviously has limited the conclusions. In
many instances, very little additional work is needed to solve complex problems, but this last step has not
been possible. This is indicated in the text (e.g., under Barbatula compressirostris, Cottus dzungaricus).
The list in this report includes all the fishes which have been recorded in the scientific literature or
observed by myself in Mongolia. Species recorded from within a few kilometres upstream or downstream
of the Mongolian border in China and Russia but not yet known from Mongolia are not included in
the Mongolian fauna, but their distribution and taxonomy is discussed. Records based on the literature
are included but only if they seem reasonably reliable; that is, either there are ways to confirm the
identification from the document itself or from voucher specimens, or the author(s) is (are) known to be
well experienced. Fisheries records have not been taken into consideration since they tend to be unreliable
and/or too superficial for serious biodiversity work. Introduced species are discussed only if they have
established self-sustaining (breeding) populations or are regularly observed.
A very small area of westernmost Mongolia drains to the Irtysh River. There is almost no data on the fishes
of that area. Data on the fishes of adjacent areas in China can be found in Anonym (1979).
2
Methods
he species concept used herein is the evolutionary species concept (see Mayden & Wood, 1995;
Methods
Kottelat, 1997). The formal synonymies include only references to original descriptions of
nominal species and only those based on material from Asia and usually ignore those nominal
species whose type locality is in Europe or North America. I did not include information
on type material as this does not seem relevant for the level of the present discussion, except in a few
exceptional cases. The spellings of scientific names in the synonymies follow exactly those in the cited
publications; this explains the apparent inconsistencies. The nomenclatural authors (the authors who
first established a name) are not used in the text because they do not add to the discussion; to the
contrary, they make the reading more difficult and often create confusion for the readers not familiar
with nomenclatural rules. Anyway, their citation is only an optional tool for retrieval of bibliography
and—contrary to a frequent misperception—their mention is not compulsory. For those who absolutely
wish to know them, they are given in Table 1 and can also be retrieved from the formal synonymies.
Nomenclaturally relevant information for species cited in the text but not part of the Mongolian fauna
are listed in Appendix 1. Place names in the formal synonymies are as they appear in the original
publication, including the then-administrative or political units. In several of these historical accounts,
“Mongolia” includes present Mongolia, as well as Inner Mongolia, Liaoning, Xinjiang, Tibet, and
adjacent areas.
There are inconsistencies in the spelling or transcription of the names of rivers, lakes, and localities used
in this report. My efforts to find maps showing all names in a consistent system failed, as well as to find
a reviewer. I could have tried to adjust them, but these could result in introducing errors and confusion
and I prefer to stick to names as they appear in the literature or in the various maps accessible to me.
Toponymy for China follows whenever feasible Zhonghua Renmin Gongheguo Fen Sheng Dituji (Hanyu
Pinyinban), Beijing, 1977.
The “Great Lakes Valley”, or “Lakes Valley” of Russian authors is called Great Lakes Basin, and the “Gobi
Valley” is called Gobi Lakes Valley. This follows WWF ecoregion terminology (ecoregions PA1316 and
PA1315, respectively; http://worldwildlife.org/science/ecoregions/palearctic.cfm).
I have not attempted to list all citations of a given name as this would have been much too time-consuming
and detrimental to more important issues within the limited time available.
The discussions on the status of Chinese species of the family Cyprinidae is based to some extent on Chen
(1998) and Yue (2000). In many cases, I have followed these conclusions, but I have to indicate some
3
4
reservations because I have a strong feeling that the species diversity is severely underestimated for many
groups.
Bibliography and references: years in bold font indicate sources that I have not been able to examine
personally; the data quoted from these sources are those repeated in the literature.
For a few species that present identification problems, the abbreviation “cf.” added between the genus
and the species names indicates a species that looks similar to a named species but possibly represents
a distinct, unnamed species. For example Brachymystax cf. tumensis indicates that the species is similar
to B. tumensis but until a revision (a comparison of all species of the genus) can be done, one cannot be
absolutely sure.
Best Guesses
As will be obvious from many of the comments below, the status of many species/populations/names
Methods
is still far from clear and cannot be elucidated without actual baseline studies, especially not without a
sampling program to obtain the material suitable for a professional taxonomic analysis. Decisions might
be needed in relation with conservation issues and it is likely that many decisions cannot be delayed until
the taxonomy can be elucidated. As this report is written within a conservation background and aim, it
seems important that users may make decisions even in the absence of complete data or final taxonomic
conclusion. This might be important, for example, when precautionary decisions have to be made, or
research targets have to be defined. For these reasons, a number of species accounts include my “best
guess” or “educated guess” of what is likely to appear once the discussed problems can be clarified.
Clearly, these best guesses are not at all scientific results but can be considered as scientific hypotheses.
They are based on a blend of knowledge of the fauna, of the topography, of the mechanisms underlying
areas of endemism, experience with the literature, with the way of working of earlier (and recent) scientists
and, the most important tool in the field, instinct.
An educated guess often is the only efficient tool (and the only available) in the absence of reliable data, in
the field or for triage, whatever academics and theoreticians may say.
5
Species Lists
he check-list below summarizes nomenclaturally valid names of Mongolian fish species, with
T taxonomic authority. The apparent inconsistency with the use of parentheses in fact is precisely
dictated by the International Code of Zoological Nomenclature. The presence of parentheses
indicates that the species was placed in a different genus by its original author. For example,
Brachymystax lenok (Pallas, 1773) was first described by Pallas in 1773 as a species of the genus Salmo,
therefore the parentheses, and B. tumensis Mori, 1930 was originally described by Mori in 1930 as a
species of the genus Brachymystax, therefore no parentheses.
Native
Species Lists
Petromyzontidae
Lethenteron reissneri (Dybowski, 1869)
Acipenseridae
Acipenser baerii Brandt, 1869
Acipenser schrenckii Brandt, 1869
Salmonidae
Hucho taimen (Pallas, 1773)
Brachymystax lenok (Pallas, 1773)
Brachymystax cf. tumensis Mori, 1930
Coregonidae
Coregonus chadary Dybowski, 1869
Coregonus migratorius (Georgi, 1775)
Coregonus pidschian (Gmelin, 1789)
Thymallidae
Thymallus arcticus (Pallas, 1776)
Thymallus baicalensis Dybowski, 1874
Thymallus brevirostris Kessler, 1879
Thymallus grubii Dybowski, 1869
Thymallus nigrescens Dorogostaisky, 1923
Thymallus sp. 1
Esocidae
Esox lucius Linnaeus, 1758
Esox reichertii Dybowski, 1869
Cyprinidae
Acheilognathus asmussii (Dybowski, 1872)
7
Carassius carassius (Linnaeus, 1758)

Carassius gibelio (Bloch, 1782)
Chanodichthys erythropterus (Basilewsky, 1855)
Chanodichthys mongolicus (Basilewsky, 1855)
Culter alburnus Basilewsky, 1855
Cyprinus rubrofuscus La Cepède, 1803
Eupallasella percnurus (Pallas, 1814)
Gnathopogon strigatus (Regan, 1908)
Gobio acutipinnatus Menshikov, 1939
Gobio cynocephalus Dybowski, 1869
Gobio sibiricus Nikolski, 1936
Gobio soldatovi Berg, 1914
Gobio tenuicorpus Mori, 1934
Gobio sp. Onon
Hemibarbus labeo (Pallas, 1776)
Hemibarbus maculatus Bleeker, 1871
Hemiculter leucisculus (Basilewsky, 1855)
Hemiculter varpachovskii Nikolski, 1904
Ladislavia taczanowskii Dybowski, 1869
Leuciscus baicalensis (Dybowski, 1874)
Leuciscus dzungaricus Koch & Paepke, 1998
Leuciscus idus (Linnaeus, 1758)
Leuciscus waleckii (Dybowski, 1869)
Microphysogobio anudarini Holcík & Pivnicka, 1969
Oreoleuciscus angusticephalus Bogutskaya, 2001
Oreoleuciscus dsapchynensis Warpachowski, 1899
Oreoleuciscus humilis Warpachowski, 1889
Oreoleuciscus potanini (Kessler, 1879)
Phoxinus cf. phoxinus (Linnaeus, 1758)
Phoxinus ujmonensis Kashchenko, 1899
Pseudaspius leptocephalus (Pallas, 1776)
Pseudorasbora parva (Temminck & Schlegel, 1846)
Rhodeus sericeus (Pallas, 1776)
Rhynchocypris czekanowskii (Dybowski, 1869)
Rhynchocypris lagowskii (Dybowski, 1869)
Rutilus rutilus (Linnaeus, 1758)
Sarcocheilichthys soldatovi (Berg, 1914)
Saurogobio dabryi Bleeker, 1871
Squalidus chankaensis (Dybowski, 1872)
Tinca tinca (Linnaeus, 1758)
Nemacheilidae
Barbatula compressirostris (Warpachowski, 1897)
Barbatula dgebuadzei (Prokofiev, 2003)
Barbatula toni (Dybowski, 1869)
Barbatula sp. Tuul
Barbatula sp. Egiin
Lefua costata Kessler, 1876
8
Triplophysa gundriseri Prokofiev, 2002
Triplophysa sp. Tuul
Cobitidae
Cobitis melanoleuca Nichols, 1925
Iksookimia lebedevi (Vasil’eva & Vasil’ev, 1984)
Misgurnus mohoity (Dybowski, 1869)
Siluridae
Silurus asotus Linnaeus, 1758
Lotidae
Lota lota (Linnaeus, 1758)
Percidae
Perca fluviatilis Linnaeus, 1758
Cottidae
Cottus szanaga Dybowski, 1869
Mesocottus haitej (Dybowski, 1869)
Leocottus kesslerii (Dybowski, 1874)
Odontobutidae
Perccottus glenii Dybowski, 1877
Introduced
a) Species that established self-sustaining populations
Species Lists
Coregonidae
Coregonus peled (Gmelin, 1789)
Cyprinus carpio Linnaeus, 1758
b) Species usually reported to be present in Mongolia but apparently result from occasional capture of
stocked or aquaculture individuals, or introduced species that did not establish
Coregonidae
Coregonus sardinella Valenciennes, 1848
Cyprinidae
Ctenopharyngodon idella (Valenciennes, 1844)
Hypophthalmichthys molitrix (Valenciennes, 1844)
Siluridae
Silurus soldatovi Nikolski & Soin, 1948
9
The list below contains the invalid names, synonyms, misidentifications, main misspellings, and erroneous
records appearing in recent Mongolian literature and Russian and Chinese literature for adjacent areas and
their valid equivalents.
INVALID NAMES VALID NAME

Acanthorhodeus asmussi Acheilognathus asmussii
Acipenser baeri baicalensis Acipenser baerii
Barbatula barbatula toni Barbatula toni
Barbatula nuda misidentified Barbatula toni
Carassius auratus gibelio Carassius gibelio
Chilogobio czerskii misidentified Sarcocheilichthys soldatovi
Chilogobio soldatovi Sarcocheilichthys soldatovi
Cobitis granoei olivai misidentified, C. melanoleuca
Cobitis granoei misidentified, C. melanoleuca
Cobitis lebedevi Iksookimia lebedevi
Cobitis taenia granoei misidentified, C. melanoleuca
Cobitis taenia sibirica misidentified, C. melanoleuca
Cobitis taenia misidentified, C. melanoleuca
Coregonus autumnalis migratorius Coregonus migratorius
Coregonus cylindraceus Prosopium cylindraceum; no actual record from
Mongolia
Coregonus mongolicus Prosopium cylindraceum; no actual record from
Mongolia
Coregonus lavaretus pidschian Coregonus pidschian
Cottus kesslerii Leocottus kesslerii
Cottus poecilopus misidentified Cottus szanaga
Cottus sibiricus no actual record from Mongolia
Culter brevicauda Culter alburnus
Culter erythropterus applied to both Chanodichthys erythropterus and
Culter alburnus
Culter mongolicus Chanodichthys mongolicus
Cultrichthys erythropterus applied to both Chanodichthys erythropterus and
Culter alburnus
Cyprinus carpio haematopterus Cyprinus rubrofuscus
Cyprinus carpio applied to both Cyprinus carpio (introduced) and C.
rubrofuscus (native)
Eleotris glehni Perccottus glenii
Erythroculter erythropterus applied to both Chanodichthys erythropterus and
Culter alburnus
Erythroculter mongolicus Chanodichthys mongolicus
Gnathopogon chankaensis Squalidus chankaensis
Gobio albipinnatus tenuicorpus Gobio tenuicorpus
Gobio albipinnatus Gobio tenuicorpus
Gobio gobio cynocephalus applied to both Gobio cynocephalus and G. sibiricus
Gobio gobio sibiricus Gobio sibiricus
10
Gobio gobio misidentified Gobio acutipinnatus, G. cynocephalus,
G. sibiricus
Gobio soldatovi tungussicus misidentified Gobio sp. Onon
Hemiculter bleekeri varpachovskii Hemiculter varpachovskii
Hemiculter lucidus Hemiculter varpachovskii
Hemiculter lucidus varpachovskii Hemiculter varpachovskii
Huso dauricus no actual record from Mongolia
Iksookimia choii misidentified Iksookimia lebedevi
Lagowskiella czekanowskii Rhynchocypris czekanowskii
Lagowskiella lagowskii Rhynchocypris lagowskii
Lampetra japonica Lethenteron camtschaticum; no actual record from
Mongolia
Lampetra reissneri Lethenteron reissneri
Lethenteron japonicum Lethenteron camtschaticum; no actual record from
Mongolia
Leuciscus leuciscus baicalensis Leuciscus baicalensis
Microphysogobio amurensis misidentified, Microphysogobio anudarini
Microphysogobio tungting misidentified, Microphysogobio anudarini
Misgurnus anguillicaudatus misidentified Misgurnus mohoity
Nemacheilus barbatula toni Barbatula toni
Nemacheilus dorsalis humilis invalid name replaced by Triplophysa gundriseri
Nemacheilus nudus misidentified Barbatula toni
Species Lists
Nemacheilus toni Barbatula toni
Nemachilus cobdonensis see under Barbatula compressirostris
Nemachilus compressirostris Barbatula compressirostris
Noemacheilus stoliczkai no actual record from Mongolia
Noemacheilus strauchi misidentified Triplophysa gundriseri
Oreoleuciscus pewzowi misidentified Oreoleuciscus angusticephalus
Orthrias dgebuadzei Barbatula dgebuadzei
Orthrias golubtsovi see under Barbatula compressirostris
Orthrias toni Barbatula toni
Paracottus kessleri Leocottus kesslerii
Paraleucogobio strigatus Gnathopogon strigatus
Parasilurus asotus Silurus asotus
Phoxinus czekanowskii Rhynchocypris czekanowskii
Phoxinus lagowskii Rhynchocypris lagowskii
Phoxinus oxycephalus misidentified Rhynchocypris lagowskii
Phoxinus percnurus Eupallasella percnurus
Phoxinus perenurus Eupallasella percnurus
Prosopium cylindraceum Prosopium cylindraceum; no actual record from
Mongolia
Rhinchocypris Rhynchocypris
Rhynchocypris costata misidentified Rhynchocypris lagowskii
Rhynchoypris oxycephalus misidentified Rhynchocypris lagowskii
Romanogobio tenuicorpus Gobio tenuicorpus
11
Rostrogobio amurensis misidentified Microphysogobio anudarini

Rutilus rutilus lacustris Rutilus rutilus
Sarcocheilichthys czerskii misidentified Sarcocheilichthys soldatovi
Sarcocheilichthys nigripinnus czerckii misidentified Sarcocheilichthys soldatovi
Saurogobio amurensis misidentified Microphysogobio anudarini
Silurus soldatovi misidentified Silurus asotus
12
Accounts of Species Recorded from Mongolia
Family Petromyzontidae habitats. It is not known from the Yenisei drainage

(lampreys) (Berg, 1948: 38; Reshetnikov, 2002: 28) and in the
Amur it is recorded only downriver of Khabarovsk.
Lampreys are eel-like fishes. The adults are All records of lampreys in Mongolia are from the
immediately recognized by their disc-shaped mouth Onon drainage and apparently refer to L. reissneri.
without jaws (that is, which cannot be closed) and
7 round branchial openings on each side of the Lethenteron camtschaticum is included in the
head. The species are mainly distinguished by the Mongolian fauna by Baasanjav & Tsendayush
kind, number and disposition of teeth. The larvae (2001: 13) because of the presence of a specimen
(called ammocoetes) spend several years hidden from River Gorlog (Mongolian name of upper
in muddy to sandy bottom. The large species live part of Yenisei) at Kargina (53.75°N 110.17°E)
several years, may migrate to the sea, are predatory in the Zoological Institute of St. Petersburg
and feed on fish to which they attach with their (ZISP 28338). This leads them to speculate on
mouth. The small species are non-predatory the presence of the species in the Selenge and in
and non migratory; after their larval stage, they Mongolia. Lethenteron reissneri is the only lamprey
metamorphose and reproduce within a few weeks definitively recorded from the Yenisei (but only
or months. The adults do not eat and they die downriver of Lake Baikal).
shortly after spawning. Their dentition is relatively
undeveloped and it is often difficult to identify Lethenteron reissneri
them. Petromyzon Reissneri Dybowski, 1869: 958 (type
locality: Russia: Amur basin: Onon and Ingoda
Two species have been reported from Mongolia, rivers)
Lampetra japonica and Lampetra reissneri. Both are Lampetra mitsukurii Hatta, 1901: 24 (type locality:
now placed in the genus Lethenteron; the species Japan: Hondo, Hokkaido)
name Lampetra japonica in fact is invalid and the Petromyzon kessleri Anikin, 1905: 10 (type locality:
correct name is Lethenteron camtschaticum (see Russia: Tom River and mouth of Kirgizka River
Kottelat, 1997). At present, there is no evidence near Tomsk, Ob drainage)
of the presence of Lethenteron camtschaticum in Lampetra mitsukurii minor Hatta, 1911: 268, pl.
Mongolia. The species is known from the lower 8 figs. 3–4 (type locality: Japan: Gifu, Sapporo,
reaches of rivers or lowlands, coastal and estuarine and numerous localities listed p. 264)
13
Remarks on systematics. Lethenteron reissneri is of the L. kessleri from northern Japan and Far
reported from Mongolia as Lampetra reissneri by Eastern Russia, and not conspecific with the two
Holcik & Pivnicka (1969), Dashdorj & Demin Japanese L. cf. reissneri species.
(1977: 142). Sokolov (1983: 104) considered these
records to be misidentified Lampetra japonica. Anyway, as the type locality of L. reissneri is the
Baasanjav & Tsendayush (2001: 13) considered Onon and Ingoda rivers, at least we know that the
that L. reissneri indeed occurs in Mongolia, in name of the Mongolian populations will not be
Onon-Khurkh drainage. They also considered that affected by the final identity of the Japanese species.
the records of Lampetra japonica by Dulmaa (1999: In any case, the name L. reissneri (established by
194) and Bogayevski (1949; Khod, Promysel, Dybowski, 1869) will remain as it is older than L.
Amur drainage) refer to this species [they do not kessleri (established by Anikin, 1905).
give the bibliographic reference to Bogayevski’s
paper]. Distribution. In Mongolia: only recorded in Onon
drainage. Distribution outside Mongolia: Eurasia,
The description in Sokolov (1983: 104) is from in Arctic Oceans basin, from Kola Peninsula to
Nikolski (1956) and is based on specimens in Anadyr drainage; Amur drainage; Sakhalin Island;
the Zoological Institute of St Petersburg (ZISP Hokkaido (Japan).
25351) collected in the stream Naleo which flows
into the liman of Amur River (liman is a Russian
terminology designating a lagoon at the estuary of Family Acipenseridae
a river created by a littoral bar blocking the access (sturgeons)
to the sea).
Three species of sturgeons are listed in the recent
The record by Holcik & Pivnicka (1969: 3) from the literature as occurring in Mongolia, but one (Huso
Onon is based on a single larva (ammocoete) 147 dauricus) is only an assumption.
mm long. They do not indicate which diagnostic
characters they used for its identification. Huso dauricus is listed by Baasanjav & Tsendayush
(2001: 15) on the basis of information by people
In her account of “L. japonica”, Dulmaa (1999) in the area of Bindir (on Onon River, Amur drain-
does not mention predatory (parasitic) feeding, age) of a specimen “as long as the platform of a
but mentions a small adult size (200 mm), A3-51” [GA3-51; a model of Russian truck; the
explicitly states that it does not migrate far, and platform is about 2 m long] caught in the 1950s.
that it has a mean number of 9,000 oocytes. This is No other information is available to identify the
a misidentification as L. camtschaticum reaches 625 species, except for the size and the drainage. Two
mm, has over 50,000 oocytes, migrates to the sea species of sturgeons are known from the Amur
where it spends 1–3 years, and is predatory. drainage, Huso dauricus and Acipenser schrenckii.
The distribution of both is mapped by Reshetnikov
The taxonomy of the various populations usually (2002: 49, 54) and is recorded to reach only to the
referred to as L. reissneri is not yet settled. confluence of the Onon and Indoga, about 400
Lethenteron reissneri is usually recorded to occur in river-km downstream of the Mongolian border.
all the rivers draining to the Pacific Ocean, from Huso dauricus is recorded to reach a maximum size
Kamtschatka to Korea, and Japan. Molecular of 5 m (Reshetnikov, 2002: 54) and A. schrenckii
studies of a number of populations in Japan have up to 2.9 m and therefore there is no way to
shown that the L. reissneri of Japanese authors in objectively identify this individual. The largest
fact represents two species, which even occur in individual A. schrenckii known from the Amur
sympatry at some localities (Yamazaki et al., 2003). drainage was 204 cm long (Dulmaa, 1999: 194)
Yamazaki et al. (2006) show that the L. reissneri and the Bindir specimen may be either one of the
from the Onon in fact is the same species as some Amur species A. schrenckii or H. dauricus.
14
Acipenser baerii in Mongolia and that it should be expected also
in its largest tributaries like the Balj and Khurkh.
Acipenser Baerii Brandt, 1869a: 175 (type locality:
Dulmaa (1999:194) report it as inhabiting “some
Russia: Siberia: Ob and Lena rivers and their
sections of the River Onon, adjacent to Russia”, but
main tributaries; also in Brandt, 1869b: 115)
without indicating the source of this information.
Acipenser stenorrhynchus Nikolski, 1896: 400 (type
Its presence in Mongolia is based on the specimen
locality: Russia: “large rivers of Siberia flowing
from Bindir identified as Huso dauricus by Baasanjav
to Arctic Ocean, but not in Lake Baikal”)
& Tsendayush (2001: 15); see family introduction
Acipenser stenorrhynchus var. baicalensis Nikolski,
for discussion.
1896: 401 (type locality: Russia: Siberia: Lake
Baikal)
The morphological data in Sokolov (1983: 107) are
Acipenser baeri chatys Drjagin, 1948: 532 (type
from Nikolski (1956) and not based on Mongolian
locality: Russia: Sakha [Yakutia]: from Lena to
specimens.
Kolyma rivers)
Distribution. In Mongolia: Onon drainage.
Remarks on systematics. Acipenser baerii is defin-
Outside Mongolia: Amur drainage. Also in Yalu
itively recorded in Mongolia, by Sokolov (1983:
River in Jilin Province of China, and North Korea
108), Baasanjav & Tsendayush (2001: 17), Dulmaa
according to map in Reshetnikov (2002: 49), but
(1999: 194) and others.
is not listed from Korea by Kim (1997) or from
Tumen drainage by Zheng et al. (1980).
Some authors have recognized up to 4 subspecies
of A. baerii, A. b. baerii (from Ob and Irtysh
drainages), A. b. stenorrhynchus (from Yenisei to
Anabar drainages), A. b. baicalensis (from Baikal Family Salmonidae
basin [in Yenisei drainage]), and A. b. chatys (from (salmons and trouts)
Lena to Kolyma drainages) (Reshetnikov et al.,
1997: 689; Sokolov, in Reshetnikov, 1998: 19). Brachymystax lenok
However, from the original descriptions and the
Salmo lenok Pallas, 1773: 716 (type locality:
scant data in Berg (1948: 88–89) and Sokolov &
Russia: Yenisei River/streams on hills of eastern
Vasil’ev (in Holcik, 1989: 268) their taxonomic
Siberia)
status cannot be decided. In the absence of
Salmo coregonoides Pallas, 1814: 362 (type locality:
diagnostic information, I do not see reason to
Russia: streams of the Altai range draining to the
distinguish these populations by formal names.
Ob and Irtin [Irtysh]/Yenisei and its tributaries
/Lake Baikal and Rivers Angara and Selenge/
Distribution. In Mongolia: Selenge drainage.
Rivers Lena, Witim and Kovyma [Kolyma])
Outside Mongolia: drainages of all large rivers
? Brachymystax lenok savinovi Mitrofanov, 1959:
flowing to the Arctic Ocean, from Ob in the west
275 (type locality: Kazakhstan: Altai: Lake
to Kolyma in the east.
Marka-Kul)
Brachymystax lenok swetowidowi Kirillov, 1962: 12
Acipenser schrenckii
(type locality: Russia: Yakutia: Vilyui River)
Acipenser Schrenckii Brandt, 1869a: 175 (type
locality: Russia: Amur River and its main Remarks on systematics. Two ‘forms’ of lenok are
tributaries; also in Brandt, 1869b: 115) recognized in Mongolia: 1) A ‘common’ “form”,
with pointed snout and large rounded reddish spots,
Remarks on systematics. Acipenser schrenckii is present in the whole country; hereunder ‘pointed-
listed by Sokolov (1983: 107) and Baasanjav & snout lenok’; 2) a ‘form’ with similar body shape, but
Tsendayush (2001: 18). This is based on Dashdorj with blunt snout, and a colour pattern consisting
(1977:143) who guessed its presence in the Onon in small black round spots on the sides, found only
15
in Kherlen River (together with the pointed-snout support the conspecificity of the East Siberian and
lenok); hereunder ‘blunt-snout lenok’. Amur pointed-snout lenok and the conspecificity
of the East Siberian and Amur blunt-snout lenok.
Their taxonomic identity is not clear and they have With two species of lenok in Siberia and Mongolia,
been at some time treated as different species or it remains to establish which one is the ‘real’ B.
subspecies (e.g., Kifa, 1976; Bogutskaya & Naseka, lenok and what the name is of the other. The type
2004: 150). Others treat them as a single species, locality of B. lenok being the Yenisei, and as only the
though acknowledging the need for further study pointed-snout species is known from the Yenisei,
(e.g., Reshetnikov et al., 1997: 692). thus on the basis of available information the
pointed-snout lenok retains the name B. lenok.
Alekseev et al. (2003) studied Brachymystax in
East Siberia, from the Olenek to the Kolyma Kifa (1976) recognized the existence of two species
drainages. The pointed-snout lenok was observed of lenok in the Amur drainage and used the names
in all drainages, while they collected the blunt- B. lenok for the pointed-snout lenok and B. savinovi
snout lenok only in some upper tributaries of the for the blunt-snout lenok. The type locality of B.
Lena (Vitim and Olekma) and along the northern savinovi is Lake Marka-Kul (in the upper Irtysh
stretch of the Lena itself. The two ‘forms’ are drainage in Kazakhstan). I could not access the
found in the upper tributaries. Alekseev et al. note original description of B. l. savinovi by Mitrofanov
that the blunt-snout lenok is more restricted to (1959) but the description in Mitrofanov (1986)
mountain tributaries and upstream lakes, and in shows a fish with a blunt snout and low gill-raker
some lakes only this ‘form’ was found. Where they count (17–24, usually 20–21) as observed in the East
occur together, individual hybrids are found, but Siberian and Amur blunt-snout lenok. Noteworthy
they do not form “hybrid swamps”. They do not is also that B. savinovi is described from an altitude
report blunt-snout lenok in the Yenisei or in the lake, which is also the frequent habitat of the Lena
Selenge drainages. blunt-snout lenok. Brachymystax lenok savinovi is
considered a synonym of B. lenok by, e.g., Shedko
The two ‘forms’ of lenok are morphologically (2001) and Bogutskaya & Naseka (2004) who use
distinct, they live in sympatry and only occasionally the name B. tumensis for the blunt-snout lenok of
hybridize and this suggests they are distinct species. the Amur drainage (see comments below).
Molecular data in Froufe et al. (2003) shows that
the same pattern is observed in the Lena and Amur The synonymy of B. lenok and B. savinovi might
drainages. They also show that the two ‘forms’ each be premature. Alekseev & Osinov (2006) studied
have unique haplotypes and that identical ‘forms’ the blunt-snout lenoks of the Ob-Irtysh drainage.
in different drainages share the same haplotypes. They conclude that they “diverge substantially
They too do not report blunt-snout lenok in the morphologically and, especially, genetically from
Yenisei or in the Selenge drainages. populations of the blunt-snout[…] lenok from
other river basins of Siberia and the Far East of
Alekseev et al. (2003) comment that in East Siberia Russia. In the Ob’-Irtysh basin populations of the
(from Lena to Kolyma drainages), individuals of the blunt- and [pointed-snout] lenoks are spatially
two forms differ from individuals of their respective segregated: the former are found in the Ob’ River
forms from the basins of the Ob, Selenge, Amur basin proper and the latter, in the Irtysh River basin,
and Uda rivers and from the rivers of Primorye. the degree of their morphological and genetic […]
They give some of the differences between the East divergence is high.” This suggests that they might
Siberian ‘forms’ and the Amur ‘forms’, but in a way be distinct species, and that B. savinovi would be
not allowing an explicit comparison, so that the the name of the blunt-snout lenok of the Irtysh.
significance of the differences cannot be evaluated.
The differences seem slight and, combined with Li et al. (1966: 42) report the presence of B. lenok
the genetic data in Froufe et al. (2003), seem to in the Ertix [Irtysh] in China and it can reasonably
16
be expected in the Bulgan River in Mongolia (an Brachymystax from Amur and Tumen are conspecific.
upper tributary of the Ertix). In their book on The name B. tumensis does not seem to appear in
fishes of Xinjiang, Anonym (1979) recorded B. the recent Korean literature. Instead, authors (e.g.,
lenok only from the Ertix; they have a figure (fig. Jeon, 1987; Kim, 1997: 356) report B. l. tsinlingensis
4) showing a fish with a moderately pointed snout from Korea; their figures show individuals with a
labelled B. lenok, the text (p. 11) mentions 21–27 blunt snout (obviously juveniles). Kim’s (1997) map
gill-rakers, which is congruent with Mitrofanov’s B. shows its distribution extending from South Korea
lenok (19–27), and slightly less than the pointed- to about Sovetskaya Gavan (49.0°N 140.2°E). The
snout lenok from Lena (22–29; Alekseev et al., source of the information is not clear and I do not
2003) (the blunt-snout lenok of Lena has 18–23). know how accurate the map is; it shows the range
When the presence of Brachymystax is confirmed extending on part of the Ussuri, a tributary of the
in the Bulgan, their identity deserves investigation Amur. It is noteworthy that all the references to
as they would probably be B. savinovi, adding one Brachymystax in Korean and immediately adjacent
more species to the Mongolian fauna. waters refer only to blunt-snout lenok and that they
are in relatively lowlands, while the Lena blunt-
Remarks on nomenclature. A neotype designation snout lenok are found in altitude rivers and lakes.
might be necessary to definitively retain B. lenok as Similarly, the records by Bogutskaya et al. (2001:
the name for the pointed-snout lenok. 41; Lake Khanka) and Shedko (2001; Primorye) are
Distribution. In Mongolia: Selenge, Yenisei and in relatively low areas.
Amur drainages; might be present in Bulgan
drainage. Outside Mongolia: rivers draining to Brachymystax tsinlingensis was originally described
Arctic and Pacific Oceans, from Ob to Amur. from the Qin Ling range in Shaanxi Province of
China, which makes the divide between the Huang
Brachymystax cf. tumensis He (Yellow River) and Chang Jiang (Yangtze)
drainages. The figure in the original description
Brachymystax tumensis Mori, 1930: 42, pl. 3 fig. 1 (Li, 1966), Chen (1987) and Anonym (1992)
(type locality: Korea: Tumen River, Yen-gan) show a deep-bodied, blunt-snouted fish. See also
Remarks on systematics. Brachymystax tumensis Chen (1986) for distribution data.
was described from Tumen River. It has been
considered as a synonym of B. lenok but Very recently, Xia et al. (2006) analysed the
Bogutskaya et al. (2001: 41), Shedko (2001) and phylogeographic structure of Brachymystax pop-
Bogutskaya & Naseka (2004: 151) consider it as ulations in Chinese waters. The study is restricted
a valid species and use this name for the blunt- to a geographic area marginal to the whole
snout lenok. This deserves more investigation. distribution of the genus. Nevertheless, some of
While Shedko recognizes a single species of lenok their observations are relevant here. They examined
in Primorye [southeasternmost province of Russia, specimens from seven populations in the Amur,
at the border with China and Korea], Bogutskaya Tumen, Yalu and Yellow Rivers. Their results
et al. (2001) and Bogutskaya & Naseka (2004) show three clearly separate lineages, 1) Amur
recognize two species, B. lenok (pointed-snout River, 2) Tumen and Yalu Rivers, and 3) Yellow
lenok) and B. tumensis (blunt-snout lenok). Mori’s River. They also discussed the published data on a
(1930) description and figure of B. tumensis show Korean population from “Hanjiang” [Hangang ?]
a fish with numerous large spots on whole body, 22 (Korea), which is placed in the Yellow River clade.
gill-rakers and a quite elongate snout; Mori (p. 42) Unfortunately, their study does not make any
even comments that the species has a more pointed connection with morphological information.
snout than B. lenok.
In conclusion, if it can be demonstrated that there
Tumen River forms the border between North Korea is a single species of blunt-snout lenok in the whole
and China. It has yet to be demonstrated that the Amur drainage and if it can be demonstrated that it
17
is conspecific with the Tumen lenok, then the name Family Coregonidae
of the blunt-snout lenok present in Mongolia is B. (whitefishes, ciscos)
tumensis. With our present knowledge, there is no
way to confirm this hypothesis. On the contrary, The taxonomy of the Eurasian species of the genus
there are indications that future work might show Coregonus is extremely confused because of a variety
they are distinct species, in which case the name of factors. First of all, being pelagic and silvery,
B. tumensis would be used for the Tumen and Yalu coregonids exhibit very few salient characters. This
species, and there would be no name available for is contrary to the situation with benthic fishes,
the Amur species. Until this is clarified, I retain the which usually have a colour pattern or peculiar
name B. cf. tumensis for all the blunt-snout lenok morphological features (for camouflage and for
of Lena, Kolyma, Amur and Tumen. The published visual recognition) and our human eyes and brain
data suggest that B. tsinlingensis is a valid species, recognize these characters, thereby helping us to
distributed in the the Yellow River drainage (and identify the fish. These characters are usually not
central Korea). prominent in pelagic fishes, which use non-visual
signals to identify conspecifics.
Distribution. In Mongolia: Onon and Kherlen
drainages. Outside Mongolia: Amur, Lena and Second, at superficial examination, most species
Kolyma drainages in Russia and china; Tumen look similar. This resulted in their identification
drainage in China and Korea. being based only on a few characters, like number of
gill rakers or lateral line scales. To people not used to
Hucho taimen taxonomic procedures, countable characters appears
Salmo Taïmen Pallas, 1773: 716 (type locality: more ‘reliable’ than characters which need to be
Russia: rivers of Siberia flowing to Arctic described, like shape of mouth parts etc. This of
Ocean; original spelling should be emended course is not true, these characters are simply easier
into taimen, Code art. 32.5.2.1) to use for untrained workers. But the unfortunate
Salmo fluviatilis Pallas, 1814: 359, pl. 73 fig. 2 (type result is that people pay more attention to some sort
locality: Russia: Siberia: tributaries of Rivers of ‘magic’ formulae than to shapes and structures.
Ob, Irtysh, Yenisei, Lena and tributaries, Lakes
Baikal, Vitim, Turuchansk, Ljala, Tura, Uba, Third, the systematics of coregonids has been
Tom, Kama and all streams flowing towards handled mainly by fishery biologists without
“Eastern Ocean” except Kamchatka) training to handle taxonomic problems. In
? Salmo lossos Günther, 1866: 140 (type locality: some areas (Russia, eastern Europe), it has
Baltic Sea, Rivers Kama, Kolva, Volga, Petschow, been addressed with ‘endemic’ concepts and a
Vitchegda, and Muilwa, Caspian Sea) nomenclatural system not entirely compatible
with international practices and rules. This system
Remarks on systematics. Despite its wide distri- recognizes a number of infrasubspecific categories,
bution across all of northern Asia, H. taimen does not always explicitly defined. Infrasubspecific
not show genetic differentiation across the Amur, names are not available for formal nomenclature,
Lena and Yenisei drainages (Froufe et al., 2003). but the occasional use of some of them at a species
There is no data for the populations of the remaining or subspecies ‘level’ automatically made them
basins. available and it is extremely time consuming to
check these and conclude (or often guess) their
Distribution. In Mongolia: Yenisei, Selenge, Onon real status. Language issues contribute to make the
and Kherlen drainages. Outside Mongolia: rivers problems more complex.
draining to the Arctic Ocean from Ob to Yana
drainages; Amur and some adjacent drainages; in Fourth, with the great confusion resulting from the
Europe, some upper tributaries of Pechora, and above, a ‘concept’ evolved that coregonids escape
Kama in Volga drainage. the general rules and that coregonid systematics
18
has to be different from the systematics of other Coregonus lavaretus is a species endemic to Lake
fishes. Then appeared an esoteric ‘Coregonus species Bourget in France. The “species” (singular)
concept’, which certainly is not an appropriate and identified as C. lavaretus elsewhere (and most
professional approach of taxonomy. especially in the Russian literature) is a collection
of maybe up to 50–100 species, each with its own
To make the whole pattern even more complex, distribution, biology etc. Some are widespread
many species/populations have been transplanted, and some are highly endemic to a single or a
introduced and/or hybridized, making it sometimes few lakes; in many lakes several species may be
impossible to sort out what the original species were. in sympatry (up to 11; in Mongolia there is at
least a record of 2 sympatric ‘forms’ in Darkhad
Tradition, the (administrative) fear to have to depression) and the conservation status of all
manage and conserve a significantly larger number should be determined individually. This is further
of species and their economic value are certainly complicated because there have been several
forces driving some scientists and agencies to close attempts to introduce Russian species (see, e.g.,
their eyes to, or to seek to negate biological reality Dulmaa, 1979), some of which were successful (in
and the existence of this taxonomic diversity, or creating self-sustaining populations and maybe in
to refuse to give it the same value that a similar eliminating native ones). Therefore, there is a need
diversity would receive had the taxa concerned to distinguish between the native and the stocked
been birds or African cichlids. species and populations. The stocked ones, even
if acclimatised, should not appear in discussion
Most authors now appear to be content simply of biodiversity issues. A difference should also be
to mention this complexity as an excuse for not made between species native at a given locality,
elaborating further on the topic. This resulted in and those native to the country but translocated
almost all populations of the family being dumped at a given locality.
at various times in a ‘mythic’ catch-all species
named Coregonus lavaretus, which was believed to be In the absence of detailed study of the Mongolian
extremely variable and plastic, able to adapt quickly coregonids and also because of the messy state of
to new habitats and which axiomatically had evolved their taxonomy throughout Siberia, it is impossible
into a great number of ecological morphs. Reviews to reach conclusions as to the status and identity of
of the diversity of coregonids in several areas is the Mongolian species and I conservatively adhere
now showing that this is much exaggerated, that to the ‘taxonomy’ in current local use.
morphological and genetic differences exist, that
species can be recognized and that the problem can Prosopium cylindraceum is listed in the Mongolian
be addressed as can be the taxonomy of any other fauna by Baasanjav & Tsendayush (2001: 38).
group of fishes. Simply, the number of the species Warpachowski (1901) described Coregonus
makes it a cumbersome and boring work. mongolicus which is considered as a synonym of P.
cylindraceum. The type locality is “lake of northern
In several lakes, several populations (up to 11 in Mongolia adjacent to sources of Yenissei River”.
Lake Onega, Russia) are morphologically (and It is not clear whether this locality is within the
genetically, for the few investigated cases) distinct, boundaries of present-day Mongolia. Actually there
live in sympatry, and have different habitats, ecology, is no subsequent record of the species in Mongolian
preferred preys and spawning seasons. These waters, and its presence is only guessed because of
populations fit the definition of species, negating its known presence in Gorlog River (upper Yenisei)
the theory that only a single species is involved. downstream of the Darkhad depression, but in
In fact, the problem is not so much to recognize Russia.
the different species, which occur in sympatry in
a given water body, but to find out whether or not Baasanjav & Tsendayush (2001: 31) list C.
species in adjacent water bodies are conspecific. sardinella as introduced in Mongolia in 1982
19
into Lakes Tolbo and Khongor-Ulen [small lakes they differ in characters other than growth rate.
in Bayan Ulgii aimag] from the hatchery of Biisk With the available data, no taxonomic value can
(Altay Province, Russia). Apparently this species be given to these ‘races’.
did not establish and is not mentioned by Dulmaa
(1995; Erdenebat M., pers. comm., February Remarks on nomenclature. This species often ap-
2006). Therefore it is not included here. pears under the name C. autumnalis migratorius
(e.g., Dulmaa, 1973; Baasanjav & Tsendayush,
Coregonus chadary 2001: 32). If one were considering that C.
autumnalis has two subspecies (migratorius and
Coregonus chadary Dybowski, 1869: 954, pl. 17
autumnalis), then their correct names should be
fig. 8 (type locality: Russia: Amur drainage:
C. migratorius migratorius (Georgi, 1775a) and
Onon River; spelled chavary p. 954, chadary in
C. m. autumnalis (Pallas, 1776) because the name
Table facing p. 958 and on pl. 17; first revised
migratorius is older than autumnalis.
[Dybowski, 1872: 222] retained C. chadary as
correct original spelling) The name C. autumnalis is now used for the
species migratory between the Arctic Ocean and
Remarks on nomenclature. Coregonus chadari is most major rivers draining to it, except the Ob.
an incorrect spelling. The name C. autumnalis is often seen in European
literature for a species from Lough Neagh, Ireland.
Distribution. In Mongolia: Onon drainage. Out- The correct name of this species is Coregonus pollan
side Mongolia: Amur drainage. (see Kottelat, 1997: 121).
Coregonus migratorius Distribution. In Mongolia: native, migratory in

Selenge, Delgermurun and Egiin Rivers. Outside
Salmo migratorius Georgi, 1775a: 182 (type
Mongolia: Lake Baikal basin. Introduced in 1956–
locality: Russia: Lake Baikal and its tributaries
1957 in Lake Khuvsgul (Dulmaa, 1973: 61),
upper Angara, Sosnowka, Tschiwirkui, Kowak,
where it is now established. The larvae came from
Bargusin and Selenge)
the hatchery of Bolsherechinsk, Russia.
Remarks on systematics. Coregonus migratorius is
treated as a subspecies of C. autumnalis in earlier
Coregonus peled
Russian literature (e.g., Berg, 1948: 342) (but see Salmo Peled Gmelin, 1789: 1379 (based on
below comment on nomenclature). Bogutskaya & Lepechin, 1780: 226, pl. 12; type locality:
Naseka (2004: 143) treat them as distinct species. “Russia boreali” [Northern Russia: Pustozersk
Coregonus autumnalis inhabits the lower part on Pechora River; Berg, 1948: 347])
of all drainages of Arctic Ocean in Eurasia from Salmo Peled Walbaum, 1792: 74 (based on
Mezen eastward (except Ob and Baikal), and north Lepechin, 1780: 226, pl. 12; type locality:
America from Cape Barrow to Coronation Bay; it “Russia boreali” [Northern Russia: Pustozersk
migrates from the sea and estuaries to spawn in on Pechora River; Berg, 1948: 347]; spelled
freshwater, migrating up to 1500 km upstream in pelcol in text, peles in index p. 714, corrected to
the Yenisei (Berg, 1948: 340). peled in emendanda)
Salmo cyprinoides Pallas, 1814: 412 (type locality:
Coregonus migratorius inhabits only Lake Baikal Russia: River Lena at Tungusis; junior homonym
basin and migrates into its tributaries for spawning. of Salmo cyprinoides Linnaeus, 1766: 514)
It does not seem to have spawning migration into Salmo Pelet Pallas, 1814: 412 (type locality: Russia:
its effluent Lower Angara. estuary of Yenisei)
Coregonus Syrok Valenciennes, in Cuvier &
Berg (1948: 366) recognizes three ‘races’ (including Valenciennes, 1848: 499 (based on Salmo wimba
a ‘Selenge race’) but his data do not suggest that of Pallas, 1814: 409; type locality: Russia: Ob
20
River and other rivers of eastern Siberia, Pechora Coregonus fera forma inarensis Järvi, 1928: 29, pl.
and lakes bordering Arctic Ocean, Lake Baikal, 4 figs. 19–20 (type locality: Finland: Lake Inari
Tungusk) at mouth of Rivers Juutuan and Niipi and at
Coregonus Rudolphianus Valenciennes, in Cuvier & Virtaniemi, Lake Muddus)
Valenciennes, 1848: 531 (based on Coregonus ? Coregonus lavaretus pidschianoides Pravdin,
pelet of Pallas, 1814: 412; type locality not 1931: 232 (type locality: Russia: Karelia: Rivers
stated, but Russia: River Yenisei) Vyg and Kem) from Berg, 1948: 395
? Coregonus lavaretus pidschianoides Pravdin, 1931:
Remarks on systematics. Russian authors (e.g., 232 (nomen nudum according to Eschmeyer,
Reshetnikov et al., 1997) recognize 4 ‘forms’, which 1998: 1338; locality: Russia: Vyg and Kem
partly occur in sympatry and which possibly are distinct rivers, Karelian coast of the White Sea)
species: 1) river peled; living in rivers, spawning in Coregonus lavaretus pidschian natio bargusini Krogius,
rivers, extending from Mezen to Yenisei drainages; 2) 1933: 85 (infrasubspecific, name not available;
lake-river peled; in same drainages but absent from locality: Russia: Barguzin River [tributary of
Ob River; 3) large lacustrine peled; spawning in lakes, Lake Baikal]) not seen, from Berg, 1948: 408
reaching up to 440 mm SL; 4) dwarf lacustrine peled; ? Coregonus lavaretus pidschian natio bergiellus
living and spawning in lakes, reaching up to 250 mm Svetovidov, 1934: 344 (infrasubspecific, name not
SL, with a shallower body and several rows of small available; locality: Russia: River Kara, Kara Bay)
black spots on body. It is presently not clear, which of ? Coregonus lavaretus pidschianoides Pravdin & Berg,
them is the ‘true’ C. peled. 1948: 13, fig. 12 (type locality: locality: Russia:
Vyg and Kem rivers, Karelian coast of the White
Distribution. Not native to Mongolia. Since 1978
Sea) adapted from Eschmeyer, 1998: 1338
introduced from Ulan Ude (Buryatia, Russia) to
? Coregonus lavaretus pidschianoides natio soldatovi
Lakes Naiman (Uburkhangai aimag), Ulaagchnii
Pravdin, in Pravdin & Berg, 1948: 15, fig. 14
Khar (Zavkhan aimag), Khongor-Ulen [a small lake
(infrasubspecific, name not available; Russia:
near Bayan Ulgii aimag ], and Thagaan (Darkhad
Lake Kildin, Kola basin) from Berg, 1948: 398
depression) (Dulmaa, 1979: 203). Outside
C. lavaretus pidschian natio delger-muren Dulmaa,
Mongolia: Arctic Ocean basin from Kolyma
1970 (infrasubspecific, name not available;
(eastern Siberia) westward to Mezen drainages.
locality: Mongolia: upper course of Delgermurun
River) from Dulmaa, 1973: 59
Coregonus pidschian
Salmo Pidschian Gmelin, 1789: 1377 (available by Remarks on biology and systematics. Coregonus
indication to Pallas, 1776b: 705; type locality: pidschian is usually treated as a subspecies of C.
not explicitly stated, but Russia: Siberia: River lavaretus in Russian literature. It is treated as a valid
Ob, is implied by statement in the description species here, following Bogutskaya & Naseka (2004:
of Salmo nasus in Pallas, 1776b: 705) 141) and for reasons discussed in Kottelat (1997:
Salmo Polcur Pallas, 1814: 400 (type locality: 118). Berg (1948: 394) recognized a large numbers
Russia: Siberia: from the Arctic Ocean migrates of local forms. They have been treated as synonyms
to the River Ob somewhat above Berezov) (and anyway some had only invalid names) by most
Coregonus sikus Cuvier, 1829: 308 (type locality: subsequent authors, but Bogutskaya & Naseka
rivers of Norway) (2004) recognize some as specifically distinct.
Coregonus baicalensis Dybowski, 1874: 389, pl. 7
figs. 1–3 (type locality: Russia: Lake Baikal) The ‘pidschian’ population of the Selenge is not
? Coregonus smitti Warpachovski, 1901: 414, pl. mentioned in the Russian literature I have accessed.
13 fig. 1 (type locality: Russia: Siberia: Lake Berg (1948) did not record C. pidschian from Lake
Teletskoe) Baikal, except (p. 408) for a ‘natio bargusini’ which
Coregonus fluviatilis Isachenko, 1925: 3 (type ascends the Barguzin River [tributary of Lake
locality: Russia: Yenisei drainage, Mana River) Baikal] for spawning.
21
In his work on the fishes of Transbaikalia, Karasev 2005); 3) The dwarf ‘form’ is cited by Dulmaa
(1987: 11) records that C. pidschian (as C. (1999: 198) who does not provide any additional
lavaretus pidschian) is present in the Lena drainage information on its biology.
but missing in the Baikal basin. He records “C. l.
pidschian natio baicalensis” from the Baikal basin. The fishermen in Darkhad depression report that
Berg (1948: 392) treated it as a subspecies of his since the introduction of C. peled, the populations
C. lavaretus but records it only from Ol’khon of C. pidschian (it is not known whether all forms
Island and the Maloe More strait north of the or a single one), Brachymystax lenok and Hucho
island. Bogutskaya & Naseka (2004: 136) treat C. taimen decreased sharply and that C. peled now
baicalensis as a valid species. constitutes about 90 percent of the catches.
It is certainly premature to conclude that the In Lake Dood Tsagaan, the catches of C. pidschian
Selenge population of C. pidschian is C. baicalensis, in 1997–2000 are said to have been 4000 kg in one
but this possibility should be investigated. If the day, with one 700 m seine, under ice; an analyzed
above distribution pattern is correct, the Selenge catch consisted of 60 C. peled, 2 C. migratorius,
‘pidschian ‘ is unlikely to be C. pidschian. and the rest was the native C. pidschian. By 2003,
C. pidschian had sharply declined, with the same
Berg (1948: 406) recognized a ‘natio’ fluviatilis effort yielding only 300 kg, mostly C. peled.
from the middle and upper Yenisei which is Coregonus pidschian prefers slow to strong current,
possibly (one of ) the species in the Darkhad while C. peled prefers standing waters (Erdenebat
depression; he did not mention Lake Baikal in M., pers. comm., July 2005).
its distribution. Coregonus ‘pidschian’ is native to
lakes of the Darkhad depression. Coregonus peled Presently it is not possible to determine if the
and C. migratorius were introduced from Ulan two forms are conspecific. The differences in
Ude in Russia, reportedly in 1985. Coregonus feeding and spawning habits suggest they might
migratorius apparently did not establish; the last be distinct. Considering the decrease of the
known observation (three individuals) was in 1997 population mentioned above (and the risk of
(information obtained by Erdenebat M.). hybridization with the introduced C. peled) this
should be investigated. (Although they normally
Three ‘forms’ of C. pidschian were originally have distinct spawning sites and periods, climatic
known from Lake Targan. There are a number variations and human alteration of the habitats
of reports containing quite contradictory could affect the spawning sites and periods).
information, and I use here the information
in the latest summary I found (Dulmaa, 1999; The population of Rivers Egiin and Uur are strictly
somewhat differing from the data in her 1973 riverine; they spend the summer in backwaters,
and 1995 works). 1) The lake ‘form’, which lives and spawn in the river.
in lakes and spawns in rivers. It starts migrating
to Sharga River in mid-August and returns to the Remarks on nomenclature. The author of C.
lakes in the second half of September. It feeds on pidschian is sometimes erroneously indicated as
bivalves and other benthic organisms. It reaches Pallas (1776b: 705). In that publication, Pallas
up to 60 cm long and has a fatty appearance and consistently indicated in the headings both the
rounded body; 2) The river-lake ‘form’, which Latin and local names. The local names were all
stays in lakes in spring and summer, enters the preceded by the name of the language in which
Shishkhed River in early October and remains it was used. Thus the heading for pidschian reads
there for some time after spawning and then “Salmo an Lavareti varietas? Ostiacis Pidschian.
returns to overwinter in lakes. It apparently feeds Samoiedis Polcur” [a Salmo variety related to
on plankton (cyclops); it is more slender. It reaches lavaretus? Pidschian in Ostiac language, Polcur in
up to 53 cm (Erdenebat M., pers. comm., July Samoyed language]. The name C. pidschian clearly
22
is not available from Pallas (1776b), but is first Remarks on systematics. This species appears as
available from Gmelin (1789). Thymallus arcticus in Mongolian literature (e.g.,
Baasanjav & Tsendayush, 2001). Russian authors
Dulmaa (1973: 59) use the name C. lavaretus recognize a number of subspecies within T. arcticus
pidschian natio delger-muren for the population (e.g. Berg, 1948: 422; Reshetnikov et al., 1997:
of River Delgermurun and refers to Dulmaa 696) but more recently others (e.g., Bogutskaya
(1970), but this does not appear to be published & Naseka, 2004: 146) recognize several of these
information. As it appears in Dulmaa (1973: 59) subspecies as distinct species: T. baicalensis, T.
the name is infrasubspecific, thus not available for brevipinnis, T. grubii, T. mertensii, and T. pallasi.
zoological nomenclature (Code art. 1.3.4).
Thymallus baicalensis inhabits Lake Baikal and the
Distribution. In Mongolia: lakes and rivers of Selenge drainage. Berg (1948: 426) considered
Gorlog drainage [upper Yenisei] in Darkhad that there was a deepwater ‘form’ which is now also
depression (Shishhed, Sharga and Tengis systems) treated as the species T. brevipinnis. Bogutskaya &
and in Selenge and its tributaries. Outside Naseka (2004) recognize the Selenge and Baikal
Mongolia: Arctic Ocean basin, from Finland ‘subspecies’ as a valid species (T. baicalensis) and
(Lapland) to eastern Siberia, Alaska, Canada the deepwater Baikal ‘form’ as another species, T.
eastward to Mackenzie drainage. brevipinnis. The populations of the Amur drainage
in Mongolia belong to T. grubii. This classification
agrees with the results of molecular studies by
Family Thymallidae Koskinen et al. (2002). This study shows T.
grubii and T. brevirostris as very distinct lineages,
(graylings) and that the remaining populations placed in T.
arcticus do not constitute a monophyletic lineage.
Thymallus arcticus It shows the populations from the Baikal drainage
Salmo (Truttac.) arcticus Pallas, 1776b: 706 (type (including Selenge) as a distinct lineage, quite close
locality: Russia: Sob River, a tributary of the to the populations from the Angara and middle
Ob, near Obdorsk [now Salekhard, lowermost Yenisei drainage. The Baikal drainage populations
Ob, 66°31’48”N 66°36’07”E] [see p. 35]) apparently are those called T. baicalensis by
Salmo digitalis Bloch, in Schneider, 1801: 421 Bogutskaya & Naseka (2004). It is not clear
(unnecessary replacement name for Salmo whether they include the Yenisei populations in
arcticus Pallas, 1776b: 706) their T. baicalensis.
Thymallus nikolskyi Kashchenko, 1899: 131 (type
locality: Russia: Altai: Ryblushka, a settlement This study also shows that the T. ‘arcticus’
close to Cherga on Rybnusska stream, Katun populations from North America and the Lena
drainage [Bogutskaya, pers. comm.]/Urusul drainage are distinct. The North American lineage
River at Ongudai/Tcharysh River at Ust-Kan/ had been called T. signifer by Russian authors (e.g.
Katun River at Nizhnii Uimon/Lake Talmenie/ Reshetnikov et al., 1997: 696 [as a subspecies]).
Tom River above Kusnetsk) The name of the Lena lineage is not yet clear;
Thymallus nikolskyi var. ongudajensis Kashchenko, Russian authors have used the name T. pallasii
1899: 134 (type locality: Russia: Altai: Urusul for it, but the status of the name depends on the
River at Ongudai) identity of the types, which apparently has not
Thymallus sellatus Kashchenko, 1899: 135, pl. 2 been investigated. Berg (1948: 428) had included
(type locality: Russia: Altai: Lake Tenga (Kenga), the Lena populations in T. pallasii. He recorded
Urusul River drainage) the type locality as Kolyma, but there is nothing
Thymallus arcticus arcticus natio alchutovi Johansen, in Cuvier & Valenciennes (1848: 448) allowing
1945: 6 (infrasubspecific, name not available; this conclusion to be reached; they only mention
locality: Russia: Lake Teletskoe) Russia as the origin.
23
Koskinen et al.’s (2002) study also shows samples based on the observation that the lower part of
of T. ‘arcticus’ from Shishkhed River as very both drainages constitutes the Western Siberian
distinct from the other populations of T. ‘arcticus’ Plain and the respective Thymallus populations
and apparently closely related to T. brevirostris. are relatively ‘close’, while the Shishked species is
Interestingly, this population is immediately in high altitude headwaters entering the plain in
distinguished from all other Thymallus in Mongolia an area diametrically opposite to the River Sob,
by its colour pattern (see below). This lineage and thus relatively ‘far’ from the type locality of
is considered a species distinct from the other T. arcticus. Again, this is a mere hypothesis which
Thymallus in Mongolia. Hereunder, it is called needs to be tested.
Thymallus sp. 1.
Whether all the populations of the Ob drainage
Two further groups of Thymallus populations have (including Irtysh) are conspecific is another open
been referred to as T. arcticus in Mongolia; one in the question. Earlier authors had recognized several taxa
Great Lakes Basin and one in the Irtysh headwaters among the Altai (upper Ob drainage) populations.
on the southern slope of Altai (Khurimt, Songinot, The type locality of T. thymallus (River Sob) is very
Yolt; Baasanjav & Tsendayush, 2001). There is no close to the mouth of the Ob, in the Arctic Ural,
information on the Irtysh populations either in on the Polar Circle. Considering the distance,
Mongolia or China, except for a description and the types of habitats in-between, the topography
figure in Anonym, 1979: 13, fig. 6), and only and the distribution pattern of Thymallus species
limited for Kazakhstan (Mitrofanov et al., 1986: in Eastern Siberia, it is reasonable to expect that
214). I could not find usable information on the the T. ‘arcticus’ from Altai, upper Irtysh and Great
populations of the headwaters of the Khovd and Lakes Basin could represent one or more additional
Zavkhan drainages. species. As this is only a guess and is currently
without supporting evidence, these populations of
Berg (1948: 424, fig. 253) illustrates a specimen course are recorded here as T. arcticus.
from the Irtysh at Ust-Kamenogorsk [now
Öskemen] in Kazakhstan and Anonym (1979:
fig. 6) figure a specimen in their book on fishes Thymallus baicalensis
of Xinjiang. They do not give locality data for the Thymallus Grubii var. baicalensis Dybowski, 1874:
figured specimen, but if the figure depicts a fish 391, pl. 8 fig. 1 (type locality: Russia: Lake
from Xinjiang, then it should be from Ertix, as this Baikal and Selenge and Angara rivers)
is the only Xinjiang drainage where they record the Thymallus arcticus baicalensis morpha angarensis
presence of this species (p. 63). Dorogostaisky, 1923: 77 (infrasubspecific, name
not available; locality: Russia: River Angara)
Beside the number of species now revealed among
Mongolian T. ‘arcticus’, a problem not yet solved
Remarks on systematics. See under T. arcticus.
is whether there are some ‘real’ T. arcticus in
Mongolia. The species was described from the
Sob, a tributary of the Ob, in Arctic Ural (Pallas, Distribution. In Mongolia: Selenge drainage.
1776b: 35). Berg (1948: 424) considered that Outside Mongolia: Lake Baikal and tributaries.
the ‘real’ T. arcticus inhabits the Kara, Ob and
Yenisei drainages, and the Great Lakes Basin of Thymallus brevirostris
Mongolia. Data are not available to objectively
decide which of the two Yenisei species mentioned Thymallus brevirostris Kessler, 1879: 306 (type
above (if any) is conspecific with the Ob species; locality: Mongolia: a tributary of Daingol
a reasonable guess could be that the lower Yenisei [Daingol Nuur = Lake Dayan, 48°23’00”N
species is more likely to be T. arcticus, but this of 88°50’00”E]/Dsapchyn River [Zavkhan River],
course requires confirmation. This hypothesis is a tributary of Lake Kara-Ussi [Lake Kar Us;
24
apparently error for Lake Khyar-gas]; also in Remarks on systematics. See discussion under
Kessler, 1880: 266) T. arcticus. Four ‘forms’ of Thymallus are present
Phylogephyra altaica Boulenger, 1898: 330, fig. (type in the Amur drainage. Knizhin et al. (2004)
locality: China: south side of Altai Mountains) distinguish three species among them; T. grubii
Thymallus brevirostris kozovi Dashdorj, Dulmaa & (with two ‘forms’: Upper-Amur and yellow-spots),
Tsendayush, 1968: 40 (type locality: Mongolia: T. burejensis (large-scale ‘form’) and T. sp. (lower-
Lakes Khoton and Khorgon) Amur ‘form’). The type series of T. grubii belongs
Thymallus brevirostris altaicus Dashdorj, Dulmaa & to the Upper-Amur ‘form’. Their analysis includes
Tsendayush, 1968: 45 (type locality: Mongolia: Lakes material of the Upper-Amur ‘form’ from the Onon
Khoton and Khorgon; junior secondary homonym and Ingoda rivers (type locality of T. grubii).
of Phylogephyra altaica Boulenger, 1898: 330) The Onon material had been obtained from the
Mongolian stretch.
Dashdorj et al. (1968) described T. brevirostris The Upper-Amur and yellow-spot ‘forms’ are
kozovi from Lakes Khoton and Khorgon, in the diagnosably distinct and have allopatric ranges.
headwaters of the Khovd drainage. Their figure It is not clear why Knizhin et al. consider them
and data and those in Dulmaa (1973: 61, 9) do conspecific, but it seems it is based on coefficients
not enable to distinguish it from other populations of differences (CD) results on a principal
of T. brevirostris. This taxon is not recognized in component analysis (PCA) of morphometric and
recent literature (e. g., Reshetnikov, 1997: 696; not meristic characters and on a mtDNA phylogeny.
mentioned Dulmaa, 1999, Bogutskaya & Naseka, A PCA indicates similarity of shape; while a
2004). Basaantjav & Tsendayush (2001: 41) and gross dissimilarity is a conclusive observation of
Sokolov (1983: 126) merely mention the name. distinctness, a similarity simply shows that the
All these authors comment that the graylings from analysed character(s) cannot be used to distinguish
Lakes Khoton and Khorgon in upper Khovd River groups. In fact their PCA of meristic characters
grow much larger than those of other populations shows only partial overlap. Similarly, CDs indicate
(up to 750 mm, vs. about 300 mm in others). degrees of similarity for given characters and are
never an argument on their own for conspecificity
The name T. b. kozovi is used on p. 38 of Dashdorj to be concluded. The same also applies to trees
et al. (1968) and the name T. b. altaicus on p. showing haplotype phylogeny. The tree in Knizhin
45 (Russian summary). The two names are used et al. shows a sister-group relationship between the
for the same taxon and the same type series and Upper-Amur and yellow-spotted ‘forms’, but the
respective precedence is determined by the first small degree of divergence is inconclusive. Certainly
reviser. In this case, however, T. b. kozovis has Knizhin et al. (2004: 68) are right when they state
precedence because T. b. altaicus is a secondary “Morphologically and genetically, the Upper-Amur
junior homonym of Phylogephyra altaica and and yellow-spotted ‘forms’ are closely related” but
therefore cannot be used. this is not enough to continue this sentence by
“and make up one species”. “Closely related” does
Distribution. Mongolia: lakes and rivers of Great not automatically imply conspecificity. To me,
Lakes Basin. Outside Mongolia: upper head of their data shows that the two ‘forms’ are distinct
Khovd [Cobdo] drainage in Russia. lineages, apparently fulfilling the criteria of species
under both the Evolutionary as well as Phylogenetic
Thymallus grubii Species Concepts.
Thymallus Grubii Dybowski, 1869: 955, pl. 18
fig. 9 (type locality: Russia: Onon and Ingoda
drainages. Outside Mongolia: Upper and middle
rivers, Amur River basin)
Amur drainage in Russia and China.
25
Thymallus nigrescens The nominal species T. arcticus dentatus described

from the area of Lakes Kara-Khol, Yenisei drainage,
Thymallus arcticus nigrescens Dorogostaisky, 1923:
is a name possibly available for the present species.
76 (type locality: Mongolia: Lake Khuvsgul)
However, it is described as having a “dark body
with a row of small dark spots”.
Distribution. Presently known only from
This species is endemic to Lake Khuvsgul. It lives in
Shishkhed River, Darkhad depression. Most likely
the lake but enters tributary streams for spawning.
occurs in upper Yenisei in Russia.
There are two spawning periods, one in mid May
to mid June in tributary streams, and the other
one in July-August along the shores (Dulmaa,
1999: 209; Tugarina, 2002; Erdenebat M., pers. Family Esocidae
comm., 2005). Research is needed to determine if (pikes)
the individuals spawning at the different seasons
and sites are morphologically and/or genetically Esox lucius
distinct. This is especially relevant since, after the Synonymy includes only nominal species whose
drought of 2002, only 20 out of 96 rivers with T. type locality is in Asia
nigrescens spawning grounds are left, and they are Esox Lucius Linnaeus, 1758: 314 (based on Artedi
now dry in May–July in dry years. If these two [1738: gen. 10 [53], syn. 26, spec. 52 [14],
spawning populations do not mix and breed only Esox rostro plagioplateo], Linnaeus [1746: 114,
among themselves, one of them might become lost n. 304, idem], and Gronovius [1754: 9, n. 28,
unnoticed if its spawning or nursery grounds are idem]; type locality: “in Europa”)
dry in May. Esox Reichertii var. baicalensis Dybowski, 1874:
392 (type locality: Russia: Siberia: all lakes and
Distribution. Endemic to Lake Khuvsgul. ponds of Lake Baikal basin [in Lake Baikal only
at the mouth of the tributaries])
Thymallus sp. 1 Esox Reichertii var. baicalensis Dybowski, 1874:
? Thymallus arcticus dentatus Gundriser, 1979b: 392 (type locality: Russia: Lake Baikal basin)
15 (type locality: Russia: Tuva: area of Lakes Esox lucius var. atrox Anikin, 1902: 109 (locality:
Kara-Khol in Kham-Syra River system, Bol’shoy Russia: Siberia: River Ob) from Berg, 1948:
Yenisei drainage) 458
Esox lucius bergi Kaganovskii, 1933: 4 (locality:
Remarks on systematics. See under T. arcticus. Russia: Siberia: River Anadyr) from Berg, 1948:
458
The Thymallus of the lakes in Darkhad depression Esox lucius lucius natio wiliunensis Kirillov, 1962:
(Yenisei drainage) have an orange caudal peduncle, 37 (infrasubspecific, name not available; locality:
dark body with indistinct yellowish area on middle Russia: Yakutia: River Vilyui basin)
of body, and indistinct black stripes. This sharp Esox lucius aralensis Pivnev, 1985: 18 (type locality:
colour contrast between the anterior part of the Kyrgyzstan: Chu River basin)
body and the caudal peduncle is apparently not
seasonal. This is also distinct in females, although Distribution. In Mongolia: Selenge drainage.
paler and somewhat less contrasted. The pattern Outside Mongolia: Most of Europe, Caspian Sea
is even more contrasted in spawning males, with basin, Siberia eastward to Anadyr drainage (Bering
bluish and greenish patches on the body. This Sea basin), North America.
pattern is seen also in specimens from the Chuluut
River [an upper tributary of Selenge], but much
less contrasted (Erdenebat M., pers. comm., July
2005).
26
Esox reichertii 23 [neotype withdrawal by Fricke, 2000: 639
not allowed by Code])
Esox Reichertii Dybowski, 1869: 956 (type locality:
Carassius carassius jacuticus Kirillov, 1956 (locality:
Russia: all lakes of the Onon and Ingoda river
Yakutia, Siberia) from Kirillov, 1962
systems [Lakes Tyrgituj, Sagtoj, Ustila, Baica
and others], rare in rivers, Amur River basin)
Remarks on systematics. The species is listed
as possibly present in the Bulgan River, e.g., by
Sokolov (1983: 199) and Baasanjav & Tsendayush
drainages. Outside Mongolia: Amur drainage and
(2001), but Sokolov explicitly stated that they
Sakhalin Island.
did not examine specimens from Mongolia and
considered records of its occurrence in Mongolia
incorrect (e.g., record from the Selenge by
Family Cyprinidae Dashdorj & Demin, 1977: 154). Their description
of the species is taken from Berg, 1949. I have
(carps, minnows) examined specimens labelled as C. carassius from
the Bulgan in the Zoological Museum in Berlin;
Acheilognathus asmussii unfortunately only the heads have been preserved
Devario Asmussii Dybowski, 1872: 212 (type and it is impossible to confirm the identification.
locality: Russia: Lake Chanka [Khanka]). The species is recorded in the Ertix [upper Irtysh]
Acanthorhodeus asmussii amurensis Holcik, 1962: but not from its tributary Ulungur River [lower
169, fig. 3 (type locality: Russia: Lake Kabar, Bulgan] by Li et al. (1966) and Anonym (1979),
Amur River near Yelabuga, about 60 km from while these authors record C. gibelio in both rivers.
Khabarowsk) Kimura et al. (1992) figure a specimen from the
lowermost Ulungur River. The species is tentatively
Remarks on systematics. This species is usually recognized as present in Mongolia, but this requires
placed in the genus Acanthorhodeus in the Russian confirmation.
literature (e.g., Reshetnikov et al., 1997: 698;
Naseka & Bogutskaya, 2004: 280; Bogutskaya & Distribution. In Mongolia: Bulgan River. Outside
Naseka, 2004: 40), which is considered a junior Mongolia: in Europe from Rhine drainage
synonym of Acheilognathus, following Arai & Akai eastwards (absent in Mediterranean basin, present
(1988). in Black Sea basin); in Asia, Caspian basin and
rivers flowing to the Arctic Ocean, eastwards to
Distribution. In Mongolia: Kherlen, Onon and Kolyma drainage.
Khalkh drainages, and Lake Buir. Outside Mongolia:
Amur drainage in Russia and China; Korea.
Carassius gibelio
Carassius carassius
Synonymy includes only nominal species whose
type locality is in Asia
type locality is in Asia
Cyprinus Gibelio Bloch, 1782: 71, pl. 12 (type
Cyprinus Carassius Linnaeus, 1758: 321 (based
locality: Churmark, Pommern, Schlesien and
on Artedi [1738: gen. 4 [29], syn. 5, spec. 29
Preussen (Prussia, now Germany and Poland))
[4], Cyprinus pinnae dorsi ...], Linnaeus [1746:
Carassius vulgaris var. kolenty Dybowski, 1877: 11
122, n. 322, idem], Gronovius [1754: 3, n. 11,
(type locality: Russia: River Amur basin)
idem; 1746: 75, n. 55, Cyprinus hamburger;
Carassius auratus gibelio morpha vovkii Ioganzen,
Wawerveen, Belgium]; type locality: Germany:
1945: 12 (infrasubspecific, name not available;
Baden-Württemberg: Neckar River at Heidel-
locality: Russia: Siberia: lakes of Baraba steppe
berg, by neotype designation by Fricke, 1999:
and Narym region)
27
Remarks on systematics. This species was stocks are known in Japan which are considered as
originally described from Europe. Its status is still distinct species (Teitler & Fujita, 1993; Hosoya, in
not clear. Several authors consider it as a ‘form’ of Nakabo, 1994: 212–213) and are treated as species
the well-known goldfish (Carassius auratus), either here under the ESC. With the present data, the
a wild ‘form’ native to Europe or an introduced Mongolian populations cannot be considered as
‘form’, or as feral stock of introduced goldfishes, conspecific with the cultivated C. auratus.
or as a result of hybridization. There is indication
that the species might have been present in Europe The discussion by Vasil’eva & Vasil’ev (2000)
before the first introduction of goldfishes from cannot be followed as it is based on a heterogeneous
Japan, which would rule out the hypothesis of the assemblage of partly second-hand data on
feral goldfish (Kottelat, 1997). morphology, genetics, ecology, and zoogeography.
Further, the authors have apparently been largely
The problem is made very complex and the original disserved by the translators and the editors of
distribution in Europe will probably never be known the journal and the paper is marred by linguistic
exactly because of introduction, transplantation, problems, making it very difficult to understand.
confusion with C. auratus and complex modes of Without inclusion of European material, their
reproduction, with diploid populations of both discussion of the identity of C. gibelio does not
sexes, as well as populations made of diploids and make sense, as the type locality is in Europe.
tetraploids, or female-only triploid populations.
The species is now invasive throughout Europe Distribution. In Mongolia: Selenge, Onon
and is a pest where it establishes. It seems that the and Kherlen drainages, and Lake Buir. Outside
invasive fishes result from populations stocked in Mongolia: western limit in Europe not clear (see
eastern Europe and imported from Siberia and above), in Asia, extends eastwards at least to the
are likely to be conspecific with the Mongolian Amur drainage, exact limits not clear.
populations.
Chanodichthys erythropterus
Although this may seem distant and of no interest
in a Mongolian context, in fact it is relevant because Culter erythropterus Basilewsky, 1855: 236, pl. 8
the name of the species present in Mongolia depends fig. 1 (type locality: China: rivers draining to
on the identity of the populations originally present Gulf of Tschili)
in Europe. The invasion from populations stocked Culter ilishaeformis Bleeker, 1871: 67, pl. 10 fig. 1
in eastern Europe of a species earlier cryptic or (type locality: China: Yangtze River)
poorly known in central Europe suggests that either Culter Sieboldii Dybowski, 1872: 214 (type
the identification of the original C. gibelio presents locality: Russia: middle course of Amur, Ussuri,
some problem, or that the invasive populations are Sungatschi and Chanka)
not conspecific with the original one. This problem Culter aokii Oshima, 1919: 250, pl. 52 fig. 1
is presently being investigated, and awaiting a (type locality: Taiwan: Jitsugetsutan, Lake
solution, I retain the name C. gibelio for the species Candidius)
present in Mongolia.
Remarks on nomenclature. For some time,
Russian and Mongolian literatures consider the the species identified as Culter erythropterus (or
local C. gibelio as a subspecies of the goldfish C. Cultrichthys erythropterus) and Culter alburnus have
auratus. The systematics of the genus Carassius been interverted in the Chinese and Southeast
in East Asia is confusing. The ancestor of today’s Asian literature. I have not attempted to track
domesticated goldfishes was introduced to Japan the source of this interversion, but it goes back at
from China at a date between 1502 and 1748 least to Wu et al. (1964: 113) and is still going
(Okada, 1959–60: 531). Available data show that on (e.g., Chen, 1998: 182; Kottelat, 2001b: 21).
at least five genetically and morphologically distinct See Bogutskaya & Naseka (1996: 21, 2004: 54)
28
for discussion. The species listed in the synonymy Culter alburnus
of C. “erythropterus” by Chen in fact are synonyms
Culter Alburnus Basilewsky, 1855: 236, pl. 8 fig. 3
of C. alburnus. Correcting this error has an impact
(type locality: China: rivers draining to the Gulf
on the genus-level nomenclature.
of Tschili)
Culter brevicauda Günther, 1868: 329 (type
The type species of Culter is C. alburnus. The
locality: Taiwan)
type species of Cultrichthys is C. brevicauda. As C.
Culter Kneri Bleeker, 1870: 252 (nomen nudum)
brevicauda is a subjective junior synonym of C.
Culter Kneri Bleeker, 1871:14 (based on Culter
alburnus, Cultrichthys is a subjective junior synonym
erythropterus of Kner, 1867: 360, pl. 14 fig. 4;
of Culter. The species placed in Erythroculter by
type locality: China: Shanghai)
Berg (e.g., 1949: 804) and other Russian authors
Culter tientsinensis Abbott, 1901: 489, fig. (type
should be called Chanodichthys as this name is
locality: China: Hebei: River Pei-Ho at Tien-
older than Erythroculter.
Tsin [Tianjin])
Distribution. In Mongolia: Lake Buir. Outside
Remarks on nomenclature. See under Chano-
Mongolia: from Amur to Red River drainages
dichthys erythropterus.
(China to Vietnam); Taiwan; Hainan.
Distribution. In Mongolia: Lake Buir and Onon
Chanodichthys mongolicus and Kherlen drainages. Outside Mongolia: from
Leptocephalus Mongolicus Basilewsky, 1855: 234, Amur to Red River drainages, Taiwan, Hainan.
pl. 4 fig. 2 (type locality: “in winter, brought to
Beijing frozen from Mongolia and Manchuria”; Ctenopharyngodon idella
spelt mongolensis on pl. 4 fig. 2; first revisers Leuciscus idella Valenciennes, in Cuvier &
[Bogutskaya & Naseka, 2004: 54] gave Valenciennes, 1844: 362 (type locality: China)
precedence to mongolicus) Leuciscus Tschiliensis Basilewsky, 1855: 233 (type
Culter Mongolicus Basilewsky, 1855: 237 (type locality: China: Gulf of Tschili and tributary
locality: “in winter, brought to Beijing frozen streams)
from Mongolia”; simultaneous secondary Ctenopharyngodon laticeps Steindachner, 1866a:
homonym of Leptocephalus mongolicus Basi- 782, pl. 18 figs. 1–5 (type locality: China: Hong
lewsky, 1855: 234, first reviser [Banarescu, Kong)
1972: 387] gave precedence to Leptocephalus Sarcocheilichthys teretiusculus Kner, 1867: 356 (type
mongolicus) locality: China: Shanghai/waters near Tianjin
? Culter Pekinensis Basilewsky, 1855: 237 (type and draining to Gulf of Tschili [Basilewsky’s
locality: China: streams draining to the Gulf of material])
Tschili) Pristiodon siemionovii Dybowski, 1877: 26 (type
Culter rutilus Dybowski, 1872: 214 (type locality: locality: Russia: Amur River, Ussuri River,
Russia: Ussuri and Chanka) Sungacha River, Lake Khanka and Sungari
Erythroculter mongolensis elongatus He & Liu, River)
1980: 483, fig. (type locality: China: Yunnan:
Lake Chenghai) Distribution. There is a single record from
Erythroculter mongolicus qionghaiensis Ding, 1990: Mongolia, around 1962 (Sokolov, 1983: 163),
246, fig. 1 (type locality: China: Sichuan: Lake in Lake Buir; the species is widely cultivated in
Qionghai, 27°53’N 102°18’E) China, and this fish likely was stocked, or escaped,
on the Chinese side of the lake. Outside Mongolia:
Distribution. In Mongolia: Lake Buir, and Onon native to East Asia, in lower and middle stretches
and Kherlen drainages. Outside Mongolia: from of major rivers from Amur to Xi Jiang drainages.
Amur to Yangtze drainages.
29
Cyprinus rubrofuscus used to produce hybrids with other species of the

genus Cyprinus.
Cyprinus rubro-fuscus La Cepède, 1803: 530, pl. 16
fig. 1 (type locality: China)
Although the diversity of the East Asian species of
Cyprinus nigro-auratus La Cepède, 1803: 547, pl.
Cyprinus has long been documented in the Chinese
16 fig. 2 (type locality: China)
literature (e.g., Wu et al., 1977; Yue, 2000), it is
Cyprinus viridi-violaceus La Cepède, 1803: 547, pl.
largely ignored, if not flatly negated, in western
16 fig. 3 (type locality: China)
literature (e.g., Balon, 1995; Banarescu & Paepke,
Cyprinus anna-carolina La Cepède, 1803: 544, pl.
2002), where it is often assumed that Asian carps
18 fig. 1 (type locality: not stated)
are derived from introduced European carps and
Cyprinus floripenna van Hasselt, 1823: 132
that the species recognized by Chinese authors are
[translated in Alfred, 1961: 85], 1824: 375
domesticated or feral forms. Kottelat (1997: 57;
(nomen nudum, Kottelat, 1987: 370)
2001a: 22, 2001b: 45) disagrees with this and
? Cyprinus flavipinnis Valenciennes, in Cuvier &
commented that the domesticated carps in East
Valenciennes, 1842: 71, fig. 457 (type locality:
Asia are one or more species distinct from the
Indonesia: Java: Buitenzorg [Bogor])
European one. The molecular data of Zhou et al.
Cyprinus haematopterus Temminck & Schlegel,
(2004) give support to this conclusion (although
1846: 189, 216, pl. 96 (type locality: Japan:
these authors did not note the taxonomic aspect of
large rivers of Kiusiu Island; junior primary
their results).
homonym of Cyprinus haematopterus Rafinesque,
1820a: 6) Remarks on nomenclature. The name C.
Cyprinus atro-virens Richardson, 1846: 287 (type haematopterus is used in the Russian and Chinese
locality: China: Canton) literature for the common Asian carp. This name
Cyprinus flammans Richardson, 1846: 288 (type (created by Temminck & Schlegel in 1846) is
locality: China: Canton) not valid for the Asian carp because it is a junior
Cyprinus hybiscoides Richardson, 1846: 289 (type homonym of Cyprinus haematopterus created by
locality: China: Canton) Rafinesque in 1820 for a North American fish.
Cyprinus sculponeatus Richardson, 1846: 290 (type The oldest name has priority and the youngest
locality: China: Canton) one is not valid. Anyway, C. haematopterus is not
Cyprinus ? fossicola Richardson, 1846: 291 (type the earliest name given to this species; the earliest
locality: China: Canton) available name for the common Asian carp is C.
Cyprinus carpio var. mürgo Dybowski, 1869: 950 rubrofuscus, a name created by La Cepède in 1803
(type locality: Russia: Transbaikalia: Onon River; (see Kottelat, 2001b: 45).
spelled murgo p. 946 [emendation as muergo (e.g.
in Eschmeyer, 1998) is erroneous as the name Distribution. In Mongolia, C. rubrofuscus is native
is not derived from a German word; Dybowski in the Amur drainage (Onon, Kherlen). Introduced
explicitly stated it is the local vernacular name in the Selenge drainage in the mid-1940s, invasive.
of the fish, Code art. 32.5.2.1]) It is not known if the introduced stocks are pure C.
Cyprinus carpio yuankiang Wu, Yang, Yue & rubrofuscus. Outside Mongolia: from Amur to Red
Huang., 1963: 43 (type locality: China) River drainages, although it is probably impossible
Cyprinus carpio triangulus Wu, Yang, Yue & Huang, to determine its exact original range.
1963: 43 (type locality: China)
Eupallasella percnurus
Remarks on systematics. There is no native
Cyprinus carpio in Mongolia, only C. rubrofuscus Synonymy includes only nominal species whose
(the C. haematopterus of Russian and Chinese type locality is in Asia
literature) is native. Cyprinus carpio is native only Cyprinus Percnurus Pallas, 1814: 299 (type locality:
to the Black, Caspian and Aral Seas basins, and it Russia: Siberia: lakes and swamps around River
is introduced elsewhere. Cyprinus carpio has been Lena)
30
Leuciscus dauricus Valenciennes, in Cuvier & and Rhynchocypris as valid. This is apparently also
Valenciennes, 1844: 149 (type locality: Russia: the conclusion reached by Bogutskaya & Naseka
waters of Daouria [Transbaikalia]; based on (2004: 92), except that they retain Rhynchocypris
Cyprinus rutilus of Pallas, 1814: 317) and Eupallasella as subgenera of Phoxinus. A recent
Phoxinus jelskii Dybowski, 1869: 952 (type molecular study suggests that Eupallasella should
locality: Russia: Siberia: lakes of Darasun, But- be included in Rhynchocypris (Sakai et al., 2006).
Durutaj, Ila and Makhojtowa valleys [River As long as the interrelationships of the three genera
Onon basin]) are not established, I do not see reason to consider
Leucaspius Fischeri Sabaneev, 1871: 277 (nomen them congeneric.
nudum; locality: Russia: Siberia: east slope of
Ural range) Remarks on nomenclature. There has been
Phoxinus perenurus var. Dauricus Dybowski, 1877: disagreement in the correct spelling of the species’
17 (type locality: Russia: Transbaikalia: lake in name, some authors using percnurus and others
basin of River Onon) using perenurus (Kottelat, 1997: 59). In fact,
Phoxinus stagnalis Warpachowski, 1886: 76, Bogutskaya et al. (2005: 93) showed that the
fig. (type locality: Russia: Kazan Prov.: Lake spelling using in the first copies of the book was
Schumjer [in basin of River Malaya Kokschaga, percnurus and that the letter ‘c’ became damaged
Berg, 1949: 578]) and was erroneously replaced by a letter ‘e’ by the
Phoxinus Sabanejewi Warpachowski, 1887a: 535 typesetter. [In 1814 printing methods were not
(type locality: Russia: lakes on eastern slope of those of today].
Urals range [district Schadrinsk, Tscheljabinsk; Distribution. In Mongolia: Selenge and Amur
Berg, 1912: 199]; also in Warpachowski, 1887b: drainages. Outside Mongolia: lakes in Arctic
688) Ocean basin (from Northern Dvina to Kolyma
Phoxinus altus Warpachowski, 1887b: 535 (type drainages) and Pacific basins (from Amur to Korea
locality: Russia: Siberia: tributaries of River and Japan); in Europe, disjunct distribution: lakes
Yenisei, Siberia [lower Tunguska, Berg; 1912: in Odra, Vistula, Dniepr, Volga drainages.
199]; also in Warpachowski, 1867b: 688)
Phoxinus variabilis Warpachowski, 1887b: 535
(type locality: Russia: Siberia: tributaries of Gnathopogon strigatus
River Ob [River Tscharysch; Berg, 1912: 199]; Leucogobio strigatus Regan, 1908: 59, pl. 2 fig. 2
also in Warpachowski, 1867b: 688) (type locality: Korea: Chong-ju)
Phoxinus percnurus mantschuricus Berg, 1907a: Gobio taeniatus mantschuricus Berg, 1914: 481,
204 (type locality: China: Amur drainage: Da- fig. 72 (type locality: Manchuria: River Schansi,
tschu-an, a tributary of River Sungari) tributary of River Hailin, at railway line,
Phoxinus crucifer Gratzianow, 1907: 128 Sungari basin/River Ussuri downstream of
(type locality: Russia: Buryatia: vicinity of Lontschakowskij)
Troitskosavsk, River Selenge drainage/small lake Paraleucogobio soldatovi Berg, 1914: 486, fig. 74
at Bain Bulyk, 20 verst of Troitskosavsk) (type locality: Russia: mouth of Ussuri River)
Phoxinus percnurus sarykul Ruzskii, 1926: 112,
fig. (type locality: Russia: Siberia: Lake Sarykul, Remarks on systematics. Synonymy follows
south of Chelyabinsk) from Berg, 1949: 575 Bogutskaya & Naseka (2004: 63). Chen (1998:
308) placed this species in the genus Paraleucogobio
Remarks on systematics. Eupallasella percnurus while they considered that Gobio taeniatus
was earlier placed in Phoxinus. Phoxinus is quite a mantschuricus is a valid species of Gnathopogon.
heterogeneous assemblage and a number of authors
have tried to divide the species into a variety of genera Distribution. In Mongolia: Lake Buir. Outside
(e.g., Dybowski, 1916; Gasowska, 1979; Howes, Mongolia: Amur drainage in Russia and China,
1985). I follow Howes in recognizing Eupallasella Korea, River Liao in China.
31
Gobio acutipinnatus Distribution. In Mongolia: Bulgan River. Outside

Mongolia: Upper Irtysh drainage in China and
Gobio gobio acutipinnatus Menshikov, 1939: 131, Kazakhstan. Possibly present further downriver in
pl. (type locality: Kazakhstan: Lake Marka-kul, Irtysh.
Irtysh basin)
Gobio cynocephalus
Remarks on systematics. Gobio acutipinnatus, G.
cynocephalus and many others have traditionally Gobio fluviatilis var. cynocephalus Dybowski, 1869:
been treated as subspecies or synonyms of G. gobio, 951 (type locality: Russia: Amur drainage:
which was considered to be a species extending Onon and Ingoda rivers)
from Spain to northeastern China (e.g., Berg, Gobio liaoensis Mori, 1927: 31 (type locality:
1949; Banarescu & Nalbant, 1973; Reshetnikov et China: Manchuria: Tai-tzu River, tributary of
al., 1997; Kottelat, 1997). Detailed examination Liao River at Chiao-tao)
of several of the populations in Europe has shown
that G. “gobio” in fact is an assemblage of several Remarks on systematics. Chen (1998: 293) treats
species, and that some even belong to the genus G. macrocephalus as a valid species, while Banarescu
Romanogobio. This is based on morphological as & Nalbant (1973: 128) treat it as a junior synonym
well as molecular characters (see, e.g., Doadrio & of Gobio soldatovi and Kim (1997: 217) treats it as
Madeira, 2004; Kottelat & Persat, 2005; Freyhof & a junior synonym of G. cynocephalus. The original
Naseka, 2005; Yang et al., 2006). Chinese authors description of G. macrocephalus by Mori (1930:
also recognize that the local G. “gobio” are also 46) is not very informative but indicates that the
an artificial assemblage and recognize a number head length is 3.6 times in SL; Chen (1998: 293)
of species (e.g., Chen, 1998). They recognize G. gives 3.3–3.8 for G. macrocephalus from Tumen
acutipinnatus as a valid species, occurring in China (type locality), 3.8–4.2 for G. soldatovi and 3.8–4.2
in the Ertix drainage (Chinese name of the upper for G. cynocephalus. Banarescu & Nalbant (1973:
Irtysh). 128) give 3.6–4.3 for G. soldatovi. The figure of G.
cynocephalus in Kim (1997: 217) shows the head
Gobio acutipinnatus was originally described from length about 4.4 times in SL; the origin of the
Lake Marka-kul, a small lake in the Irtysh drainage, specimen is not stated. With the available data, I
in Kazakhstan, close to the border with China. The exclude G. macrocephalus from the synonymy of G.
Bulgan River [Ulungur in China] is a tributary cynocephalus and consider it as a valid species.
of the Ertix. I have examined various samples of
Gobio from the Bulgan and Ertix, and the syntypes Distribution. In Mongolia: Onon, Kherlen and
from Lake Marka-kul in Zoological Institute, St. Khalkhiin rivers, Lake Buir. Outside Mongolia:
Petersburg, and I agree with Chen (1998: 294) that Amur drainage in Russia and China; River Liao
they belong to a distinct species. Records of both He in China.
G. gobio and G. g. cynocephalus from the Bulgan in
Baasanjav & Tsendayush (2001: 95, 96) refer to Gobio sibiricus
this species. Gobio gobio sibiricus Nikolski, 1936: 470 (type
locality: Kazakhstan: Akmolinskaya Oblat:
Gobio acutipinnatus is distinguish from other Nura River, where it enters Lake Dzhanybek
Gobio in Mongolia by the combination of the /Russia: Krasnoyarskiy Krai: Minusinskiy
following characters: anus midway between pelvic- District: Yenisei, Minusinkaya channel)
fin base and anal-fin origin; throat naked in front ? Gobio gobio magnicapitata Gundriser, 1967: 70
of pectoral-fin base; 12–16 [usually 14] scale rows (type locality: Russia: Tuva: Lake Kara-Khol
around caudal peduncle; snout blunt; head depth basin, Kham-Syra river system, Bol’shoy Yenisei
about 1.5 times in its length; a mid-lateral row of drainage, 53.43°N 95.25°E [Gundriser, 1979a:
10–14 dark spots; 40–43 total lateral line scales. 9])
32
Remarks on systematics. The map in Baasanjav Distribution. In Mongolia: Selenge drainage.
& Tsendayush (2001: 99) shows G. cynocephalus Outside Mongolia: Ob (except Irtysh), Yenisei and
at different localities in the Selenge drainage. It Nura drainages.
is not recorded in the Selenge by Sokolov (1983:
168). The map in Reshetnikov et al. (2002: 250) Gobio soldatovi
shows no species of Gobio in Lake Baikal basin
Gobio gobio var. soldatovi Berg, 1914: 461, fig.
(including Selenge). Banarescu & Nalbant (1973:
63 (type locality: Russia: lower Amur River at
133) consider that the range of G. cynocephalus is
Tscheptschiki [Chepchiki])
restricted to the Amur drainage and northeastern
? Gobio gobio tungussicus Borisov, 1928: 105,
China and (p. 125) that the earlier records of
165, pl. 6 figs. 14–15 (type locality: Russia:
G. cynocephalus in the upper Yenisei and Baikal
Sakha-Yakutia: Lena River near Zhigansk)
drainages are G. gobio sibiricus. Sideleva (2003: 10)
records G. g. cynocephalus in Lake Baikal basin.
Remarks on systematics. Gobio soldatovi was
first recorded from Lake Buir by Baasanjav
Nikolski (1936) described G. g. sibiricus on the
& Tsendayush (2001: 97). Berg (1949: 651),
basis of material from several localities in the
Nikolski (1956) and Banarescu & Nalbant (1973:
Yenisei, Ob and Nura drainages. Berg (1949: 645)
129) recorded it only from the lower Amur
commented that the Yenisei and Nura specimens
drainage (downriver of Khabarovsk). Reshetnikov
are not conspecific and he treated syntypes from
et al. (1997: 701; 2002: 252) record it from the
the Yenisei (Minussinsk) as G. g. cynocephalus
whole Amur drainage, but this includes the G. s.
and syntypes from the Nura (a recently formed
“tungussicus” of Nikolski (1956), Sokolov (1983),
endorheic drainage, still occasionally connected
and Baasanjav & Tsendayush (2001: 171), which
with the Irtysh drainage) as G. g. lepidolaemus.
is not conspecific (see under Gobio sp. Onon) and,
Banarescu & Nalbant (1973: 125) examined
on the basis of the available data, the presence of
material from Nura and treated the two populations
the real G. soldatovi in the upper Amur remains to
as conspecific, as C. g. sibiricus, and distinct from
be confirmed.
G. g. cynocephalus. They, however, consider the
Nura specimens as ‘intergrades’ between their G.
If G. soldatovi is naturally absent from the upper
g. sibiricus and G. g. lepidolaemus.
Amur, the resulting range (Lena and lower Amur
drainages) would be strangely disjunct and the
Banarescu & Nalbant (1973: 134) distinguished
synonymy and status of the nominal taxa G. g.
their G. g. sibiricus from G. cynocephalus by lateral
tungussicus should be re-examined (see under Gobio
line scale count (40–44 in G. g. sibiricus, vs. 43–
sp. Onon).
48) and shape of dorsal fin (distal edge more-or-
less straight, vs. distinctly ‘notched’ [concave]).
Even if absent in upper Amur, the presence (and
Without access to specimens, the difference
apparently recent discovery) of G. soldatovi in
seems convincing and I treat them as distinct. I
Lake Buir could be explained by introduction or
use the name G. sibiricus for the Yenisei and Ob
accidental release, together with many other species,
populations. As there is a possibility that these
on the Chinese side of the lake. The natural habitat
two populations may be distinct from the Nura
of the species in lower Amur is slow flowing and
population, a lectotype designation will be needed
standing water.
to definitively fix the name to the Yenisei and Ob
populations. Clearly, this would be premature at
this stage.
Mongolia: Amur drainage in Russia and China;
Sakhalin Island; ? Lena drainage in Russia.
It remains to compare material from the Yenisei
and Selenge to confirm that they are effectively
conspecific, but I tentatively accept it.
33
Gobio tenuicorpus Gobio sp. Onon

Gobio gobio tenuicorpus Mori, 1934: 13, pl. 4 fig. 2a Remarks on systematics. Sokolov (1983: 171)
(type locality: China: Hopei: Hsing-lung-shan) and Baasanjav & Tsendayush (2001: 97) recorded
G. soldatovi tungussicus from the Onon drainage
Remarks on systematics. Listed as Gobio in Mongolia. Reshetnikov et al. (1997: 701) con-
albipinnatus tenuicorpus by Holcik & Pivnicka sidered that this ‘subspecies’ is restricted to the Lena
(1969: 7) and Sokolov (1983: 169), and “Gobio drainage. Bogutskaya & Naseka (2004: 65) treated
albipinnatus Mori, 1934” by Baasanjav & G. g. tungussicus as a synonym of G. soldatovi.
Tsendayush (2001: 94). Gobio albipinnatus Without access to specimens, it is impossible to
is a different species from the Volga and Ural know which is the species reported by Sokolov
drainages, placed in the genus Romanogobio (1983) and Baasanjav & Tsendayush (2001). The
(Bogutskaya & Naseka, 2004: 70). Gobio description and illustration in Sokolov are from
tenuicorpus was also moved to Romanogobio Nikolski (1956) and based on specimens from “a
by Banarescu & Nalbant (1973: 143), who small unnamed lake in the Shargol’dzhin River
considered Romanogobio as a subgenus of Gobio. system (a tributary of Ingoda), in Bal’zinskoye
Romanogobio is now considered as a valid genus Lake in the Tura River system, a tributary of Ingoda
(Bogutskaya & Naseka, 2004: 70), also including River, and in lake-like puddles of Onon River at
the former subgenus Rheogobio as a synonym (see Novo-Kazachinskoye”, in Russia.
Naseka & Freyhof, 2004). The genus Romanogobio
includes species from the Black Sea and Caspian Gobio gobio tungussicus was originally described
Sea basins and species from East Asia, a strange from the Lena River drainage (Borisov, 1928: 105).
zoogeographic pattern; their actual relationships Nikolski’s figure of G. s. tungussicus shows a fish
require further investigation. Indeed a molecular quite different from that of Borisov (reproduced
analysis of the relationships within Gobioninae in Berg, 1949: 646). It shows a deeper bodied fish,
shows G. tenuicorpus included in the Gobio with a more slender caudal peduncle, a blunt snout
lineage and clearly distinct from the Romanogobio and a colour pattern made of a row of blotches
lineage (Yang et al., 2006). along the flank, while Borisov’s drawing shows a
dark longitudinal band. A topotype of G. soldatovi
Gobio tenuicorpus is known from the Amur to the figured in Berg (1949: 650) looks similar to G.
Luang Ho drainages. In the Amur drainage, it has a s. tungussicus. Bogutskaya & Naseka (2004: 65)
broadly disjunct range, with the lower and middle consider them as synonyms.
stretches on the one hand and Lake Buir on the
On the basis of the published data, the fish
other hand (e.g., Reshetnikov et al., 2002: 316).
figured by Nikolski (1956) as G. s. tungussicus is
Sokolov (1983: 170) compared the data of Holcik
misidentified. I am unable to identify it as any of
& Pivnicka (1969: 7) with materials from the delta
the species of Gobio recorded from the Amur in the
of stream Khalkhiin and with materials from Amur
Russian and Chinese literature.
and found no morphological differences. Their
description of the species is taken from Nikolski Distribution. In Mongolia: Onon River drainage.
(1956: 185). It is not clear whether this disjunct Outside Mongolia: Onon and Ingoda drainages in
range reflects a collecting bias, or reality. The data Russia.
and drawing in Sokolov (1983: 170) are those
of Nikolski (1956: 185) based on material from
Hemibarbus labeo
middle and lower Amur.
Cyprinus labeo Pallas, 1776a: 207 (nomen nudum),
Distribution. In Mongolia: In River Khalkhiin 1776b: 703 (type locality: Russia: “streams
and in Lake Buir. Outside Mongolia: from Amur flowing through Dauria and draining to the
to Luang Ho drainages. Amur” [p.207]/Dauria is mentioned p. 703 as
34
“Russis in Dauria [kon] (Equus)” which means ? Hemibarbus longibarbis Fang, 1938: 269 (type
“called [kon] (horse) by the Russians of Dauria”]; locality: China: Kiangsi: Sau-hsui)
also in Pallas, 1778: appendix, p. 14)
Gobio barbus Temminck & Schlegel, 1846: 198, Remarks on systematics. The genus Hemibarbus
pl. 99 fig. 1 (type locality: Japan: Nagasaki; was revised by Yue (1995; Yue, in Chen, 1998:
junior secondary homonym of Cyprinus barbus 491).
Linnaeus, 1758: 320, when placed in Barbus by
Günther, 1868: 135) Distribution. In Mongolia: drainages of Lake Buir
? Barbus abramoides Brandt, in Maak, 1861: 196 and Rivers Onon, Kherlen and Khalkhiin. Outside
(nomen nudum; locality: Russia, Ussuri) Mongolia: from Amur to Yangtze drainages (Russia,
Barbus schlegelii Günther, 1868: 135 (replacement China) and Taiwan.
name for Gobio barbus Temminck & Schlegel,
1846: 198) Hemiculter leucisculus
Acanthogobio oxyrhynchus Nikolski, 1904: 358
Culter Leucisculus Basilewsky, 1855: 238 (type
(type locality: Russia: Lake Chanka at mouth of
locality: China: streams draining to Gulf of
River Santacheza)
Tschili)
Pseudogobio chaoi Evermann & Shaw, 1927: 106
Squaliobarbus annamiticus Tirant, 1883: 97 (type
(type locality: China: Nanking)
locality: Vietnam: River of Hué)
Hemibarbus longianalis Kimura, 1934: 123, pl. 4
Culter Balnei Sauvage, 1884: 213, pl. 8 fig. 4 (type
fig. 1 (type locality: China: Sichuan: Suining
locality: Vietnam: vicinity of Hanoï)
and Howchwan)
Hemiculter Schrencki Warpachowski, in Warpa-
chowski & Herzenstein, 1887: 46, pl. fig. 4
Remarks on systematics. The genus Hemibarbus was
(type locality: China: Fu-Tschau [Fuchow]; also
revised by Yue (1995; Yue, in Chen, 1998: 491).
in Warpachowski, 1888: 18)
Hemiculter kneri Warpachowski, 1888: 17 (based
Distribution. In Mongolia: drainages of Lake Buir
on Culter leucisculus of Kner, 1867: 362; type
and Rivers Onon, Kherlen and Khalkhiin. Outside
locality: China: Shanghai)
Mongolia: from Amur to Mingjiang drainages
Hemiculter kneri Kreyenberg & Pappenheim,
(Russia, China) and Taiwan.
1908: 105 (based in part on Culter leucisculus of
Kner, 1867: 362; type locality: China: Shanghai
Hemibarbus maculatus
and Hankau; junior primary homonym of
Hemibarbus maculatus Bleeker, 1871: 19, pl. 4 fig. Hemiculter kneri Warpachowski, 1888: 17)
3 (type locality: China: Yangtze River; junior Parapelecus eigenmanni Jordan & Metz, 1913: 21,
secondary homonym of Barbus maculatus pl. 3 fig. 1 (type locality: Korea: Suigen, south
Valenciennes, in Cuvier & Valenciennes, of Seoul)
1842: 195, when placed in Barbus by Günther, Cultriculus akoensis Oshima, 1920: 132, pl. 3 fig. 4
1889: 224; these taxa are no longer considered (type locality: Taiwan: Ako)
congeneric and the substitute name [Barbus Hemicultur [sic] clupeoides Nichols, 1925c: 7 (type
semibarbus Günther, 1889: 224] is not in use, locality: China: Hunan: Lake Tungting)
so Hemibarbus maculatus is not rejected; Code Kendallia goldsboroughi Evermann & Shaw, 1927:
art. 59.3) 108 (type locality: China: Hangchow)
Barbus semibarbus Günther, 1889: 224 (replacement
name for Hemibarbus maculatus Bleeker, 1871: 19) Remarks on systematics. Most records of the
Hemibarbus joiteni Jordan & Starks, 1904: 241, pl. presence of H. leucisculus in Mongolia (Lake
64 (type locality: China: Pei Ho at Tientsin) Buir) in fact are based on misidentification of H.
Acanthogobio paltschevskii Nikolski, 1904: 356 varpachovskii, see list of citations in Sokolov (1983:
(type locality: Russia: Lake Chanka at mouth of 194). The two species are cited as present in Lake
River Santacheza) Buir by Baasanjav & Tsendayush (2001: 113),
35
Bogutskaya & Naseka (1996: 26) and Reshetnikov presence of their two subspecies only in Lake
et al. (1997: 702). Khanka [Xinkaihu; H. lucidus lucidus] and Lake
Hulun Hu [Lake Dalai Nur, adjacent to Lake Buir;
Distribution. In Mongolia: Lakes Buir and Bayan. H. lucidus varpachovskii], that are at or very close
Outside Mongolia: from Amur to Red River to their respective type localities. The fish figured
drainages; Taiwan. by Chen (1998: 169, 171) as H. lucidus lucidus and
H. lucidus varpachovskii look quite different (head
Hemiculter varpachovskii length, body depth, length of last simple dorsal
ray) and do not appear conspecific. I tentatively
Hemiculter varpachovskii Nikolski, 1904: 359 (type
retain H. varpachovskii as a valid species.
locality: Mongolia: Lake Buir)
Distribution. In Mongolia: Khalkh River and
Notes on systematics. This species was originally
Lake Buir. Outside Mongolia: Lake Hulun [Dalai
described from Lake Buir. It was later treated
Nur] in China and possibly upper Argun drainage
as a synonym of Hemiculter leucisculus (e.g.,
in Russia and China.
Berg, 1949: 365), as a subspecies of Hemiculter
bleekeri (e.g., Wu et al., 1964: 89), as a subspecies
Hypophthalmichthys molitrix
of Hemiculter leucisculus (e.g., Sokolov, 1983:
194), as a synonym of Hemiculter lucidus (e.g., Leuciscus molitrix Valenciennes, in Cuvier &
Bogutskaya & Naseka, 1996: 26), as a subspecies Valenciennes, 1844: 360 (type locality: not
of Hemiculter lucidus (e.g., Chen, 1998: 170) or stated [China ?])
as a valid species (e.g., Baasanjav & Tsendayush, Leuciscus hypophthalmus Richardson, 1845: 139,
2001: 112). pl. 63 fig. 1 (type locality: China: Canton)
Cephalus Mantschuricus Basilewsky, 1855: 235, pl.
Baasanjav & Tsendayush (2001: 113) record 7 fig. 3 (type locality: China: Beijing/Manchuria
the presence of a second species of Hemiculter /Mongolia)
in Lake Buir, as H. leucisculus. Bogutskaya & Hypophthalmichthys Basilewskii Kner, 1867: 350
Naseka (1996: 26) and Reshetnikov et al. (1997: (unnecessary replacement name for Cephalus
702) record the presence of H. leucisculus and H. mantschuricus Basilewsky, 1855: 235)
lucidus (including H. varpachovskii as a synonym) Abramocephalus microlepis Steindachner, 1869:
in Lake Buir. Sokolov (1983: 194) and Bogutskaya 150 (type locality: China; also in Steindachner,
& Naseka (1996: 26) list a number of citations of 1870: 302)
H. lucidus as H. leucisculus. They consider that H. ? Hypophthalmichthys Dabryi Bleeker, 1871: 84
leucisculus leucisculus, H. leucisculus lucidus and (not available, name listed in synonymy)
H. leucisculus varpachovskii of Nikolski (1956: ? Hypophthalmichthys Dabryi Bleeker, 1878: 210
290, 298, 301) are H. lucidus and that his H. (type locality: China: River Yang-tse-kiang)
eigenmanni in fact is H. leucisculus. To me the Hypophthalmichthys Dybowskii Herzenstein, in
figures of H. l. leucisculus and H. l. varpachowskii Warpachowski & Herzenstein, 1888: 38 (type
show two distinct species. locality: Russia: Amur River/Amur River
between Emoro and Chilusa [China: Fuchow;
The specimen I have examined from Lake Buir Berg, 1949: 846])
agrees with the description of H. leucisculus Pseudolaubuca clupeoides Duncker, 1904: 183, pl.
varpachovskii in Sokolov (1983: 194) and Chen 1 figs. 1–1a (type locality: Malaysia: Sungai
(1998: 170, 484) and their drawings. Bungus near Kuala Lumpur [introduced])
While Reshetnikov et al. (1997: 703) consider Distribution. In Mongolia: not native. Outside
H. lucidus to be distributed from the Amur to Mongolia: from Amur to Xi Jiang [Pearl River]
southern China, Chen (1998: 484) report the drainages.
36
Hypophthalmichthys molitrix was recorded a from the Ertix drainage, whose identity is not yet
few times in Lake Buir (Sokolov, 1983: 205; clear, see below]).
Tsendayush, 1968) and a few were also caught
from deep places in the upper part of Orshuun I could not find studies of L. leuciscus sensu
River and at the mouth of Khalkhiin River. lato throughout its recorded range (Europe and
This fish is commonly cultivated in China and northern Asia), posterior to that of Berg (1949).
these individuals have probably been stocked on Berg recognized the Siberian populations as a
the Chinese side of Lake Buir. It is unlikely to subspecies L. l. baicalensis distinguished from the
establish. Where introduced, H. molitrix usually European L. l. leuciscus by a terminal mouth (vs.
survives only by stocking. Adults are migratory in subterminal) and usually 9–10 branched rays in
large rivers. To successfully reproduce in the wild, the anal fin (vs. 8). The mouth position recorded
they need to have large rivers, with deep water and by Berg disagrees with his figure 313 which shows
a minimum speed; eggs are spawn at the surface a fish from the Kolyma with a subterminal mouth,
and must remain drifting in the current until they but much less than his figure 312 of a specimen
hatch and the larvae settle. If the river is blocked from the Neva. Koch & Paepke (1998: 162) also
or is too short, the eggs drop to the bottom and considered that the populations of the Selenge
fail to develop. have a subterminal mouth.
Ladislavia taczanowskii The difference in the number of branched anal-fin

rays (9–10, vs. 8) is more significant. The number
Ladislavia Taczanowskii Dybowski, 1869: 954, of anal-fin rays is usually very stable in species
pl. 17 fig. 7 (type locality: Russia: Onon and of the subfamily Leuciscinae, and differences in
Ingoda rivers) number of anal-fin rays have often been used as
a reliable character for the diagnosis of species.
Distribution. In Mongolia: in Rivers Onon, Balj Mitrofanov (2000) examined the variability of four
and Khurkh. Outside Mongolia: Amur and Yalu meristic characters in Leuciscus populations from
drainages; Korea. Kazakhstan and Europe and showed that they differ
in the number of branched anal-fin rays and gill
Leuciscus baicalensis rakers. Unfortunately he did not give ranges but
Squalidus baicalensis Dybowski, 1874: 388, pl 5 only mean values; nevertheless, as the number of
fig. 1 (type locality: Russia: Lake Baikal [rare] examined individuals are high, and the values very
and all its tributary streams) distinctive, they show clear differences. The average
Squalius suworzewi Warpachovski, 1889b: 17 number of branched anal-fin rays are between 7.5
(type locality: Kazakhstan: Irtysh River at and 8.1 for 7 European populations (11 to 226
Semipalatinsk) from Berg, 1949: 546 specimens per population) and 9.2 to 10.3 for 16
Squalius mehdem Warpachovski, 1897: 255, pl. 12 populations of the Irtysh drainage in Kazakhstan (9
fig. 1 (type locality: Russia: Ob River at Atlym) to 102 specimens per population. This corresponds
to the 8 vs. 9–10 values indicated by Berg (1949).
Remarks on systematics. The populations of dace
(Leuciscus species) of Mongolia have been reported Mitrofanov (2000) also recorded differences in
as L. leuciscus baicalensis by most authors (e.g., gill-raker counts. The 7 European populations
Sokolov, 1983: 142), as well as those from most of have mean values between 6.7 and 9.4, while the
Siberia (e.g., Reshetnikov et al., 1997: 703, 2002: 16 Irtysh populations have mean values between
277; Bogutskaya & Naseka, 2004: 85). Other 8.2 and 10.7.
authors consider them as a distinct species (e.g.,
Baasanjav & Tsendayush, 2001: 78, Luo, in Chen, These data show that the European and Siberian
1998: 70 [but this is based only on populations populations are diagnosably distinct and they are thus
37
distinct species, the European one being L. leuciscus in the lowermost part of the Irtysh. Further
and the Siberian one L. baicalensis. It remains to be downriver on the Irtysh (the name of the Ertix in
shown that all Siberian populations are conspecific. Kazakhstan), V. P. Mitrofanov et al. (1987: 80)
The identity of the Leuciscus populations inhabiting record 8–15 gill rakers in their L. “baicalensis” from
the endorheic basins of Central Asia, from Aral to the Irtysh and I. V. Mitrofanov (2000: 37) records
Balkash, requires clarification. mean values of 8.8–10.5 in 6 populations of the
upper Irtysh drainage in Kazakhstan. Kimura et
The analysis of the populations from westernmost al. (1992: 94) examined 4 specimens of Leuciscus
Europe (Loire, Garonne and Adour drainages in from the Ulungur, which they consider as possibly
France), earlier identified as L. leuciscus, shows that representing three species. They have 10–11 gill
they are not conspecific with those from the rest of rakers.
Europe (pers. obs.) and the same may possibly be
the case when populations from the rest of the range Koch & Paepke (1998) mention other characters
are examined at a closer geographic resolution. distinguishing the two species. Leuciscus
dzungaricus has a terminal mouth (vs. subterminal
Distribution. In Mongolia: Selenge drainage. in L. baicalensis), fewer vertebrae (usually 42, vs.
Outside Mongolia: rivers draining to the Arctic 44), the presence (vs. absence) of a dense cover of
Ocean, from the Ob to the Kolyma. small tubercles (or unculi ?) on the pectoral-fin
rays, and some body proportions.
Leuciscus dzungaricus
Leuciscus dzungaricus Paepke & Koch, in Koch The figures in Anonym (1979: fig. 16) and Luo
& Paepke, 1998: 162, fig. 6 (type locality: (in Chen, 1998: 70) show fishes more slender
Mongolia: Bulgan-Gol) than the holotype of L. dzungaricus, with a slightly
subterminal mouth. Luo records 41–42 vertebrae.
Remarks on systematics. This species was recently The specimens reported by Kimura et al. (1992)
described from the Bulgan River shortly upstream have 42 or 43 vertebrae. The mouth appears
of the Mongolia-China border (Koch & Paepke, terminal in their L. sp. 1, possibly subterminal in
1998). Koch & Paepke distinguished it from their L. sp. 2, and its position cannot be observed
L. baicalensis of the Selenge drainage in having, on their figure of L. sp. 3 but they report it as
among others, more gill-rakers on the first gill arch subterminal.
(14–17, vs. 7–10). Berg (1949: 546) records 7–11
gill rakers in L. baicalensis, and this agrees with the As the differences noted by Koch & Paepke allow
data of Mitrofanov (2000) discussed above, under to distinguish it from L. baicalensis, I consider
L. baicalensis. L. dzungaricus as a distinct species, but clearly,
direct examination and comparison of material
Luo (in Chen, 1998: 70) records 9–12 gill rakers in from different localities along the Bulgan-Ertix-
the L. “baicalensis” specimens they examined from Irtysh is needed to confirm their distinctness,
Lake Ulungur and the Ertix and Ulungur rivers, the limits of their ranges, and that the different
and this seems to show they are not conspecific authors used the same method for taking counts
with L. dzungaricus. Ulungur is the name of the and measurements. The presence of two species in
Bulgan in China; it is a tributary of the Ertix. One sympatry, or the existence of an introgression zone,
reasonably expects that a species from the Bulgan or clinal variation within a single species should
stretch could also occur in the Ulungur stretch. not be excluded.
If L. dzungaricus and L. baicalensis are effectively
distinct, they possibly occur in sympatry and it
Kimura et al. (1992) also record the presence of
cannot be ruled out that L. dzungaricus occurs in
L. bergi in Ulungur River. It is distinguished from
the uppermost part of Ulungur and L. baicalensis
L. baicalensis and L. dzungaricus in having more
38
gill rakers on the first gill arch (22–24, vs. 7-–15). Leuciscus farnumi Fowler, 1899: 179 (type locality:
Leuciscus bergi was earlier known only from Lake China: Tore River, tributary of Sungari River)
Issy-Kul in Kirghizistan. Its presence in the Ulungur Leuciscus waleckii sinensis Rendahl, 1925: 197
is surprising considering the very great distance (type locality: China: Shansi Prov.: Hoangho
between the two drainages, but, if confirmed, its [Huang He River], Ping-lu-hsien/Honan Prov.
presence in the Bulgan might be possible. [Henan]: Hsien-an-hsien; in title as Leuciscus
(Idus) waleckii sinensis, on p. 197 as Idus waleckii
Distribution. In Mongolia: Bulgan River. Out- sinensis)
side Mongolia: expected in Ertix drainage in China. ? Leuciscus mongolicus Oshima, 1926: 103 (type
locality: China: Jehol: Chih-fang [from Mori,
Leuciscus idus 1934: 23]; secondary junior homonym of
Squalius mongolicus Kessler, 1876: 21 when
Synonymy includes only nominal species whose placed in Leuciscus; see also Oshima, 1929
type locality is in Asia (Abstracts): 83)
Cyprinus Idus Linnaeus, 1758: 324 (based on Leuciscus brevirostris Mori, 1927: 31 (type locality:
Linnaeus [1746: 121, n. 320, Cyprinus ... radiis China: Manchuria: Hun River, tributary of Liao
13], Artedi [1738: gen. 5 [6], syn, 14, spec. 6 River)
[5], Cyprinus iride sublutea ...] and Gronovius Leuciscus waleckii tumensis Mori, 1930: 44 (type
[1754: 3, n. 13, idem]; type locality: “in Europae locality: Korea: Mo-san and Kai-nei)
aquis dulcibus”; type material: NT) Leuciscus (Idus) waleckii Joholensis Kimura, 1934:
Squalius oxianus Kessler, 1877: 124 (type locality: 13, pl. 3 fig. 1 (type locality: “Eastern Mongolia”
Uzbekistan: mouth of Amu Darya River and [China: Hebei Prov.]: Cheng-the [Chengde],
Kunja-Urgentsch in delta of Amu Darya River) River Je-Ho)
Idus oxianus Kessler, 1877: 129 (type locality: ? Leuciscus oshimae Fowler, 1958: 12 (replacement
Uzbekistan: lower part of Amu Darya River) name for Leuciscus mongolicus Oshima, 1926:
Idus melanotus var. orientalis Sinitzyn, 1900: 45 103)
(nomen nudum; Russia: Siberia: Lake Baikal) ? Leuciscus jeholi Howes, 1984: 291 (unnecessary
from Berg, 1912: 166 replacement name for Leuciscus mongolicus
Leuciscus idus idus natio sibiricus Kirillov, 1958 (infra- Oshima, 1926: 103, a junior secondary
subspecific, name not available; locality: Russia: homonym of Squalius mongolicus Kessler, 1876:
Siberia: River Leny) from Kirillov, 1962: 47 21 when placed in Leuciscus)
Distribution. In Mongolia: Selenge drainage.
Remarks on systematics. The Leuciscus population
Outside Mongolia: in Asia, from Ob to Lena
from Tumen River (border between China and
drainages; Aral basin. In Europe, Baltic, Black,
Korea) is considered as a distinct subspecies L. w.
northern Caspian and North Sea basins, Atlantic
tumensis by Luo (in Chen, 1998: 67), as a synonym
basin southward to Loire drainage (France).
of L. waleckii by Kim (1997: 251), and as probably
Reported from the Ertix in China (Anonym, 1979: a valid species L. tumensis by Bogutskaya & Naseka
26; Kimura et al., 1992: 93; Luo, in Chen, 1998: (2004: 86). Populations from upper Huang Ho
477) and thus presence in Bulgan River should not earlier referred to as L. waleckii are considered as
be excluded. a distinct species L. chuanchicus by Luo (in Chen,
1998: 68).
Leuciscus waleckii (Dybowski) Howes (1984: 289) established the genus Genghis,
Idus Waleckii Dybowski, 1869: 953, pl. 16 fig. 5 with Squalius mongolicus Kessler, 1876 as type
(type locality: Russia: Onon and Ingoda rivers, species. Howes’ Genghis mongolicus is considered
Amur drainage) to be misidentified L. chuanchicus by Luo (in
39
Chen, 1998: 68), who treated it as a species of Microphysogobio anudarini

Leuciscus; they listed L. mongolicus as a synonym
Microphysogobio tungtingensis anudarini Holcik
of L. waleckii. This would make Genghis a junior
& Pivnicka, 1969: 8, figs. 1–3 (type locality:
synonym of Leuciscus. Howes listed a number
Mongolia: Lake Buir-Nur [Lake Buir])
of characters distinguishing his Genghis from
Leuciscus and which suggest that the identity of the
Remarks on systematics. The Microphysogobio
type species of Genghis is not yet clear.
species from Lake Buir was described as M.
tungtingensis anudarini by Holcik & Pivnicka (1968:
Also there seem to be confusion as to the identity
8). Banarescu & Nalbant (1973: 263) considered it
of L. waleckii and L. mongolicus. Indirectly, Howes
one of five subspecies of M. tungtingensis. The other
(1984: 291) considered that the drawing of Idus
subspecies were M. t. amurensis, M. t. suifuensis, M.
waleckii in Dybowski (1869: pl. 16 fig. 5) shows
t. uchidai and M. t. tungtingensis.
a species distinct of that of Squalius mongolicus in
Kessler (1876: pl. 2 fig. 2). Indeed, the two differ
Microphysogobio t. suifuensis from the middle and
in head shape, shape of caudal peduncle and eye
upper Yangtze is considered a valid species by
size. The size of the eye is the character used in
Banarescu (1992: 326) and as a synonym of M.
Luo’s (in Chen, 1998: 478) key to distinguish L.
kiatingensis by Yue (in Chen, 1998: 358); this will
waleckii from L. chuanchicus (6 times or less in
not be discussed here, as all authors agree at least
head length, vs. 6 or more). Berg (1912: 92, 110,
that it is not M. tungtingensis. Microphysogobio t.
1949: 547) and Banarescu (1970: 48) considered
uchidai from Korea is considered as a subspecies
that S. mongolicus and S. chuanchicus are synonyms.
of M. tungtingensis by Banarescu (1992: 326), as a
Bogutskaya (1994: 617) examined the holotypes
valid species by Kim (1997: 244) and as a synonym
of S. mongolicus (ZISP 2472) and S. chuanchicus
of M. yaluensis by Kim & Yang (1999: 6). This later
(ZISP 2483) and found them to be distinct species.
conclusion is followed here.
Berg (1912: 92, 110) gave precedence to the name
S. mongolicus over the name S. chuanchicus.
Microphysogobio t. amurensis from the Amur drain-
age is considered as a synonym of M. tungtingensis by
The type locality of Squalius mongolicus is given as
Bogutskaya & Naseka (2004: 68) and Reshetnikov
“a southern Lake Dalai-nor (no outlet)” by Berg
et al. (2003: 283), as a subspecies by Banarescu
(1949: 547), as “Dalai Nor, Huang-Ho drainage”
(1992: 326) and Reshetnikov et al. (1997: 703), as
by Banarescu (1970: 47), and as “Hu-lun [Dalai-
a valid species by Baasanjav & Tsendayush (2001:
Nor] Lake, China” by Eschmeyer (1998: 1114).
104), and as a valid species in the genus Rostrogobio
There are a number of lakes named Dalai Nor in
by Yue (in Chen, 1998: 372). Microphysogobio t.
Mongolia and northern China. Hu Lun is one
anudarini from Lake Buir is treated as a synonym
of them, in Amur drainage, and connected with
of M. amurensis by Baasanjav & Tsendayush
Lake Buir in Mongolia. In fact, the type locality
(2001: 104) and Sokolov (1983: 181; but placed
seems to be the Dalai Nur located at 43°18’00”N
in Rostrogobio) but is overlooked in the Chinese
116°37’00”E, as is clear from the map in
literature. Microphysogobio t. tungtingensis inhabits
Przewal’skii (1876) whose expedition collected the
the lower Yangtze.
specimens. It is in an endorheic basin in Liaoning
Province of China. It is not connected with Huang
Yue (in Chen, 1998: 358, 372) places the Yangtze
He drainage. Maps suggest it might have had an
and Amur populations in two different genera.
earlier connection with Amur or Liao drainages.
In the key (p. 490) she distinguishes them by the
position of the oval pads in the lower lip (close
Distribution. In Mongolia: in Rivers Kherlen,
together in Microphysogobio, vs. slightly separate
Onon, Balj, Adraga, Khalkh, Orshuun, and
in Rostrogobio), the scalation of the belly (naked
Numrug, and in Lake Buir. Outside Mongolia:
only in front of the pectoral fins, vs. naked in
from Amur to Huang Ho drainages.
40
front of the pelvic fins), and the shape of the 61, by original designation). Gender mas-
caudal peduncle (short and deep, vs. slender). culine.
Without other information and without access
to comparison material, it is difficult to see these Remarks on systematics. The genus Oreoleuciscus
characters as distinguishing two genera and I retain is almost endemic to Mongolia. Outside Mongolia,
both in Microphysogobio. But these characters and it is known only in the Russian part of the Lake
the figures and descriptions of the two populations Uvs basin, and in the upper reaches of Chuya
in Yue (in Chen, 1998) show two species grossly (near Kosch-Agach) and Chulyshman rivers, two
distinguished by snout shape, body depth, shape tributaries of Ob River in Tuva (Russia).
of caudal peduncle, lateral line scale counts; added
to the widely disjunct ranges, they are distinct The taxonomical history of Oreoleuciscus is intricate.
species. It has been investigated by various authors with
various successes, and with very different and
Holcik & Pivnicka (1969: 10) report the following often conflicting theoretical approaches. It is not
differences between the Lake Buir (anudarini) the place here to discuss all these studies in detail
and the Amur populations (amurensis): three and I will only give a very raw summary.
simple anal-fin rays in Lake Buir, vs. two in Amur
[this last count is most likely erroneous]; distance In recent years, some Russian and Mongolian authors
between anus and anal-fin origin 17.8–19.6 % recognized a single species with a variety of forms, or
SL, vs. 23.0–29.5; and “some other features”. morphs, or ecotypes. These are mainly ecologically-
The data for the Amur population are from orientated publications, and this probably explains
Nikolski (1956). In the four specimens from the problems with the taxonomic concepts and with
Lake Buir that I examined the distance between the nomenclature. I could not find information on
the anus and the anal-fin origin is 19.0–20.8 % the species concept they used or a definition of what
SL. On the basis of this difference, I tentatively they call a ‘form’. Others have recognized some of
retain M. anudarini as a distinct species, pending these “forms” as valid species.
examination and direct comparison with material
of M. amurensis. Sokolov (1983: 146–154; 1985: 87–120)
recognized a single species, O. potanini, with
Berg (1949: 669) reported the distribution of 4 ‘forms’: dwarf, herbivorous, piscivorous and
M. amurensis as “middle and lower Amur as far sharp-snout. Baasanjav & Tsendayush (2001:
upstream as Blagoveshchensk and further” but 69–77) recognized 3 species, O. potanini (for the
it has since been collected in Shilka, Ingoda and herbivorous ‘form’), O. pewzowi (for the piscivorous
Onon systems in Russia (N. Bogutskaya and A. ‘form’) and O. humilis (for the dwarf ‘form’); they
Naseka, pers. comm.). do not mention the sharp-snout ‘form’. Bogutskaya
(2001) reviewed the literature, the systematics
Distribution. In Mongolia: Lake Buir. Outside and the nomenclature of the genus and provided
Mongolia: Probably in Lake Hu-Lun, China, and morphological and anatomical descriptions. She
possibly elsewhere in upper Amur drainage. recognized three species: O. humilis (for the dwarf
‘form’), O. potanini (for the herbivorous and the
sharp-snout ‘forms’) and O. angusticephalus (for
Oreoleuciscus
the piscivorous ‘form’). Bogutskaya did not have
Oreoleuciscus Warpachowski, 1889a: 27 (type access to usable material of the sharp-snout ‘form’
species: Chondrostoma potanini Kessler, 1879: and this is apparently the reason why she did not
306, by subsequent designation by Berg, 1912: recognize it as distinct.
81). Gender masculine.
Acanthorutilus Berg, 1912: 81 (type species: My examination of a variety of specimens and
Oreoleuciscus dsapchynensis Warpachowski, 1889a: of the literature makes me largely agree with the
41
taxonomic and nomenclatural conclusions of Dgebuadze, 1985; Golubtsov et al., 1999) also
Bogutskaya (2001). She recognized two species studied the Lake Tana barbs in Ethiopia (e.g., Mina
(O. potanini, O. angusticephalus) in the lakes and et al., 1996, 1998) where they also considered as
rivers of the Great Lakes Basin and one species (O. ‘forms’ or ‘morphotypes’ what a later taxonomic
humilis) in the Gobi Lakes Valley, Lake Uvs basin analysis demonstrated to be a species flock of 15
and isolated localities of the Selenge drainage. species (Nagelkerke, 1997; Nagelkerke & Sibbing,
The material I examined includes samples with 1997, 2000). If ‘form’ and ‘morphotype’ have
a modified dorsal fin that constitute a distinct consistent definitions in their different works,
species, for which O. dsapchynensis seems to this suggests that use of appropriate taxonomic
be the valid name (see below for reservations). procedures to the study of the Oreoleuciscus ‘forms’
Unfortunately, I have not been able to access case might lead to a greater taxonomic diversity.
material of the sharp-snout ‘form’ figured by
Sokolov (1983: 151) and Golubtsov et al. (1999:
891); its striking, unique morphology makes it
Oreoleuciscus angusticephalus
difficult to decide whether it is conspecific with Oreoleuciscus angusticephalus Bogutskaya, 2001: 35,
O. dsapchynensis or belongs to an additional fig. 7 (type locality: Mongolia: channel between
species. For the time being, I tentatively treat it as Lakes Khirgis-Nur [Lake Khyar-gas] and Airik-
conspecific with O. dsapchynensis. Nur [Lake Airag])
Bogutskaya’s conclusions are supported by Remarks on biology and systematics. This is the
morphological data. Sokolov (1983) summarized piscivorous ‘form’ of Sokolov (1983: 149, 1985:
the then available information on the biology of 99) and the O. pewzowi of Baasanjav & Tsendayush
the different ‘forms’. The presence in sympatry (in (2001: 72) and various authors. Bogutskaya (2001:
different combinations in different lakes) of O. 28) showed that the holotype of O. pewzowi in
potanini, O. angusticephalus, O. dsapchynensis and/ fact is a specimen of O. potanini, thus the name
or the sharp-snout ‘form’, with different feeding O. pewzowi is a junior synonym and cannot be
habits, and habitat preferences are additional used for the piscivorous ‘form’. There was no other
indications that they are distinct species. name available for this species and a new name (O.
angusticephalus) had to be created.
But the situation is possibly more complex in the
lakes of the Great Lakes Basin. Although I have The biology of O. angusticephalus is reported by
not examined as much material as Bogutskaya did, Sokolov (1983: 147, 1985: 88) and summarized
those specimens I examined are additional to hers by Bogutskaya (2001: 40). It is an exclusively
and partly from other localities and they suggest that lacustrine species, reaches a size up to 1000 mm
the exact distribution of the different species still SL, matures around 200–240 mm SL, and lives
requires investigation, and that the populations of up to at least 40 years. Juveniles feed on plankton,
some lakes escape the pattern presently recognized. adults prey on fishes. The species spawns in May-
For example, some samples from Lake Airag share June.
the diagnostic characters of O. humilis (see below),
a species otherwise not known in the Great Lakes Distribution. Endemic to Mongolia: lakes of
Basin. Lakes Uureg (an endorheic basin adjacent the Great Lakes Basin (Khar-Us, Khar, Nogoon,
to Khovd drainage and Lake Uvs basin) and Achit Khyargas, Achit, Tolbo, Uureg and Tal).
have populations sharing some of the characters of
O. dsapchynensis.
Oreoleuciscus dsapchynensis
Some of the authors who contributed to the study ? Leuciscus Pewzowi Herzenstein, 1883: 244
and discussion of ‘forms’ of Oreoleuciscus (e.g., (nomen nudum; locality: Mongolia: Tchon’-
Sokolov, 1983, 1985; Borisovets et al., 1985; Kharikha [Tchon-Kharnkha on Herzenstein’s
42
map; Tchonocharajch-Gol, channel between follow Sokolov et al. in treating it as mere ‘form’
Lakes Khar and Khar-Us]) of O. potanini, without a proper demonstration
? Oreoleuciscus Pewzowi Warpachowski, 1889a: (and without a clear definition of the word ‘form’).
41, pl. 1 fig. 2 (type locality: Mongolia: Tschon- This ‘form’ is found only in Lakes Khar-Us, Khar,
Charicha [Tchon-Kharikha; Tchonocharajch- Nogoon. On the other hand, it has some similarity
Gol, channel between Lakes Khar and Khar- with the O. dsapchynensis that I examined from
Us]) Lake Airag. Unfortunately I have not been able
? Oreoleuciscus Pewzowi var. altus Warpachowski, to examine material of the sharp snout ‘form’
1889a: 45, pl. 1 fig. 3 (type locality: Mongolia: from Lakes Khar-Us, Khar or Nogoon and
River Tatche-teli [Tatchen-Tel, a channel cannot conclude on their conspecificity with O.
between Lake Khar and Zavkhan River]) dsapchynensis.
? Oreoleuciscus Pewzowi var. longicaudus Warpa-
chowski, 1889a: 48, pl. 3 fig. 3 (type locality: The biology of O. dsapchynensis is described by
Mongolia: River Tatche-teli [Tatchen-Tel, a Sokolov (1983: 150, 1985: 110). It reaches a
channel between Lake Khar and Zavkhan maximum size of 280 mm and matures around
River]) 110 mm. It inhabits lakes and the lowermost part
Oreoleuciscus dsapchynensis Warpachowski, 1889a: of River Khovd. It spawns in May-June. It feeds
61, pl. 2 fig. 2 (type locality: Mongolia: on phytoplankton, plants (especially Chara) and
Dsapchyn River [Zavkhan River]) invertebrates.
Remarks on biology and systematics. I tentatively The figures of O. pewzowi, O. p. longicaudus and
identify as O. dsapchynensis material from Lake O. p. altus in Herzenstein (1889: pl 1) show fishes
Airag that I examined in Museum für Naturkunde with relatively large eye, flushed with dorsal profile
in Berlin (ZMB). Diagnostic features include the of head, relatively slender caudal peduncle, and
narrow head with long snout, a relatively large slightly falcate dorsal fin. But the branched dorsal
eye flushed with the dorsal profile of the head, rays seem ‘normally’ shaped, with membranes
frontal with a well marked shelf above orbit, the between them. Bogutskaya (2001: 32, 33) already
simple dorsal-fin rays very long, rigid almost to discussed their head shape and other details and
tip, segmented only close to tip, branched rays commented that some of them might be hybrid
1–2 rigid, not separated by membranes or only by between O. potanini and O. angusticephalus.
narrow membranes, and slender caudal peduncle Apparently, the material of the sharp-snout ‘form’
longer than head. The narrow head, the large available to her was not enough to suggest the
eye, the shelf, the length of the caudal peduncle, hypothesis that they represent a distinct species. I
the structure of the last simple ray are visible in tentatively include these 3 nominal species in the
the figure of the holotype of O. dsapchynensis in synonymy of O. dsapchynensis.
Warpachowski (1889a: pl. 2) or described by
Bogutskaya (2001: 29). The shape of the anterior Distribution. Endemic to Mongolia, presently
branched dorsal-fin rays is not very distinct in my known only from Lake Airag and Zavkhan River.
copy of Warpachowski’s plate, but the membranes The sharp-snout ‘form’ is found only in Lakes
seem narrower than in the figures of the other Khar-Us, Khar, Nogoon.
species. I tentatively conclude that the holotype
of O. dsapchynensis is conspecific with this ZMB
Oreoleuciscus humilis
material.
Oreoleuciscus humilis Warpachowski, 1889a: 50,
Oreoleuciscus sp. ‘sharp snout’ or ‘long nose’ pl. 2 fig. 3 (type locality: Mongolia: Ulaangom
figured by Sokolov (1983: 151) and Golubtsov et [in basin of Lake Uvs])
al. (1999: 891) has a uniquely shaped head among Oreoleuciscus humilis var. phoxinoides Warpachow-
Oreoleuciscus populations and it is very difficult to ski, 1889a: 54, pl. 2 fig. 4 (type locality:
43
Mongolia: Ulaangom [in basin of Lake Uvs]/ The phenomenon is very interesting and a precise
Russia: Tuva: Kosch-Agach) description of the yearly cycles at precise localities
? Oreoleuciscus pewzowi natio polybranchialis is needed to test this hypothesis and to show that
Gundriser, 1962: 250, figs. 1–2 (infrasubspecific, the pattern is exactly the same in the different lakes.
name not available; locality: Russia: Tuva: Lake An explicit description of the ontogeny of the two
Tere-Khol, south of Erzinskiy District [Uvs basin]) ‘forms’ and of the morphology of the juveniles
? Oreoleuciscus potanini infraspecies fluviatilis (separately for each lake basin) should confirm
Gundriser, 1962: 252, fig. 3 (type locality: that the juveniles of the two ‘forms’ cannot be
Russia: Tuva: Erzin River (tributary of Tess- distinguished.
Khem) at Erzin, also Naryn and Tess-Khem
rivers [Lake Uvs basin]) Although the species is not present in the Great
Lakes Basin according to Bogutskaya (2001), I
Remarks on biology and systematics. Two ‘eco- have examined specimens from Lake Airag (ZMB
morphological forms’ are recognized in O. humilis, 24046, 29051) that agree with O. humilis in
dwarf ‘form’ and lake ‘form’ (Sokolov, 1983: 148, having flexible simple dorsal rays and 7½ branched
1985: 88; summarized by Bogutskaya, 2001: dorsal-fin rays.
21). The dwarf ‘form’ occurs in shallow lakes and
their tributaries. It reaches a maximum length of Distribution. In Mongolia: basin of Lake Uvs (e.g.,
210 mm SL, matures at about 70 mm, and feeds Tes River, Lake Sangiin Dalai); lakes and rivers of
mainly on invertebrates. The lake ‘form’ inhabits Gobi Lakes Valley (Lakes Taatsyn Tsagaan, Boon
only lakes. It reaches up to 550 mm SL, matures Tsagaan and Orog, Baydrag and Ongiin rivers); a
at about 200 mm SL; by about 200 mm SL, it few isolated populations in Selenge drainage (e.g.
becomes piscivorous. Both ‘forms’ spawn in river Lake Ust). Outside Mongolia: basin of Lake Uvs;
deltas and in flood plains during floods, in June- upper Chuya River near Kosch-Agach, Tuva (Ob
August, but the dwarf ‘form’ slightly earlier than drainage), but this last locality needs confirmation
the lake ‘form’. by fresh material.
In the lakes of the Gobi Lakes Valley, which may
dry out periodically, the two ‘forms’ disappear Oreoleuciscus potanini
from the lakes and only the dwarf ‘form’ survives
Chondrostoma Potanini Kessler, 1879: 306
in the rivers from which it would re-invade the
(type locality: Mongolia: “Quellzuflussen des
lakes once they are filled again, and re-develop
Daingol” [source tributaries of Daingol Nuur =
the two ‘forms’ (Dgebuadze, 1995). The actual
Lake Dayan, 48°23’00”N 88°50’00”E]; repeated
data supporting this hypothesis have apparently
in Kessler, 1880: 267)
not been published, and a number of questions
Leuciscus latifrons Herzenstein, 1883: 244
remain unanswered, at least to the reader.
(nomen nudum; locality: Mongolia: Ulaangom
Especially, it would be very useful to have a
[erroneous, Bogutskaya, 2001: 33])
detailed list of the observation sites, the dates at
Oreoleuciscus Potanini var. recurviceps Warpachow-
which observations have been made at each site,
ski, 1889a: 38, pl. 3 fig. 2 (type locality:
and the details of the observations. The list of
Mongolia: probably Naryn River in upper
localities and dates on p. 239 suggests the above
Kungui system [Khüngiy] [Bogutskaya, 2001:
scenario is based on isolated visits at the different
23])
lakes and rivers and that they were not all visited
Oreoleuciscus similis Warpachowski, 1889a: 57,
the same year (e.g., Lake Bon Tsagaan: summer
pl. 2 fig. 1 (type locality: Mongolia: Dsapchyn
time of wet period of 1975, 1982 and 1983,
River [Zavkhan River])
summer time of dry period of 1986 and 1988;
Oreoleuciscus Herzensteini Warpachowski, 1889a:
and its tributary Baidrag River: summer time
65, pl. 1 fig. 1 (type locality: Mongolia: upper
of wet period of 1975, 1979, 1980 and 1982,
Kungui River [Khüngiy River])
summer time of dry period of 1988).
44
Oreoleuciscus gracilis Warpachowski, 1889a: 68, 26 figs. 2–3; type locality: Russia: Siberia:
pl. 1 fig. 4 (type locality: Mongolia: Ulaangom Catharinopolis)
[erroneous, Bogutskaya, 2001: 33]) ? Cyprinus Galian Gmelin, 1789: 1421 (based on
? Oreoleuciscus choerocephalus Warpachowski, Lepechin, 1771a: 491, pl. 26 figs. 2–3; 1772:
1889a: 72, pl. 3 fig. 4 (type locality: Mongolia: pl. 9 figs. 4–5; type locality: Russia: Siberia:
Lake Airik Nor [Lake Airag]; misspelled once Catharinopolis)
hoerocephalus in caption to pl. 3)
Oreoleuciscus ignatowi Nikolski, 1903: 188 (type Remarks on systematics. A number of species
locality: Russia: Tuva: Lake Tschoëbok-kul earlier placed in Phoxinus are now in the genera
[Tscheibok-kol or Choebak-kol, Baschkaus Rhynchocypris and Eupallasella, partly following
drainage], Altai Range) Howes (1985).
Oreoleuciscus warpachowskii Dulmaa, 1999: 213
(nomen nudum) The species P. phoxinus has traditionally been
considered as extending throughout Europe and
Remarks on biology and systematics. Biology northern Eurasia, reaching eastwards to Anadyr
is described by Sokolov (1983: 148, 1985: 112) and Korea (Berg, 1949; Kottelat, 1997), but recent
and summarized by Bogutskaya (2001: 33). This studies have shown that the European populations
species inhabits lakes and stretches of rivers with in fact represent several species (Kottelat,
slow currents. It feeds mostly on aquatic vegetation unpublished). A detailed study of Asian populations
and invertebrates, but larger individuals also eat is missing. My examination of material of P.
fishes. Maximum known size about 500 mm SL. “phoxinus” from the Selenge and Kherlen drainages
Mature around 180 mm SL, but much smaller in shows that they are immediately distinguished
some populations. They spawn in May-August, in from the various European species known to me by
rivers and along lake shores. a more slender caudal peduncle and also by general
appearance. These two populations also differ from
The name Oreoleuciscus warpachowskii is cited by
each other, suggesting that they are distinct species.
Dulmaa (1999: 213) but to my knowledge this
But, considering that Phoxinus are possibly the
name does not exist. It is a nomen nudum in this
most common fish in Mongolia, present in most
paper. It is probably an error from the use of the
water bodies, and with a wide, continuous range in
generic name Oreoleuciscus and the name of its
northern Asia, it is impossible to reach conclusions
nomenclatural author Warpachowski.
on their distinctness or conspecificity without
Bogutskaya (2001) includes the sharp-snouted examining and comparing material from many
‘form’ in her O. potanini. I consider that it is a more localities in the same drainages. Meanwhile
distinct species, O. dsapchynensis. they can only be treated as conspecific.
Distribution. In Mongolia: lakes and rivers of These populations are not conspecific with the
the Great Lakes Basin and Khovd and Zavkhan European species which is now called P. phoxinus but
drainages (Lakes Khar, Khar-Us, Nogoon, Dorgon, their actual name is not clear. A number of nominal
Airag, Kyar-gas, Telten, Bayan, Hoton, Horgon, species now treated as synonyms of P. phoxinus have
Horomdog, Dayan; Rivers Khovd, Zavkhan, been described from Asia and the identity of each
Khüngiy). Outside Mongolia: upper reaches of has to be investigated. The earliest name proposed
Chuya (near Kosch-Agach) and Chulyshman, two for Siberian Phoxinus outside Altai are Cyprinus
tributaries of Ob River in Tuva. isetensis (and its objective synonym C. galian),
described using material from Ekaterinenburg.
Without material I cannot comment on whether
Phoxinus cf. phoxinus it is distinct from P. phoxinus or not, or from the
? Cyprinus isetensis Georgi, 1775b: 621 (available Selenge and Kherlen populations. I therefore retain
by indication to Lepechin, 1771a: 491, pl. all as P. cf. phoxinus.
45
The Altai populations are discussed under P. 157) and Baasanjav & Tsendayush (2001: 86)
ujmonensis. I have not researched the status of record P. phoxinus from the Bulgan, a tributary of
the Balkash population for which the name P. the Ertix. I have not seen material from the Bulgan
laevis balchaschanus would be available; it is not in Mongolia, but I have examined in the Institute
unreasonable to expect that it would be distinct. of Zoology, Chinese Academy of Sciences, Beijing,
Dulmaa (1973: 64, table) lists the presence of the material from the Ulungur (the name of the Bulgan
genus Phoxinus in the Great Lakes depression. in China) at Ertai, about 80 km downstream of
Baasanjav & Tsendayush (2001: 87) explicitly state the Mongolia-China border, which I assume are
that there is no Phoxinus in that basin. conspecific with those of the Mongolian stretch.
Distribution. In Mongolia: Selenge, Onon, and I compared the Bulgan material side by side
Kherlen drainages, lakes of Darkhad depression. with material from the Selenge drainage (Alag
Outside Mongolia: exact range and number of Tsar). Although time was not sufficient to make
species not clear, see discussion above. comparison on a large number of characters on
many specimens, striking differences were observed.
Phoxinus ujmonensis They differ in having a deeper body (20.4–25.8
? Cyprinus rivularis Pallas, 1773: 717 (type % SL in the Ertix material, vs. 16.4–20.2 in the
locality: Russia: Siberia: small streams in the Selenge material), deeper caudal peduncle (9.1–
Altai range [p. 616: Zmeinogorsk (51°11’N 10.8 % SL, 2.5–2.8 times in its length, vs. 7.2–8.5
82°14’E), basin of Alei River]) and 3.2–3.6, respectively), possibly less developed
Phoxinus laevis ujmonensis Kashchenko, 1899: sexual dimorphism, and may have a different
144 (type locality: Russia: Altai: River Katun at colour pattern in life. I consider that they are
Nizhnii Uimon, Ob drainage) different species.
Phoxinus laevis mikrosquamatus Kashchenko,
1899: 145 (type locality: Russia: Altai: Lake It is presently impossible to know how many of
Karalachinskoie in Argut drainage [a tributary the five nominal species described from the Altai
of Katun River]) are valid, and which will be the valid name for
Phoxinus saposchnikowi Kashchenko, 1899: 146 the Bulgan species. In order to avoid creating
(type locality: Russia: Altai: lake on plateau of greater confusion I follow current usage and retain
Ukëk, source of River Kalguty, in Argut drainage P. ujmonensis. Phoxinus laevis mikrosquamatus
[a tributary of Katun Argut River]) and P. saposchnikowi are presently considered as
Phoxinus czekanowskii sedelnikowi Berg, 1908: simultaneous junior subjective synonyms of P.
226 (type locality: Kazakhstan: Lake Saissan ujmonensis, and precedence is given to P. ujmonensis.
[Zaisan], upper Irtysh basin, Karasuat-Busen) Cyprinus rivularis is a potential senior synonym,
but this can be clarified only when fresh material
Remarks on systematics. A number of nominal from the type locality and additional samples from
species of Phoxinus have been described from the the upper Irtysh become available.
Altai region (upper Irtysh and upper Ob drainages).
Until recently, they have been considered as It also remains to be checked whether the Ertix
synonyms of P. phoxinus. See under P. cf. phoxinus material is conspecific with the material from Altai
for discussion. (Ob drainage). The figure of the lectotype in Berg
(1949: 592, fig. 349) shows a fish with a more
Two populations of Phoxinus are known from slender body (body depth 16 % SL; depth of caudal
China. Chinese authors (e.g. Luo, in Chen, 1998: peduncle 10 % SL, 2.8 times in its length).
76, 478) use the name P. phoxinus for the Amur
population (P. cf. phoxinus above) and the name Distribution. In Mongolia: Bulgan drainage.
P. ujmonensis for the population of the Ertix (the Outside Mongolia: upper Irtysh and upper Ob
name of the upper Irtysh in China). Sokolov (1983: drainages in China, Kazakhstan and Russia.
46
Pseudaspius leptocephalus Remarks on systematics. Rhodeus amarus has long
been considered a subspecies of the East Asian R.
Cyprinus leptocephalus Pallas, 1776a: 207 (nomen
sericeus (e.g., Holcik & Duyvené de Wit, 1964;
nudum), 703 (type locality: Russia: “stony and
Arai & Akai, 1988: 211). The two species are
fast flowing streams draining to the eastern
broadly disjunct, R. amarus occurring (roughly)
Ocean” [Onon River, p. 207])
in central and eastern Europe and northern Asia
Minor, and R. sericeus in the Amur basin and
Distribution. In Mongolia: in Rivers Onon,
Sakhalin Island. They are treated here as distinct
Kherlen and Khalkhiin; Lake Buir (rare). Outside
species because they are distinct lineages separated
Mongolia: Amur drainage, Sakhalin Island.
for an estimated 2 to 4 million years (Holcik &
Jedlicka, 1994: 160) by about 4000 km and they
Pseudorasbora parva
are diagnosable (Kottelat, 1997: 75).
Leuciscus parvus Temminck & Schlegel, 1846:
215, 216, pl. 102 fig. 3 (type locality: Japan: Distribution. In Mongolia: in Lake Buir and in
Nagasaki) Rivers Kherlen, Onon, Khalkh and Orshuun.
Leuciscus pusillus Temminck & Schlegel, 1846: 216, Outside Mongolia: from Amur drainage to
pl. 102 fig. 4 (type locality: Japan: Nagasaki) southern China, Sakhalin Island.
Fundulus virescens Temminck & Schlegel, 1846:
225, pl. 102 fig. 6 (type locality: Japan: Rhynchocypris
Nagasaki)
Rhynchocypris Günther, 1889: 225 (type species:
Micraspius mianowskii Dybowski, 1869: 954
Rhynchocypris variegata Günther, 1889: 225, by
(type locality: Russia: Onon and Ingoda basins
monotypy). Gender feminine.
[numerous localities listed])
Lagowskiella Dybowski, 1916: 101, 106 (as
Pseudorasbora altipinna Nichols, 1925c: 5 (type
subgenus of Phoxinus Rafinesque, 1820b: 236;
locality: China: Sichuan: Yen-ching-kao)
type species: Phoxinus lagowskii Dybowski,
Pseudorasbora fowleri Nichols, 1925c: 5 (available
1869: 953, by original designation; subjective
by indication to Aphyocypris chinensis of Fowler,
simultaneous synonym of Czekanowskiella
1924c: 383, fig. 1; type locality: China: Anhwei:
Dybowski, 1916: 102, 109; first reviser
Ningkwo [Suancheng])
[apparently Howes, 1985: 63] gave precedence
Pseudorasbora depressirostris Nichols, 1925c: 5 (type
to Lagowskiella). Gender feminine.
locality: China: Shansi: Chin-ssu)
Czekanowskiella Dybowski, 1916: 102, 109 (as
Pseudorasbora monstrosa Nichols, 1925c: 6 (type
subgenus of Phoxinus Rafinesque, 1820b: 236;
locality: China: Fukien: near Yenping)
type species: Phoxinus czekanowskii Dybowski,
Pseudorasbora parva parvula Nichols, 1929: 8, fig.
1869: 953, by original designation; subjective
5 (type locality: China: Shantung: Tsinan)
simultaneous synonym of Lagowskiella
Pseudorasbora parva tenuis Nichols, 1929: 10, fig. 6
Dybowski, 1916: 101, 106; first reviser
(type locality: China: Shantung: Tsinan)
[apparently Howe, 1985: 63] gave precedence
to Lagowskiella). Gender feminine.
Distribution. In Mongolia: in Rivers Onon,
Moroco Jordan & Hubbs, 1925: 180 (type species:
Kherlen, Ulz, Khalkh, and Orshuun and in Lake
Pseudaspius bergi Jordan & Metz, 1913: 22 by
Buir. Outside Mongolia: from Amur drainage to
original designation). Gender masculine.
northern Vietnam; Japan; Taiwan; Hainan.
Rhynchocypris czekanowskii
Rhodeus sericeus
Phoxinus Czekanowskii Dybowski, 1869: 953
Cyprinus sericeus Pallas, 1776a: 208 (nomen
(type locality: Russia: Onon and Ingoda rivers,
nudum), 704 (type locality: Russia: Dauria
Amur drainage [Ila Bukdurga on Table; lakes
[Onon River; p. 208])
47
in valley of Ili River; Berg, 1949: 579]; spelled the first problem is to know how many species are
crebanowskii in Table, an obvious inadvertent present, and only then to decide about their
error [also corrected in Dybowski, 1872: 222]) names.
Phoxinus Strauchi Warpachowski, 1887a: 534
(type locality: Russia & Kazakhstan: tributaries Rhynchocypris czekanowskii was originally described
of Irtysh River [near Tyumen; Berg, 1949: 579]; from the Onon drainage. Material from Kherlen
also in Warpachowski, 1887b: 687) River (Amur drainage) is figured by Travers (1989:
Phoxinus sublaevis Warpachowski 1887a: 535 (type 196).
locality: Russia: tributaries of Lena River [Vilyui
River; Berg, 1949: 579]; also in Warpachowski, Distribution. In Mongolia: Rivers Tuul, Onon
1887b: 689) and Kherlen, and Lake Buir. For Mongolia, this
Phoxinus czekanowskii ignatowi Berg, 1907a: 209 is the first record of the species outside the Amur
(type locality: Kazakhstan: mouth of Seletny drainage. Outside Mongolia: Arctic Ocean basins,
River into Lake Seletytengiz [Berg, 1949: 581]) from Kara to Kolyma; Amur drainage in Russia
Phoxinus czekanowskii czerskii Berg, 1912: 225, pl. and China.
1 figs. 6, 6a (type locality: Russia: Lake Khanka
basin) Rhynchocypris lagowskii
Phoxinus czekanowskii suifunensis Berg, 1932:
Phoxinus lagowskii Dybowski, 1869: 952, pl. 15
361 (type locality: Russia: Suifun and Kangauz
fig. 4 (type locality: Russia: Onon and Ingoda
[rivers near Vladivostok])
rivers, Amur River drainage)
Remarks on systematics. Species of Rhynchocypris
Remarks on systematics. There is apparently
were earlier placed in Phoxinus. Phoxinus is quite a
some confusion as to the identity of L. lagowskii
heterogeneous assemblage and a number of authors
and some species of Rhynchocypris.
have tried to recognize and name a number of
lineages (genera or subgenera) within the genus (e.g.,
The species identified and figured as R. steindachneri
Dybowski, 1916; Gasowska, 1979; Howes, 1985).
in Travers (1989: 197) is apparently a large adult of
I partly follow Howes, but I treat Rhynchocypris and
R. lagowskii. It has the general body shape figured
Lagowskiella as synonyms. This is apparently also
by Dybowski (1969: pl. 15) for the types of L.
the conclusion reached by Bogutskaya & Naseka
lagowskii, except for the fleshy rostral process, which
(2004: 92), except that they retain Rhynchocypris
Howes use as an autapomorphy of Rhynchocypris.
and Eupallasella as subgenera of Phoxinus. A recent
But Dybowski already reported “nose protruding,
molecular study suggests that Eupallasella should
swollen during spawning season”. The photograph
be included in Rhynchocypris (Sakai et al., 2006).
in Travers (1989) also agrees with other published
As long as the interrelationships of the three genera
figures of L. lagowskii, e.g. in Luo (in Chen, 1998:
are not established, I do not see reason to consider
85; Howes, 1985: 65, fig. 5d) and with the keys
them congeneric and I see no reason to retain
in Chen (1988), Shedko (2001: 233). Further, R.
the subgenus classification which implies close
steindachneri is a species native to Japan and Korea
relationships between the different subgenera.
and never reported north of Tumen River (Kim,
1997: 280; Fujita et al., 2005).
The identity of a number of species placed in
Rhynchocypris is obscure. The small size of most
species and their non-descript colour pattern Travers (1989: 198) also recorded R. costata from
possibly explains a number of problems and Mongolia. He does not provide information on
confusions. Without abundant material of a these specimens, merely stating the characters
number of species from a number of localities distinguishing them from R. steindachneri in
throughout their ranges it is probably impossible Howes (1985). He comments that the vertebrae
to elucidate their taxonomy. In Mongolian waters, numbers distinguish the two species that Howes
48
identified under that name, but it is not clear Rutilus rutilus lacustris natio menschikowi Kirillov,
whether this character had been checked on the 1962: 40 (not available because infrasubspecific;
Mongolian specimens. locality: Russia: Yakutia: Vilyui River drainage)
Rhynchocypris costata was earlier treated as a synonym Remarks on systematics. The Siberian populations
of R. oxycephalus. Howes recorded 36 vertebrae in of Rutilus rutilus have been considered as a distinct
his R. oxycephalus, 37–38 in his R. steindachneri subspecies R. r. lacustris (e.g., by Berg, 1949: 499;
and 42–44 in his R. costata, which clearly indicates Sokolov, 1983: 140; Luo, in Chen, 1998: 89)
that he examined at least two different species. or species Rutilus lacustris (e.g., by Baasanjav &
Luo (in Chen, 1998) records 40–42 vertebrae in Tsendayush, 2001: 66). Bogutskaya & Naseka
R. lagowskii; this is the highest vertebrae number (2004: 88) consider R. lacustris as a junior
he records for all Chinese Rhynchocypris. Howes synonym of R. rutilus. Reshetnikov et al. (1997:
also recorded 40–44 vertebrae in his R. lagowskii. 705; 2002: 319) do not mention the name but the
A number of Travers’ R. costata were collected maps shows its distribution included in that of R.
together with his R. steindachneri and I suspect rutilus. The characters used by Berg to distinguish
that they too are R. lagowskii. This can only be his subspecies R. r. rutilus and R. r. lacustris are
confirmed by examination of the specimens. minor and show much overlap, and do not justify
recognizing them as distinct taxa. Holcik &
The type locality of R. costata is Duolun Skorepa (1971) and Ruban & Libosvarsky (1987)
[42°11’09”N, 116°28’39”E], Inner Mongolia, did not find differences. For the time being there
Luan He drainage [draining to the Gulf of Bohai, is thus no reason to retain R. r. lacustris as distinct.
Yellow Sea]. The species (or probably its valid
senior synonym R. oxycephalus) is not expected to Distribution. In Mongolia: Selenge drainage;
be present in Mongolia. For the time being, there is might be present in Bulgan River as it is recorded
no reason to list it as part of the Mongolian fauna. further downriver in China. Outside Mongolia:
from Europe north of Pyrenees and Alps, eastward
to Lena drainage, Aral Sea basin.
drainages, Lake Buir; Selenge drainage:
Kharuukhiin stream (Khara Bukhin, a left-hand Sarcocheilichthys soldatovi
tributary of Tuul River) (Hensel & Dashdorj, Chilogobio soldatovi Berg, 1914: 492, figs. 76–77
1978, cited by Sokolov, 1983) and Tuul (Holcik (type locality: Russia: Lower Amur, Lake Tschlja
& Pivnicka, 1969: 6). Outside Mongolia: upper /Amur, downriver of Tscheptschiki/Amur, 10
Lena and Amur drainages, in Russia and China. versts [earlier Russian unit of length equivalent
to 1.07 km] upriver of Chabarowsk/Amur, 6
Rutilus rutilus versts downstream of Chabarows /Amur, mouth
of Maginskaja Protoka/Ussuri, at confluence
Synonymy includes only nominal species whose with Buldsin/River Kamenuschka upstream of
type locality is in Asia Scheremetewa, Ussuri system)
Cyprinus Rutilus Linnaeus, 1758: 324 (based on
Linnaeus [1746: 124, n. 329, Cyprinus ... radiis Remarks on systematics. This species appears
12], Artedi [1738: gen. 3 [10], syn. 10, spec. 10 as Chilogobio czerskii in Sokolov (1983: 176)
[3], Cyprinus iride pinnis ...], Gronovius [1754: and as Sarcocheilichthys nigripinnus czerckii
2, n. 8, idem; 1746: 74, n. 51, idem, n. 52, [sic] in Baasanjav & Tsendayush (2001: 99).
Cyprinus Rex van Ruy: Waverveen, Belgium]; Sarcocheilichthys czerskii is treated as a subspecies
type locality: ”in Europae lacubus”) of S. nigripinnis by Banarescu & Nalbant (1973:
Cyprinus lacustris Pallas, 1814: 314 (type locality: 46) and as a valid species by Naseka (1996: 156),
Russia: Siberia as far as River Lena, Lake Baikal Reshetnikov et al. (1997: 705, 2002: 321), Yue
[Berg, 1949: 499])
49
(in Chen, 1998: 279) and Bogutskaya & Naseka into chankaensis, Code art. 32.5.2.1)
(2004: 71). Naseka (1996: 155), Bogutskaya & Gobio ussuriensis Berg, 1914: 473, figs. 70–71
Naseka (2004: 71) and Reshetnikov (1997: 705, (type locality: Russia: Lower Amur drainage,
2002: 325) recognize S. soldatovi as a distinct River Ussuri at mouth of River Bira)
species whose range largely overlaps that of S. Gobio ussuriensis morpha brevicirris Berg, 1914,
czerskii, except that only S. czerskii is present in 476, fig. 70 (infrasubspecific, not available;
Mongolia (Reshetnikov et al., 2002: 321). Berg locality: Russia: Ussuri drainage, River Viti)
(1914, 1949: 661) distinguished the two species Gobio ussuriensis morpha longicirris Berg, 1914,
by body shape and mouth position, subterminal 476, fig. 71 (infrasubspecific, not available;
in S. czerskii and inferior in S. soldatovi. In 1949 locality: Russia: Ussuri drainage, River Viti)
(p. 663) he recorded the presence of S. soldatovi in
Lake Buir. The single specimen I examined from Distribution. In Mongolia: Lake Buir. Outside
Lake Buir clearly has an inferior mouth, and a Mongolia: Amur drainage in Russia and China.
relatively slender body as illustrated by Berg; it is
S. soldatovi. The description and figure in Sokolov Tinca tinca
(1983: 177) are from Nikolski (1956).
Synonymy includes only names based on material
from European localities; thus only original
description listed here:
Mongolia: from Amur to Liaoning drainages (not
Cyprinus Tinca Linnaeus, 1758: 321 (based on
in central and southern Korea).
Artedi [1738: gen. 4 [27], syn. 5, spec. 27 [4],
Saurogobio dabryi Cyprinus mucosus ...] and Linnaeus [1746:
122, n. 321, Cyprinus ... ossiculorum 11]; type
Saurogobio Dabryi Bleeker, 1871: 27, pl. 5 fig. 1 locality: “in Europae stagnis, lacubus”)
(type locality: China: ? Yangtze River)
Gobiosoma amurensis Dybowski, 1872: 211 (type Distribution. In Mongolia: Bulgan River. Outside
locality: Russia: Amur River drainage) Mongolia: native in most of Europe, naturally
Pseudogobio productus Peters, 1881: 1035, pl. fig. 6 absent only in Ireland, Scandinavia north of
(type locality: Hong Kong) 61°30’N, eastern Adriatic basin and western and
Pseudogobio drakei Abbott, 1901: 486, fig. southern Greece where it is now introduced. In
(type locality: China: Hebei Prov.: Tien-Tsin Asia, native eastward to western Yenisei drainage
[Tianjin]: Pei-ho River) south of 60°N.
Longurio athymius Jordan & Starks, 1905: 197, fig.
3 (type locality: Korea: Chemulpo)
? Saurogobio longirostris Wu & Wang, 1931: 229,
fig. 4 (type locality: China: Szechwan)
Family Nemacheilidae
? Saurogobio dabryi chenghaiensis Dai & Yang, (stone loaches)
2002: 307, fig. 1 (type locality: China: Yunnan:
Lake Chenghai, 26°27’-26°38’N 100°38’- Members of the family Nemacheilidae were
100°49’E) earlier placed in the family Cobitidae and more
recently in the family Balitoridae (or its synonym
Distribution. In Mongolia: Lake Buir. Outside Homalopteridae; Kottelat, 1988). They are
Mongolia: from Amur to Xi Jiang [Pearl River] now considered as a distinct family on the basis
drainages; Korea. of molecular studies. Many species have been
placed in the genus Nemacheilus (also misspelled
Squalidus chankaensis Noemacheilus, Nemachilus, etc.) but real Nemacheilus
species live only in Southeast Asia. Several species
Squalidus chankaënsis Dybowski, 1872: 215
from Europe and northern Asia have been called
(type locality: Russia: Lake Chanka [Khanka];
Orthrias, but this name is not valid; the correct
original spelling chankaënsis must be emended
50
genus name for these fishes is Barbatula. Some Khol [50.35°N 89.83°E] and Ak-Khol Khol
authors still use Orthrias but their argument for [50.25°N 89.6°E] and River Mogen-Buren,
using the name contradicts the International Code basin of River Kobdo [Khovd in Mongolia])
of Zoological Nomenclature. ? Barbatula barbatula morpha tigris Gundriser,
1975 (infrasubspecific, name not available)
I have examined material of Barbatula from a from Prokofiev, 2003: 695
number of localities in the Amur and Selenge ? Orthrias golubtsovi Prokofiev, 2003: 698, fig. 1
drainages and from the Gobi Lakes Valley. (type locality: Russia: Tuva: Chedi-Tei River,
This material suggests that there are a number outflow of Ak-Khol Lake, Mogen-Buren
of unnamed species in Mongolia, most with drainage part of Kobdo drainage, about 60 km
restricted distribution ranges. In several cases two west of Mugur-Aksy)
species may occur in sympatry. As a revision of
the Central Asian species of the genus Barbatula Remarks on systematics. The species reported as
is supposed to be published soon by others, there Noemacheilus barbatulus by Baasanjav & Tsendayush
is no point entering into a detailed discussion of (2001: 124), as Barbatula toni by Reshetnikov
these species at this point. I include in the list below et al. (1997: 707; 2002: 356 [but figure shows a
two unnamed species present in the Mongolian Triplophysa]), and as Nemachilus barbatulus toni
material I examined. by Sokolov (1983: 207) probably include all the
species of Barbatula recognized here.
Another genus present in Mongolia is Triplophysa.
Nemachilus compressirostris was the first species of
The genus is distributed throughout Central
Barbatula recorded from western Mongolia. It was
Asia, and is very diverse, most species inhabiting
originally based on two specimens from “lakes in
a relatively small range. As for Barbatula,
northwestern Mongolia” (Warpachowski, 1887).
examination of various materials from Mongolia
Warpachowski also commented that the species
and northern China shows that there are a greater
came from the same lakes as Oreoleuciscus potanini.
number of species than reported in the literature.
These specimens were obtained for the Nizhniy
Definitive identification or formal description
Novgorod Fisheries Exhibition in 1896. The
of the unnamed species is not possible without
species is now considered to be a synonym of B.
more extensive sampling, and without material
toni (e.g., Vasil’eva, in Reshetnikov et al., 1998: 95);
of the numerous species described from adjacent
its identity is not discussed by Prokofiev (2003).
areas for comparison. I include in the list below
Sokolov (1983: 209) reported the presence of B.
an unnamed species present in the Mongolian
toni in the Khovd drainage in Mongolia, which
material I examined.
may refer to B. compressirostris, B. cobdonensis or B.
golubtsovi, but this can be established only on the
Furthermore, the exact limits and species
basis of specimens. Also, it cannot be excluded that
composition of Barbatula and Triplophysa are not
B. cobdonensis or B. golubtsovi are synonyms of B.
clear. Triplophysa obviously is a heterogeneous
compressirostris.
assemblage and some species of Triplophysa seem
to have more affinities with Barbatula than with I have examined the syntypes of B. compressirostris
other Triplophysa. (ZISP 11298). They seem to have been partly dried
at some time, which is already suggested by the figure
Barbatula compressirostris in Warpachowski (1897: pl. 11 fig. 1). The head
and snout shape figured by Warpachowski (1897:
Nemachilus compressirostris Warpachowski, 1897:
pl. 11 fig. 1) is identical to that of the holotype of
270, pls. 11 figs. 1–3 (type locality: lakes in N.
B. golubtsovi figured by Prokofiev (2003: fig. 1).
W. Mongolia)
This is certainly not enough to conclude that B.
? Nemachilus cobdonensis Gundriser, 1973: 77
golubtsovi is a synonym of B. compressirostris but
(type locality: Russia: Tuva: Lakes Khintiktig-
this hypothesis should be investigated.
51
Barbatula golubtsovi is presently known only from case of real ambiguity, the designation of a neotype
Lake Ak-Khol and its outflow Chedi-Tei River would have immediately and definitively cleared
in Mogen-Buren drainage, part of the Kobdo the problem.
[Khovd] drainage, about 60 km west of Mugur-
Aksy (Russia: Tuva). It is diagnosed (among other Barbatula cobdonensis has not yet been formally
characters) by the presence of numerous skin reported in Mongolia. I have examined
projections on the whole body. The skin of the photographs of a Barbatula species from Khovd
syntypes of B. compressirostris is smooth but the River (Erdenebat M., pers. comm.) which is
specimens have not been preserved in an optimal possibly this species, based on the appearance and
way and the skin has been abraded so that no patterning more similar to that of B. cobdonensis
conclusion can be reached. The other diagnostic on the figure in Gundriser (1979c) than that of O.
characters listed by Prokofiev (2003: 698) can be golubtsovi on the figure in Prokofiev (2003). This
checked only on the skeleton. should be confirmed by examination of specimens.
Considering the geographic isolation of the Khovd
Sokolov (1983: 209) reports 44–47 vertebrae in population and the endemicity pattern observed
material of their B. ‘toni’ from the Khongor-Ulen in the genus in adjacent areas (each basin or
River [upper Khovd drainage in Bayan-Ulgii aimag] sub-basin inhabited by one or more endemics, a
and Prokofiev (2003: 700) comments that this pattern further observed in most Nemacheilidae),
agrees with the counts in B. golubtsovi (45–47). I hypothesise that B. compressirostris and/or B.
cobdonensis are specifically distinct from topotypical
Barbatula cobdonensis was described from Lakes material of B. toni (see below).
Ak-Khol (50.25°N 89.6°E) and Khindiktig-Khol
(50.35°N 89.83°E) in Tuva Republic, Russia. In fact it is expected that two species of Barbatula
These lakes are connected to the River Mogen- are present in the Mongolian part of the Khovd
Buren and are located about 40 km upstream of drainage.
the Mongolia-Russia border. The Mogen-Buren
becomes Bökhmörön in Mongolia and enters Lake I have observed ‘tubercles’ similar to those
Achit. described by Prokofiev in B. golubtsovi in B.
sturanyi from Lake Ohrid (Europe) and a number
Prokofiev (2003: 700) comments that the original of species in Mongolia and China. Several samples
description of B. cobdonensis by Gundriser (1973: include specimens with and without ‘tubercles’.
77) is apparently based on two species, which he They are also present in some Mongolian species
identifies as B. toni and B. golubtsovi, which he had of Triplophysa.
also identified in the material he examined from
Khovd drainage in Tuva. He described B. golubtsovi, Distribution. In Mongolia: Khovd River drainage.
distinguished by the presence of ‘coarse tubercles’. Outside Mongolia: probably Cobdo [Khovd] River
He further comments that the identification of B. drainage in Tuva Republic, Russia.
cobdonensis remains uncertain and that it cannot
be excluded that a third species is present in
Barbatula dgebuadzei
Khovd drainage. He concludes in deciding that B.
cobdonensis is a ‘nomen nudum’. Judging from all Orthrias dgebuadzei Prokofiev, 2003: 700, fig. 2
the data he mentions from Gundriser’s description, (type locality: Mongolia: Zag River, basin of
the name is not a nomen nudum. At best it could Baidrag River)
be a nomen dubium, but after examining material
from the area where the nominal species was Remarks on systematics. Barbatula dgebuadzei
collected and reaching his conclusions, there is is endemic to Mongolia. It is formally recorded
absolutely no point in leaving the case as a nomen (on the basis of identified specimens) from Zag
dubium. In the absence of type material and in River (Prokofiev, 2003) and Tuy River (pers. obs.,
52
based on far from optimally preserved material). northern Asia, from the Urals to Japan (e.g., Chen,
The records of B. toni from other lakes and 1989: 29; Reshetnikov, 2002: 356; Prokofiev,
rivers of the Gobi Lakes Valley (e.g., Dgebuadze, 2003). A number of nominal species are considered
1995) are referred to B. dgebuadzei by Prokofiev. to be synonyms of B. toni. My examination of type
Considering the isolation of the basins, I think material or topotypical material of a number of
that the populations of the different lakes should them shows that a number of these synonyms in
first be compared based on fresh material before fact are valid species of Barbatula and even some
reaching this conclusion. are Triplophysa.
Distribution. Endemic to Mongolia. In lakes and Dybowski (1869) originally described B. toni from
rivers of the Gobi Lakes Valley. the Rivers Onon and Ingoda (upper Amur drainage)
and there is thus a reasonable likelihood that at least
the Onon populations in Mongolia will retain the
Barbatula toni
name. The populations referred to B. toni from the
? Nemacheilus nudus Bleeker, 1865: 12 (type Kherlen and Selenge drainages are tentatively
locality: Mongolia [p. 13, but ‘brought from retained in B. toni but this requires confirmation,
China’ on p. 14; see below]) especially since several species of Barbatula occur in
Cobitis toni Dybowski, 1869: 957 (type locality: sympatry in at least the Selenge drainage. The
Russia: “common in both river systems” [Rivers specimens which I examined and identified as B.
Onon and Ingoda, Amur drainage]) toni show great variation in body shape and
Nemachilus pechiliensis Fowler, 1899: 181 (type appearance but I have not seen enough samples to
locality: China: Pechili Prov. [Nei Mongol]: reach conclusions on the taxonomic significance of
Tan Lan Ho, tributary of Shu Lan Ho, about this variability. Prokofiev (2003: 702) figured
30 miles NE of Lama-Miau or Dolon-Nor specimens of B. toni from the Onon which can be
[Duolun, Inner Mongolia]) considered topotypical.
Orthrias oreas Jordan & Fowler, 1903: 769, fig.
2 (type locality: Japan: Hokkaido: Chitose, in In recent years, Chinese and Korean authors (e.g.,
Iburi) Zhu, 1989: 29; Wang et al., 2001: 168; Kim,
Nemacheilus sibiricus Gratzianow, 1907: 167, 168 1997: 283) have used the name B. nuda for the
(type locality: Russia: Altai: Bija [Biya] River B. toni of earlier authors. This is not without
near Bijsk [Biysk]) creating a number of problems. The type locality
Nemacheilus barbatulus tomianus Ruzskii, 1920: is usually listed as Mongolia. Indeed, Bleeker
36, figs. 1–2 (type locality: Russia: basins of (1864: 13) indicated the type locality as Mongolia,
the Rivers Tom’, Ob [Katun’, Cherga, Ursul which at his time could have meant present day’s
and Charysch rivers] and Yenisei [Abakan and Mongolia, Inner Mongolia, or some other place in
Minusinka rivers]) northern China. But on p. 14, he wrote “described
Barbatula toni fowleri Nichols, 1925b: 3 (type from a single specimen and brought from China
locality: China: Chihli [Hebei] Prov.: Eastern by the missionary David”. This holotype still
Tombs) exists (preserved in the collection of the Muséum
Nemacheilus barbatulus markakulensis Men’shikov, National d’Histoire Naturelle in Paris–MNHN
1939: 141, fig. (type locality: Kazakhstan: Lake 1450, Bertin & Estève, 1948: 98) but I have not
Marka-kul, Irtysh drainage) had the opportunity to examine it and to compare
Barbatula toni kirinensis Tchang, 1932: 115, fig. 2 it side by side with the Mongolian and Chinese
(China: Kirin [Jilin Province]) material.
Remarks on systematics. Barbatula toni has Armand David (27 September 1826–10 November
been considered as a valid name for a species or 1900; for a biography, see Boutan, 1993) travelled
a subspecies whose range extends throughout in ‘southern Mongolia’ in 1866. As Bleeker
53
described B. nuda in 1864, the specimen must flowing to the Arctic and Pacific Oceans between
have been collected earlier. In 1862, David visited the Ob and the Huang He.
the “Siwantze”, 25 km NE of Kalgan. In 1863,
he explored the mountains bordering the west of Barbatula sp. Tuul
the plain of Beijing. In 1864, he travelled in Jehol,
north-east of Beijing. Jehol is a former Chinese Remarks on systematics. An unnamed species of
province that included part of today’s Hebei, Barbatula reaching at least 140 mm SL. The colour
Shanxi and Nei Mongol provinces. This excludes pattern evolves with increasing age and size from
present Mongolia as type locality. mottled to irregular blotches with paler middle
area. Anterior and posterior nostrils conspicuously
I have examined a number of samples identified separate. Caudal fin truncate or with slightly
as B. nuda from northern China in the collections concave posterior edge. Dorsal-fin origin behind
of the Institute of Zoology, Chinese Academy of pelvic-fin origin.
Sciences, in Beijing. They turned out to represent
several species of Barbatula and Triplophysa; at Distribution. Presently known only from the
some localities, two species occur in sympatry. Selenge drainage in Mongolia (Tuul, Yeruu).
Names are available for some of them, but others Expected in Selenge drainage in Russia.
are apparently still undescribed. Until the holotype
of B. nuda can be identified with one of these Barbatula sp. Egiin
species, I see no reason to use the name to replace Remarks on systematics. An unnamed species
B. toni. Also, it would first be necessary to compare of Barbatula, with body entirely covered by soft
material from northern China with material from skin projections, with adjacent nostrils, slightly
the upper Amur to decide if one (and which one) emarginate caudal fin, dorsal-fin origin above pelvic-
of these species is conspecific with B. toni. fin origin, deep body and stout and short caudal
peduncle. The largest examined specimen is 45 mm
[Prokofiev (2003: 703) comments that the type SL but this is probably not its maximum size.
locality of B. toni is possibly China because ZISP
has a specimen received in the 18th century [sic; Distribution. Presently known only from the
that is 1701–1800 ?] from MNHN, identified as Selenge drainage in Mongolia (Egiin River).
N. nudus and with locality data “Setschun occid.” I Possibly endemic to Mongolia.
assume the material was received in 19th century if
it had a name created in 1864. I do not see reason Other Barbatula species recorded in
to speculate on the origin of a specimen on the sole Mongolian waters
ground that the name on its label is used on another
label from the same museum; by that time MNHN Dulmaa (1973: 55) and Sokolov (1983: 209)
had already received several collections from China, mention that Barbatula toni is present in the
including various nemacheilines (noteworthy are Bulgan River, but do not provide any data which
the collections of P. Dabry de Thiersant; see Dabry would allow to decide of the identity of this
de Thiersant, 1872; Sauvage & Dabry de Thiersant, population.
1874). This ZISP 4471 specimen was already
mentioned by Herzenstein (1888: 21) who gave the Barbatula toni is also reported from the Ertix and
locality as “Sse-tschuan occid.” [western Sichuan]. Ulungur drainages (Xinjiang) by Anonym (1979:
There is no record of B. nudus from Sichuan in 48, 66, fig. 41). This is apparently the species
today’s Chinese literature (e.g., Ding, 1994)]. described as B. altayensis by Zhu (1992: 241) from
Anyway, David first arrived in Sichuan in 1868]. a tributary of the Ertix at Altay.
Distribution. In Mongolia: Onon, Kherlen and A specimen identified as B. nuda [the name used
Selenge drainages. Outside Mongolia: all rivers in China for the B. toni of Russian authors, see
54
above] from the Ertix in Altay (Xinjiang, China) is species. Molecular data on Japanese and Korean
figured in Kimura et al. (1992). It does not seem populations show that additional, unnamed
to be conspecific with B. altayensis and I could not species of Lefua exist (Sakai et al., 2003). The type
identify it as a species known to me. locality of L. costata is Lake Dalai Nur in Liaoning
(China). There are several Dalai-Nur (or alternative
I have examined the type series of B. altayensis in spellings). Berg (1949: 887) commented that this
the Institute of Hydrobiology in Wuhan as well Dalai Nur is located at 43°N. This is apparently
as material which seems to be the species figured the one at 43°18’00”N 116°37’00”E [not to be
by Kimura et al. It is likely that one of these two confused with the one called Hu-Lun in Chinese
species, or both, is the B. toni recorded from the and adjacent to Lake Buir].
Bulgan by Dulmaa (1973) and Sokolov (1983)
but without specimens it is impossible to conclude Distribution. In Mongolia: streams Azargiin,
and this is the reason why I do not formally list B. Tamcagiin Bulag, Shine Usnii of Lake Buir basin
altayensis in the fauna of Mongolia. (Baasanjav & Tsendayush, 2001: 127). Outside
Mongolia: from Amur to Huang He drainages;
ZISP 12576 is a sample from “Lake Alik-Nur, Sakhalin Island.
Burkhan-Budda, Mongolia” collected in 1900
by Kozlov and Kaznakov and identified as B.
Triplophysa gundriseri
compressirostris. Burhan Buubay Uul is a mountain
located at about 45°40’N 96°43’E. I cannot find Nemacheilus dorsalis humilis Gundriser, 1962: 253,
an Alik-Nur, but there is Alag-Nuur about 150 fig. 4 (type locality: Russia: Tuva Republic: Tes-
km west of Burhan Buuday Uul. The map in Khem River, 25 km northwest of Erzin, 50°27’N
Przewal’skii (1876) shows Alak-Nor at about 44°N 95°01’E [original type locality: Russia: Tuva:
95°E and a later map (Przewal’skii, 1883) shows Erzin and Tes-Khem rivers]; junior homonym
apparently the same lake (but named Alyk-nur) of Nemachilus humilis Lin, 1932: 515)
at about 45.5°N 95°E. I am not aware of any fish Triplophysa gundriseri Prokofiev, 2002: S47
collection obtained in this area since 1900. (replacement name for Nemacheilus dorsalis
humilis Gundriser, 1962: 253)
? Triplophysa gundriseri chandagaitensis Prokofiev,
Lefua costata
2002: S55, fig. 4a (type locality: Russia: Tuva
Diplophysa costata Kessler, 1876: 29, pl. 3 fig. 4 (type Republic: Chandagaity River in Chandagaity
locality: “Mongolia: Lake Dalai-Nor at 43°N” village, 50°44’N 92°08’E)
[Berg, 1949: 887], apparently China: Liaoning,
Lake Dalai Nur, 43°18’00”N 116°37’00”E) Remarks on systematics. This species was
Nemacheilus dixoni Fowler, 1899: 181 (type reported from Tesiin River and Lake Sangiin Dalai
locality: China: Pechili Prov. [Nei Mongol]: Tan as Noemacheilus strauchii by Holcik & Pivnicka
Lan Ho, tributary of the Shu Lan Ho, about (1969: 12) and from the Tes-Khem [Tesiin] in
30 miles NE of Lama-Miau or Dolon-Nor Russia as Nemacheilus dorsalis humilis by Gundriser
[Duolun, Inner Mongolia]) (1962, 1979c) and Prokofiev (2002). Prokofiev
Elxis coreanus Jordan & Starks, 1905: 201, fig. 7 also described a subspecies T. g. chandagaitensis
(type locality: Korea: Gensan) from the Chandagaity River [Khara-Modo-Gol],
Lefua andrewsi Fowler, 1922: 1 (type locality: another tributary of Lake Uvs-Nuur; it is not clear
China: Shing Lung Shan, Eastern Tombs) whether they constitute one or two species.
Remarks on systematics. Naseka & Bogutskaya Triplophysa strauchii is a species originally

(2004) showed that the L. costata of Russian described from Lake Balkash basin [Kazakhstan].
authors (e.g., Berg, 1949: 887; Reshetnikov et Several nominal species have been considered to
al., 1997: 707, 2002: 358) contains at least two be synonyms of T. strauchii by Berg (1949: 851):
55
Nemachilus ruzskyi and N. strauchii reuniens also Triplophysa stolickai [note correct spelling] was
from Lake Balkash basin, N. ulacholicus, Diplophysa originally described from Lake Tso Morari, a small
strauchi ulacholica var. pedaschenkoi (a non- endorheic basin in Rupshu (Kashmir), surrounded
available infrasubspecific name) and N. strauchi by the Indus River drainage. A number of nominal
dorsaloides from Lake Issyk-kul basin, N. strauchi species described since have been considered to be
zaisanicus from Lake Zaisan basin. Considering synonyms of T. stolickai. They have been collected
that these basins are very isolated from each other, in localities extending from the Helmand basin
this synonymy need to be re-examined. Species of (Afghanistan), northern Pakistan, upper Amu-
the Triplophysa strauchii group have a typical body Darya River (Aral Sea basin), Tarim basin, and
shape, with a relatively deep body in front of the upper Huang He. That a single species occupied a
anal fin, and a short and shallow caudal peduncle, range extending across all drainages of High Asia,
with contrasted dark spots on the body more or with a number of endorheic basins seems highly
less aligned in vertical rows. unlikely. Some of the figures accompanying the
descriptions show very different fishes.
The figure of “Noemacheilus strauchi” in Baasanjav
& Tsendayush (2001) shows a species similar to T.
strauchii but more slender. It is not known if it is Family Cobitidae
based on an actual specimen or on a figure from (spiny loaches)
literature.
Cobitis melanoleuca
Distribution. In Mongolia: Tesiin River and Lake
Sangiin-Nuur. Outside Mongolia: basin of Lake Cobitis taenia melanoleuca Nichols, 1925a: 3 (type
Uvs-Nuur in Tuva Republic, Russia. locality: China: Shansi: Chin-ssu)
Cobitis taenia granoei Rendahl, 1935: 332, figs. 5–6
Triplophysa sp. Tuul (type locality: Russia: Irtysh River near Omsk)
Cobitis taenia sibirica Gladkov, 1935: 73 (type
Remarks on systematics. An unnamed species, locality: Russia: Lake Turgoïak, southern Urals)
apparently belonging to the genus Triplophysa, with Cobitis granoei olivai Nalbant, Holcik & Pivnicka,
the body entirely covered by soft skin projections, no 1970: 121 (type locality: Mongolia: Archangaj
scales, a complete lateral line, a deeply emarginate Co.: Lake Ögijn-nuur and Narijn River, an
caudal fin, a slender body and caudal peduncle. The upper right tributary of Orkhon River, Selenge
largest examined specimen is 67 mm SL but this is drainage)
probably not its maximum size.
Remarks on systematics. The taxonomy of the
Distribution. Mongolia: Tuul River. northern Asian species of Cobitis is not yet clear.
The synonymy of C. melanoleuca and C. granoei
Other Triplophysa species recorded in follows Nalbant (1993: 108), but a convincing
Mongolian waters demonstration is still needed. It is quite astonishing
Baasanjav & Tsendayush (2001: 124) report to have a species of Cobitis with such a huge range
the presence of “Noemacheilus stoliczkai” in extending from the Black Sea basin to the Amur
Lake Sangiin Dalai, Tesiin River, and “maybe in and Huang He drainages. It seems likely that a
Bulgan”. There is no way to know to which species pattern of a mosaic of species may be discovered, as
they refer. Their figure apparently is based on fig. happened for the European species in recent years
595 in Berg (1949: 865) of a specimen from the (see, e.g., Nalbant, 1993; Kottelat, 1997; Freyhof et
distant Chu River in (Kazakhstan). It is puzzling al., 2000).
that the recorded distribution is the same as they
report for their T. strauchii (except for the “maybe Nalbant et al. (1970) described C. granoei olivai
in Bulgan”). based on material from the Orkhon system.
56
Their figures suggest a species with a mid-lateral considered the Baikal-Selenge populations as a
series of a few small blotches (their figure 2), distinct subspecies, for which the name C. m. olivai
or somewhat elongated, poorly contrasted, and would be available. Vasil’eva refers to Vasil’ev &
somewhat connected by a dark stripe (their figure Vasil’eva (1994–a short conference abstract) for
1) differing from the large blotch pattern (see supporting karyological data. With the published
figures) illustrated by most authors for Chinese, data, this reasoning is flawed. While the karyotypes
Korean and Siberian populations (e.g., Anonym, of the Volga and Selenge populations effectively
1979: fig. 43; Chen, 1987: 39; Liu & Qin, 1987: differ and suggest they represent different taxa, this
190; Berg, 1949: 892; Kim, 1997: 310). I have does not allow recognition of a taxon peculiar to the
examined material from the Kherlen, Selenge and Selenge and Baikal. Without information on the
Bulgan drainages and could not find differences. karyotypes of populations adjacent to the Selenge
The samples from the Egiin (Selenge) and the (from the rest of the Yenisei drainage as well as from
Tuul rivers (Orkhon) each include specimens with the basins surrounding Mongolia), the identity of
the smaller and poorly contrasted pattern and the populations in intermediate areas cannot be
specimens with the large and bold pattern (see assessed. If two species are involved, it is likely that
figures). To me, these are the kind of differences the Baikal-Selenge populations will be conspecific
expected within a population, and often related with some other Siberian or East Asian ones and
with the turbidity, or substrate, or the way the one of the three senior synonyms listed above could
individual specimens have been handled before have precedence as a name for this species.
fixation. Presently there is no data to justify
retaining C. granoei as a distinct taxon. Specimens from the Ertix River (Xinjiang, China)
are figured in Anonym (1979: fig. 43) and Kimura
Specimens of C. melanoleuca with the small spot et al. (1992).
pattern have also been figured from China (e.g.,
Shaanxi: Nichols, 1943: 198; Anonym, 1992: 78, Distribution. In Mongolia: Kherlen, Onon,
fig. 3–81; Nalbant, 1993: 103; Hebei: Wang et al., Khalkhiin, Selenge and Bulgan drainages; Lakes
2001: 176) and Korea (e.g., Kim, 1997, pl. 25). Ugii, Khuvsgul and Buir. Outside Mongolia: from
the Don drainage eastwards to Amur and Huang
Sokolov (1983: 210) and Baasanjav & Tsendayush He drainages (China).
(2001: 125) recognized the presence of two
subspecies in Mongolia, which they distinguished
Iksookimia lebedevi
as C. t. taenia, with the head length greater than
the length of the caudal peduncle, and C. t. Cobitis lebedevi Vasil’eva & Vasil’ev, 1985: 464,
sibirica, with the head length about equal to the fig. 1A (type locality: Russia: Amur River near
length of the caudal peduncle. They reported Elabuga)
both from the Selenge drainage and only their ‘C.
t. taenia’ from the Kherlen and Onon drainages. Remarks on systematics. Kottelat & Lim (1992:
All the specimens I have examined show a reverse 216), Nalbant (1993: 105) and Reshetnikov et al.
situation; the Selenge drainage specimens have the (1997: 707; 2002: 360) considered I. lebedevi as a
head longer than the caudal peduncle, while it is synonym of I. choii. Iksookimia choii is endemic to
only slightly longer or as long in the specimens Kum River drainage, a very small area in Korea.
from the Kherlen drainage. Kim et al. (1999: 377) and Bogutskaya & Naseka
(2004: 104) treat I. lebedevi as a valid species.
Vasil’ev & Vasil’eva (1994: 67) reported differences
in karyotype between the populations from the Distribution. In Mongolia: Kherlen River
Volga and those from Baikal-Selenge drainages. (Vasil’eva, 1994; Reshetnikov, 1997). Outside
Later, Vasil’eva (in Reshetnikov et al., 1997: 708) Mongolia: Amur drainage in China and Russia.
57
Misgurnus mohoity Silurus asotus

Cobitis fossilis var. mohoity Dybowski, 1869: 957 Silurus Asotus Linnaeus, 1758: 304 (type locality:
(type locality: Russia: Transbaikalia: Onon and Asia)
Ingoda rivers) Silurus dauuricus Pallas, 1787: 359, pl. 11 fig. 11
? Misgurnus maculatus Bleeker, 1873: 146 (nomen (type locality: Russia: Dauuria: Onon, Ingoda
nudum; locality: China) and Argun drainages)
? Misgurnus spilurus Bleeker, 1873: 146 (nomen Silurus punctatus Cantor, 1842: 485 (type locality:
nudum; locality: China) China: Chusan Island; primary junior homonym
Nemachilus bipartitus Sauvage & Dabry de of Silurus punctatus Rafinesque, 1818a: 355)
Thiersant, 1874: 16 (type localities: North Silurus xanthosteus Richardson, 1845: 133, pl. 56
China & Central China) figs. 12–14 (type locality: China: Chusan Island
Misgurnus cestoideus Kessler, 1876: 34 (type locality: and Canton)
China: Liaoning Prov.: Dalai Nur [43°18’00”N Silurus japonicus Temminck & Schlegel, 1846:
116°37’00”E]) 226, pl. 104 fig. 1 (type locality: Japan: Higo,
Cobitis adjan Dybowski, in Sinicyn, 1900: 49 Satzuma and Nagasaki)
(nomen nudum) Silurus cinereus Dabry de Thiersant, 1872: 189, pl.
Ussuria leptocephala Nikolski, 1904: 362 (type 47 fig. 1 (type locality: China: Yang-tse-kiang
locality: River Ussuri / River Cherulu in eastern [Yangtze River])
Mongolia) Silurus bedfordi Regan, 1908: 61, pl. 2 fig. 3 (type
Misgurnus erikssoni Rendahl, 1922: 3 (type locality: locality: South Korea: Kimhoa and Chong-ju)
China: Nei Mongol: Djaggaste) Parasilurus asotus var. longus Wu, 1930b: 255, fig.
1 (type locality: China: Tchekiang: creek on the
Remarks on systematics. Systematics follows hill of Tian-Tai)
Vasil’eva et al. (2001, 2003). The Mongolian
populations were identified as M. anguillicaudatus Remarks on systematics. The genus Parasilurus
in earlier literature. is a junior synonym of Silurus (see Kobayakawa,
1989).
Distribution. In Mongolia: Rivers Onon, Kherlen,
Ulz and Khalkh; Lake Buir and Selbe River. Outside Distribution. In Mongolia: in Lake Buir and in
Mongolia: Amur drainage; northeastern China. Rivers Orshuun, Khalkh, Onon, Kherlen and
their tributaries. In 1932, introduced into Lake
Shashka, from where it entered River Selenge and
Family Siluridae rapidly invaded rivers of this drainage, until Rivers
(wels, catfishes) Orchon, Kharaa, Tuul, Eyruu, and Lake Ugii.
Outside Mongolia: from the Amur drainage to
The inclusion of S. soldatovi in the Mongolian central Vietnam; Japan; Taiwan.
fauna by Baasanjav & Tsendayush (2001: 130) is
based on a single specimen collected in 1956 in
Lake Buir and identified by Bord & Tsendayush. Family Lotidae
Otherwise, the species is known only in lowermost (burbots)
Amur, below Khabarovsk. I consider it as either a
misidentification, or resulting from an introduction Lota lota
or accidental release, or escapee on Chinese side of
Lake Buir. Synonymy includes only nominal species whose
type locality is in Asia.
58
Gadus Lota Linnaeus, 1758: 255 (based on in connection with the populations from the Ertix
Linnaeus [1746: 109, n. 292, Gadus dipterygius and Bulgan drainages.
... aequalibus], Artedi [1738: gen. [spec.] 22,
syn. 38, Gadus dorso ... ore cirrato, spec. 107, Distribution. In Mongolia: Selenge drainage:
Silurus ciro in mento unico]; type locality: “in Rivers Ider, Delgermurun, Orkhon, Tuul, Eyruu,
lacubus Europaeis”) and their tributaries, Lakes Ugii, Terhiin Thagaan,
Lota vulgaris var. obensis Anikin, 1902: 108 (type Khuvsgul and Khagiin Khar; Bulgan River; absent
locality: Russia: Siberia: River Ob) from Berg, in rivers and lakes of Darkhad depression. Outside
1949: 943 Mongolia: throughout Europe to northernmost
Lota lota asiatica Kirillov, 1972: 279 (type locality: extremity of Scandinavia; in Siberia, in rivers
Russia: Yakutia) draining to Arctic Ocean eastward to Kolyma.
Distribution. In Mongolia: rivers and lakes of

Selenge drainage, in Lake Khuvsgul and its tributaries,
Lakes Ugii, Terkhiin Tsagaan; rivers and lakes of Family Cottidae
Amur drainage, in upper reaches of Rivers Onon, (sculpins, bullheads)
Kherlen, Khalkhiin; Lake Buir. Outside Mongolia:
northern Eurasia, from France to River Lena, Black Cottus sibiricus is listed in the Mongolian fauna
and Caspian Seas basins. Populations from eastern by Baasanjav & Tsendayush (2001: 141). They
Siberia and North America earlier referred to L. lota comment that the species is possibly present in
represent a distinct species, L. maculosa. Selenge and possibly in Bulgan River. The map in
Reshetnikov et al. (2002: 170) does not show it
present in Lake Baikal basin and Selenge drainage,
Family Percidae neither is it recorded by Sideleva (2003). There is
no actual record of the species in Mongolia. To
(perch, perches)
my knowledge, there is no record of any species
of Cottus in Bulgan River, but the presence of C.
Perca fluviatilis
dzungaricus should be expected.
type locality is in Asia The taxonomy of the Siberian Cottus species is not
Perca fluviatilis Linnaeus, 1758: 289 (based on settled. The recent revision of the European species
Linnaeus [1746: 106, n. 285 (sic; 284)], Artedi by Freyhof et al. (2004) has shown that the single
[1738: gen. 39 [74], syn. 66, spec. 74 [39], Perca ‘widespread’ species C. gobio recognized by most
lineis utrinque ...] and Gronovius [1754: 42, n. authors in fact is an aggregate of at least 14 species.
96, idem]; type locality: “in Europae lacubus A similar situation is most likely to appear when
imprimis”) the Siberian populations are examined in detail.
Perca fluviatilis zaissanica Dianov, 1955 (type As presently recognized, C. sibiricus extends from
locality: Kazakhstan: Lake Zaisan) from the Ob to the Lena drainages. The type series of
Svetovidov & Dorofeyeva, 1963: 637 C. sibiricus includes material from the Yenisei.
Perca fluviatilis intermedius Svetovidov & Berg (1949: 1148) also mentions the Irtysh at
Dorofeyeva, 1963: 639 (type locality: Russia: Ust-Kamenogorsk as part of the type locality but
Siberia: River Kolyma) without supporting data.
Remarks on systematics. The population from There is no actual record of Cottus in Bulgan River
Lake Zaisan was regarded as a distinct subspecies in Mongolia, but the genus is known from the Ertix
by Dianov (1955). Its status should be investigated, River (Xinjiang, China) and its tributary Ulungur,
59
the downstream name of the Bulgan. The Ertix is Berg (1949: 1143). Holcik & Pivnicka (1969: 18)
the upper Irtysh. The subspecies C. sibiricus altaicus described topotypical material from the Onon River
was described based on material from the Ertix in Mongolia and considered C. szanaga as a valid
(Li et al., 1966: 49). It is presently not possible to species. Sokolov (1983) commented that there is no
know if the syntype(s) (?) of C. sibiricus from the difference in meristic and morphometry between
Irtysh [if they exist, see above] could be conspecific European and Amur and Onon populations of their
with C. s. altaicus. A lectotype designation for C. C. poecilopus and that Holcik & Pivnicka’s decision
sibiricus will probably be needed. to consider C. szanaga as valid was done without
comparison (which is not true, these authors
The Xinjiang C. s. altaicus is a primary junior compared the two species).
homonym of C. poecilopus altaicus, described from
the Katun drainage [Ob drainage] in Altai by Bogutskaya & Naseka (2004: 188) consider
Kashchenko (1899: 151). The junior homonym C. szanaga as a valid species. Cottus poecilopus is
C. s. altaicus has to be replaced by a new name; described from Europe and is redescribed by
it is not possible to retain it under art. 23.9.1 of Freyhof et al. (2005: 167) who did not include C.
the Code as the name C. altaicus of Kashchenko szanaga in its synonymy.
has been used as a valid name after 1899 (e.g.,
Sinicyn, 1900: 55). I establish Cottus dzungaricus I could examine only very briefly and without
as a new replacement name for Cottus altaicus optical equipment a single specimen about 50 mm
Li et al., 1966. Cottus altaicus belongs to the C. SL from the Onon drainage in the Institute of
poecilopus group and is considered to be a valid Biology, Mongolian Academy of Sciences. The first
species by Bogutskaya & Naseka (2004: 185, 188) dorsal fin is very low, much lower than on figures
and Ostroshabov & Naseka (2005). in Sokolov (1983) and Baasanjav & Tsendayush
(2001). The pelvic fin has about 4 dark bands on
For the time being, I tentatively accept that all the first ray. Prickles were not distinct but might
syntypes of C. sibiricus are conspecific. Cottus be present.
dzungaricus is distinct from C. sibiricus as described
by Berg (1949: 1148). I examined the holotype Distribution. In Mongolia: Onon drainage.
and 31 specimens of C. dzungaricus from the Ertix Outside Mongolia: Amur drainage.
and Ulungur, at Fu Hai, Fu Yun, Ke Ke Tuo Hai
and Ertai (IZCAS 50059–75, 50268–288, 55073– Leocottus kesslerii
077, 56824). Ertai is on the Ulungur, about 80
Cottus Kesslerii Dybowski, 1874: 384, pl. 2 fig. 1
km downriver of the Mongolia-China border and
(type locality: Russia: Lake Baikal and Angara,
it is very likely that the species is also present in
Irkut and Selenge Rivers)
Mongolia (it seems impossible that the suitable
Cottus trigonocephalus Gratzianov, 1902: 32 (type
habitat is missing). Cottus dzungaricus is easily
locality: Russia: Lake Baikal: Ushkan Island)
distinguished from C. sibiricus in having a naked
Cottus kessleri var. nudus Dybowski, 1908: 545
body (vs. body entirely covered by prickles).
(type locality: Russia: Lake Baikal)
Cottus szanaga Cottus kessleri bauntovi Taliev, 1946: 744, fig. 2
(type locality: Russia: Buryatia: Lake Baunt,
Cottus szanaga Dybowski, 1869: 949, pl. 14 fig. east of Lake Baikal, Vitim River drainage)
1 (type locality: Russia: Onon River and its Paracottus (Leocottus) kessleri lubricus Taliev,
tributaries, Amur River basin) 1955: 250, figs. 97–98 (type locality: Russia:
Lake Baikal)
Remarks on systematics. Recorded from Mongolia Paracottus kessleri arachlensis Tarchova, 1962:
as C. poecilopus by Sokolov (1983: 223) and 103 (type locality: Russia: Lake Arakhlei, Lake
Baasanjav & Tsendayush (2001: 141). Cottus szanaga Baikal Basin)
had been considered a synonym of C. poecilopus by
60
Paracottus kessleri gussinensis Tarchova, 1962: 108 Eleotris pleskei Warpachowski, in Warpachowski &
(type locality: Russia: Lake Gusinor, Lake Baikal Herzenstein, 1888: 19, pl. fig. 2 (type locality:
Basin) Russia: Lefu River [Ilistaya] near Nikolajewka,
Khanka Lake basin)
Distribution. In Mongolia: lower Selenge drainage Eleotris Dybowskii Herzenstein & Warpachowski,
(Dgebuadze & Dulmaa, 1996: 33). Outside in Warpachowski & Herzenstein, 1888: 21
Mongolia: Lake Baikal and small lakes in its basin; (type locality: China: swamp near Chingan
lower Selenge, Angara and Baingol Rivers. [Khingan] mountains, Amur River basin)
Mesocottus haitej Distribution. In Mongolia: Lake Buir; in Russia,

invasive in Selenge River and likely to arrive in
Cottus haïtej Dybowski, 1869: 949, pl. 14 fig. 2
Mongolia soon. Outside Mongolia: from Amur
(type locality: Russia: Onon and Ingoda Rivers
drainage to northeastern China, Korea. Introduced,
and their tributaries; spelling haïtej must be
accidentally or not, in other areas of Russia, very
emended in haitej, Code art. 32.5.2.1)
invasive and gluttonous, dangerous for the native
fauna.
Distribution. In Mongolia: Onon drainage.
Outside Mongolia: Amur drainage; northern
Sakhalin; Yaludzyan drainage. Unidentifiable Records
A ‘black bream’ was collected in 2002 in Lake
Buir (Erdenebat M., pers. comm., 2005). Without
Family Odontobutidae more information, specimen or photograph, it is
(‘sleepers’) impossible to comment on its identity. It could
possibly be Parabramis pekinensis or Megalobrama
Perccottus glenii terminalis, both of which are cultivated in China.
Perccottus Glenii Dybowski, 1877: 28 (type locality:
Russia: Golovechka [Ussuri drainage; Berg,
1948: 1056])
61
Accounts of Species Recorded from Adjacent Areas
T he following species are known from areas immediately adjacent to Mongolia and their
presence in Mongolia might be expected either as permanent or temporary inhabitants, isolated
populations, or vagrant individuals.
Accounts of Species Recorded from Adjacent Areas

Amur drainage
Family Gasterosteidae
Pungitius sinensis
Present in Dalai Nor according to Berg (1949: 968).
Family Bagridae
Pseudobagrus herzensteini
Present in Onon drainage (Berg, 1949: 918). Map in Reshetnikov (2002b: 21) shows it immediately
adjacent to Mongolia. Earlier placed in genera Macrones or Leiocassis. Generic position follows Mo
(1990: 135).
Irtysh (Ertix) drainage

The following species are unknown in the Bulgan River in Mongolia but are recorded in the Chinese part of
the Irtysh drainage (Ertix and Ulungur Rivers) (Li et al., 1966; Anonym, 1979; Kimura et al., 1992; pers.
obs. of material in the Institutes of Hydrobiology and Zoology of the Chinese Academy of Sciences, in
Wuhan and Beijing respectively). Some of them are probably restricted to the downstream part (especially
Lakes Ulungur Hu and Ji Hu). Most of them are present in the Selenge drainage in Mongolia. Mention
of A. ruthenus in the Ertix in Anonym (1979) is based on Berg’s (1948: 77) records from ‘Irtysh’ (see Li et
al., 1966: 53) and it is not retained here.
Family Acipenseridae
Acipenser baerii (see above)
Family Salmonidae
Hucho taimen (see above)
Brachymystax lenok (see above)
63
Family Coregonidae
Stenodus nelma
Family Esocidae
Esox lucius (see above)
Family Cyprinidae
Abramis brama
Often reported as A. b. orientalis. I have not researched the status of this ‘subspecies’.
Aspiopsis merzbacheri
Not previously reported from Ertix drainage, but I examined material from Burqin (47°43’00”N
86°53’00”E) on the Ertix downriver of Lake Ulungur Hu in the Institute of Hydrobiology in
Wuhan. Appears usually as Leuciscus merzbacheri in the literature. Generic placement follows Howes
(1984).
Carassius carassius (see above)
Carassius gibelio (see above)
Leuciscus idus (see above)
Rutilus rutilus (see above)
Barbatula altayensis (see above)
Barbatula sp. (see above)
Family Lotidae
Lota lota (see above)
Family Percidae
Perca fluviatilis (see above)
Gymnocephalus cernuus
Appears often as Acerina cernua in the literature.
Family Cottidae
Cottus dzungaricus (see above)
64
Recommendations
Recommendations
he present evaluation of the systematics and nomenclature of the freshwater fishes of Mongolia
T shows that, although not very diverse, this fauna is still very poorly known and its diversity
underestimated. The first, most obvious, work that should be undertaken is a survey of the
whole country.
Some priority areas should be surveyed:
• Bulgan River: there is almost no information on the local fauna; because it is the only water body in
Mongolia belonging to the Irtysh drainage it is potentially inhabited by species not found elsewhere
in the country, as suggested by the existing information on the fauna of the Ertix drainage in China
(Bulgan is a tributary of Ertix);
• Khovd drainage and lakes of the Great Lakes Basin, because of their high level of endemicity, the
potential for the discovery of several additional species, and the need to understand the status and
distribution of the Oreoleuciscus species, forms, or population;
• Onon drainage;
• springs of the Gobi Lakes Valley and along the Mongolia-China border: there is historical information
that some are inhabited by fishes and this should be verified (see under “Other Barbatula”).
Attention should especially be given to small size species (Nemacheilidae, Cobitidae, Cottidae, Phoxinus,
Rhynchocypris) as close comparison of various populations shows that a number of additional species are
likely to be found. These small-sized species often have very restricted distribution ranges and several of
the wide-ranging species presently recognized are expected to be, in fact, artificial aggregates of a number
of species with smaller ranges.
Considering the important scientific interest of Oreoleuciscus, their systematics should absolutely be
revisited, based on large scale surveys, well-preserved material and modern concepts. Morphological as
well as molecular methods should be applied. Their ecology should be revisited once their taxonomy is
understood. This could also be the topic of one or several academic research projects.
The taxonomy of Brachymystax should be resolved, but this will not be feasible without access to material
from Russia, China, Kazakhstan and Korea. The status of several populations of Thymallus and Coregonus
should be examined, especially in the case of sympatric ‘forms’.
65
Comparison material from adjacent areas of China and Russia will be needed to solve a number of the
taxonomic problems concerning the Mongolian fish fauna.
Future introductions or stocking with non-native fishes should absolutely be preceded by an Impact
Assessment, following international standards (e.g., FAO, 1996). The impact of past introductions should
be critically reviewed. The level of endemicity of the Mongolian fish fauna was earlier underestimated and
what was viewed as possibly a minor impact on widely distributed species in fact could have been a major
impact on a narrow endemic. In particular, the introduction of fish-eating and other predatory species
should be avoided.
Several species with economical importance are now shown to be in fact assemblages of several distinct
species. Fisheries policies should take this into account in management and legislation. Transbasin stocking
should be forbidden as in many cases this may introduce a species into a new drainage, with the resulting
problem of competition with the native species and the risk of hybridization.
There is an obvious need for training in techniques and methods for fish sampling, preservation,
examination and identification, as well as in practical aspects of taxonomy. It does not mean that there is a
need for full-time researchers working on fish taxonomy, but a reasonable understanding of the methods,
procedures, results, expectations, and use of the data is necessary.
66
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the goldfish, Carassius auratus (Cyprinidae)]. Musée Zoologique de l’Académie Impériale des
Voprosy Ikhtiologii, 40 (5): 581–592 [in Russian, Sciences de St-Pétersbourg], 2: 241–271, pls.
translated in Journal of Ichthyology], 40 (8): 11–12.
553–563. Warpachowski, N. 1901. [The fishes of Lake
Vasil’eva, E. D., V. P. Vasil’ev & M. O. Teletskoe]. Ezegodnik Zoologiceskago Muzeja
Skomorokhov. 2003. [Loaches (Misgurnus, Imperatorskoj Akademii Nauk, Sankt-Peterburg
Cobitidae) of Asiatic Russia: II. Morphology, [Annuaire du Musée Zoologique de l’Académie
synonymy, diagnoses, karyology, biology, Impériale des Sciences de St-Pétersbourg], 5 (4)
and distribution]. Voprosy Ikhtiologii, 43 (4): (1900 [1901]): 412–427, pl. 13.
447–456 [in Russian; translated in Journal of Warpachowski, N. & S. Herzenstein. 1887.
Ichthyology, 43 (7): 501–510. [Notizen über die Fischfauna des Amur-
82
Beckens und der angrenzenden Gebiete]. Trudy inferred from mitochondrial cytochrome b gene
St.-Petersburgskago Obscestvo Estestvoispytatelej, sequences. Hydrobiologia, 553: 255–266.
Otdeleniye Zoologii [Travaux de la Société des Yue, P.-Q. 1995. [A revision of the cyprinid fishes of
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58, 1 pl. [in Russian, Latin diagnoses] [authors Cyprinidae)]. Acta Zootaxonomica Sinica, 20
listed as Herzenstein & Warpachowski in German (1): 116–123 [in Chinese, English summary].
title]. Yue, P.-Q. (ed.) 2000. [Fauna Sinica. Osteichthyes.
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83
Appendix 1
Nomenclatural information on species mentioned in the text but not belonging to the
Mongolian fauna.
Appendix 1
Family Petromyzontidae
Lethenteron camtschaticum (Tilesius, 1811)
Petromyzon marinus Camtschaticus Tilesius, 1811: 240, pl. 9 (type locality: Japan: Jeddo and Yokohama,
by neotype designation [originally “portus Divi Petri et Pauli Camtschatici” (Russia: Kamchatka:
Petropavlovsk Kamtchatskii)])
Petromyzon Japonicus Martens, 1868: 3, pl. 1 fig. 2 (type locality: Japan: Jeddo and Yokohama)
Family Acipenseridae
Huso dauricus (Georgi, 1775)
Acipenser dauricus Georgi, 1775a: 352 (type locality: Russia: Amur, Argun, Schilka and Onon Rivers)
Family Salmonidae
Brachymystax tsinlingensis Li, 1966
Brachymystax lenok tsinlingensis Li, 1966: 92, fig. (type locality: China: Shaanxi: Chow-Tze-Hsien, How-
Chen-Tze, in Tsinling range [Qin Ling])
Family Coregonidae
Coregonus autumnalis (Pallas, 1776)
Salmo autumnalis Pallas, 1776a: 32, 1776b: 705 (type locality: Russia: ascends Pechora and Yenisei Rivers
/Lake Baikal from which it enters Angara River and Tuba River to Lake Madsharein [see Berg, 1948:
342 for more details])
85
Coregonus clupeoides La Cepède, 1803

Coregonus clupeoïdes La Cepède, 1803: 698 (type locality: Scotland: Island Inchtonachon, Lochlomoud
[Loch Lomond]; syntypes: LU)
Coregonus lavaretus (Linnaeus, 1758)

Salmo Lavaretus Linnaeus, 1758: 310 (type locality: France: Lake Bourget, by neotype designation by
Kottelat, 1997: 104)
Coregonus pollan (Thompson, 1835)

Coregonus Pollan Thompson, 1835: 78 (type locality: UK: Northern Ireland: Lough Neagh)
Coregonus sardinella (Pallas, 1814)

Salmo clupeoides Pallas, 1814: 410 (type locality: Russia: Siberia: Kolyma River from mouth to
Srednekolymsk and further upstream, Alazeya, Indigirka at Zashiversk, and Arctic Ocean [Berg, 1948:
328; Valenciennes, in Cuvier & Valenciennes, 1848: 517 also mentions Irtysh, which is not mentioned
by Pallas]; junior secondary homonym of Coregonus clupeoides La Cepède, 1803: 698 when placed in
Coregonus)
Prosopium cylindraceum (Pennant, 1784)

Salmo cylindraceus Pennant, 1784: ciii, xxxvii (type locality: Russia: Lena, Indigirfka [Indighirka], and
Kowyma [Kolyma] Rivers)
Family Thymallidae
Thymallus brevipinnis Svetovidov, 1931

Thymallus arcticus var. brevipinnis Svetovidov, 1931: 85 (type locality: Russia: Lake Baikal)
Thymallus burejensis Antonov, 2004

Thymallus burejensis Antonov, 2004: 443, fig. 2 (type locality: Russia: Bureye River [a tributary of Middle
Amur]; holotype: MGU P-20928)
Thymallus mertensii Valenciennes, 1848

Thymalus [sic] Mertensii Valenciennes, in Cuvier & Valenciennes, 1848: 453 (type locality: Russia:
Kamchatka)
Thymallus pallasii Valenciennes, 1848

Thymalus [sic] Pallasii Valenciennes, in Cuvier & Valenciennes, 1848: 448 (type locality: Russia)
Thymallus signifer (Richardson, 1823)

Coregonus signifer Richardson, 1823: 711, pl. 26 (type locality: Canada: rivers north of Great Slave
Lake)
86
Family Cyprinidae
Abramis brama (Linnaeus, 1758)
Abramis brama bergi Grib & Vernidub, 1935: 112 (type locality: Aral Sea at Muinak / Lake Yaskhan in
Uzboi/River Sary-su; junior primary homonym of Abramis sapa bergi Belyaev, 1929) from Berg, 1949:
774
Abramis brama orientalis Berg, 1949: 774 (replacement name for Abramis brama bergi Grib & Vernidub,
1935)
Aspiopsis merzbacheri Zugmayer, 1912

Aspiopsis merzbacheri Zugmayer, 1912: 682 (type locality: China: Manas River near Manas city, northwest
of Urumtschi [Urumqi] on northern slopes of Thian-Shan range; also in Zugmayer, 1913: 13, pl.)
Barbus barbus (Linnaeus, 1758)

Cyprinus Barbus Linnaeus, 1758: 320 (based on Artedi [1738: gen. [spec.] 4, syn. 8, Cyprinus maxilla
superiore longiore ...], Gronovius [1754: 5, n. 20. idem; 1756: 3, n. 20, idem] and on data from the
then unpublished Linnaeus, 1764: 107 [from Spain]; type locality: River Ijssel at Deventer, Netherlands
Appendix 1
[by lectotype designation by Kottelat, 1997: 48; original locality: “in Europa australis”)
Genghis Howes, 1984

Genghis Howes, 1984: 289 (type species: Squalius mongolicus Kessler, 1876: 21, by original designation).
Gender masculine.
Gobio macrocephalus Mori, 1930

Gobio gobio macrocephalus Mori, 1930: 46 (type locality: Korea: Kai-nei/Kei-Ko)
Gobio tungussicus Borisov, 1928

Gobio gobio tungussicus Borisov, 1928: 105, 165, pl. 6 figs. 14–15 (type locality: Russia: Sakha-Yakutia:
Lena River near Zhigansk)
Leuciscus chuanchicus (Kessler, 1876)

Squalius chuanchicus Kessler, 1876: 23 (type locality: China: Huang He River)
Squalius mongolicus Kessler, 1876: 21, pl. 2 fig. 2 (type locality: China: Liaoning Prov.: Lake Dalai-Nor
[endorheic lake at 43°18’00”N 116°37’00”])
Luxilus cornutus (Mitchill, 1817)

Cyprinus haematopterus Rafinesque, 1820a: 6 (type locality: USA: New York: streams falling into the
Hudson River)
Microphysogobio amurensis (Taranetz, 1937)

Rostrogobio amurensis Taranetz, 1937: 114 (type locality: Russia: middle and lower Amur River and Khanka
Lake)
87
Microphysogobio kiatingensis (Wu, 1930)

Pseudogobio kiatingensis Wu, 1930a: 70, fig. 1 (type locality: China: Sichuan: Kiating [Lo-Shan], upper
Yangtze drainage)
Pseudogobio suifuensis Wu, 1930a: 71, fig. 2 (type locality: China: Sichuan: Suifu)
Microphysogobio tungtingensis (Nichols, 1926)

Pseudogobio tungtingensis Nichols, 1926: 4, fig. 4 (type locality: China: Hunan Prov.: Huping, Tungting
Lake)
Microphysogobio yaluensis (Mori, 1928)

Pseudogobio yaluensis Mori, 1928: 59 (type locality: Korea: Yalu River at Tsao-ho-kou)
Microphysogobio tungtingensis uchidai Banarescu & Nalbant, 1973: 264, fig. 139 (type locality: South
Korea: Sinch’on-ni, 35°16.5”N 128°50.7’E, about 25 km west-northwest of Pusan)
Phoxinus phoxinus (Linnaeus, 1758)

Cyprinus Phoxinus Linnaeus, 1758: 322 (based on Artedi [1738: syn. 12, Cyprinus tridactylus ...]; type
locality: “in Europa”)
? Phoxinus laevis var. balchaschana Kessler, 1879: 283 (type locality: Kazakhstan: River Ajagus [Ayaguz]
near Sergiopol [Ayaguz], Lake Balkash basin; also in Kessler, 1880: 234)
Rhynchocypris oxycephalus (Sauvage & Dabry, 1874)

Pseudophoxinus oxycephalus Sauvage & Dabry de Thiersant, 1874: 11 (type locality: China: Pékin [Beijing],
Si-wan, and southern Shen-si)
Leuciscus costatus Fowler, 1899: 180 (type locality: China: Pechili [Nei Mongol]: Tan lan Ho River,
tributary of Shu lan Ho, approximately 30 miles northeast of Lama-miau or Dolon-nor [Duolun,
Inner Mongolia])
Rhynchocypris steindachneri (Sauvage, 1883)

Phoxinus Steindachneri Sauvage, 1883: 148 (type locality: Japan: Lake Biwa)
Sarcocheilichthys nigripinnis (Günther, 1873)

Gobio nigripinnis Günther, 1873: 246 (type locality: China: Shanghai)
Sarcocheilichthys czerskii (Berg, 1914)

Chilogobio czerskii Berg, 1914: 490, fig. 75 (type locality: Russia: Sintukha, Lake Khanka basin)
(stone loaches)
Barbatula altayensis Zhu, 1992

Barbatula altayensis Zhu, 1992: 241, figs. 1-2 (type locality: China: Xinjiang: Kelang He River, a tributary
of the Ertix River, near Altay City, 47°52’N 88°06’E)
88
Barbatula sturanyi (Steindachner, 1892)
Nemachilus Sturanyi Steindachner, 1892a: 131 (type locality: FYROM: Lake Ohrid at Pestani, between
Ohrid City and Naum monastery; also in Steindachner, 1892b: 378, pl. 2 fig. 3)
Triplophysa stolickai (Steindachner, 1866)

Cobitis stolickai Steindachner, 1866b: 793, pl. 14 fig. 2 (type locality: India: Kashmir: Rupshu: rivulets in
the vicinity of Lake Tso Morari)
? Nemachilus dorsonotatus plagiognathus Herzenstein, 1888: 33, pl. 5 fig. 5, pl. 7 fig. 2 (type localities:
China: Lake Kuku-nor, eastern Mongolia [Qinghai Hu, China]/Eastern Zaidam [Tsaidam]/spring at
Galmyk/Gan-ssu/Dabsun-gobi [apparently Caka Yanhu 36°42’00”N, 99°06’00”E, Dalai Dabassu on
map in Przewal’skii, 1876]/Chami [Hami, Xinjiang; 42°48’00”N, 93°27’00”E ]/Lake Alak-nor [Alag
Hu]/Chuan-che R. near Gomi [upper Huang He about 50 km upriver of Guide]; syntypes: ZISP 7252
[1], ZISP 7306 [2], ZISP 7312 [5], ZISP 7315 [2], ZISP 7319 [more than 6], ZISP 7371 [1], ZISP
7853 [2], ZISP 7854 [1], ZISP 7855 [1])
Triplophysa strauchii (Kessler. 1874)

Diplophysa Strauchii Kessler, 1874: 58, pl. 8 fig. 40 (type locality: Kazakhstan: River Ili, tributary of Lake
Appendix 1
Balkash, and Ich-Balya River)
Nemachilus ulacholicus Anikin, 1905: 3, 18 [of reprint] (type locality: Kirghizistan: Lake Issyk-kul at the
mouth of Ulakhol River)
Diplophysa strauchi ulacholica var. pedaschenkoi Berg, 1931b: 312, fig. 2 (an infrasubspecific name, not
available)
Nemachilus strauchi zaisanicus Menschikov, 1937: 437 (type locality: Kazakhstan: Karasu River at Akdzhar,
Tarbagatai District, basin of Lake Zaisan, 40 km from the lake) from Berg, 1949
Nemacheilus strauchi dorsaloides Turdakov, 1947: 155 (type locality: Kirghizistan: Tyupsky Bay of Lake
Issyk-kul, USSR)
Nemachilus ruzskyi Nekrashevich, 1948: 121 (type locality: Kazakhstan: Lake Alakul, east of Lake
Balkash)
Nemachilus strauchi reuniens Turdakov, 1952: 57 (type locality: Kirghizistan: Irisu River, tributary of
Karkara River [tributary of Charyn River, Lake Balkash basin])
Family Cobitidae
(spiny loaches)
Iksookimia choii (Kim & Son, 1984)

Cobitis choii Kim & Son, 1984: 50, fig. 1 (type locality: South Korea: Chungcheongbug-do Prov.: Miheo-
cheon stream, a tributary of Geum River at Yeocheon-ri, Ochang-myon, Cheongwon-gun)
Misgurnus anguillicaudatus (Cantor, 1842)

Cobitis anguillicaudata Cantor, 1842: 485 (type locality: China: Chusan Island)
89
Family Siluridae
(wels, sheatfishes)
Silurus soldatovi Nikolski & Soin, 1948
Silurus soldatovi Nikolski & Soin, 1948: 1359, fig. 1 (type locality: Russia: Khabarovskyi Krai: Amur
River, Lake Kabar at Elabuga)
Family Bagridae
(bagrid catfishes)
Pseudomystus herzenszeini (Berg, 1907)
Macrones herzensteini Berg, 1907b: 421 (type locality: Russia: mouth of Onon River)
Family Gasterosteidae
(sticklebacks)
Pungitius sinensis (Guichenot, 1869)
Gasterosteus sinensis Guichenot, 1869: 204, pl. 12 fig. 4 (type locality: China [Yangtze River])
Family Percidae
(perches)
Gymnocephalus cernuus (Linnaeus, 1758)
Perca Cernua Linnaeus, 1758: 294 (based on Linnaeus [1746: 107, n. 286, Perca ... radiis 27], Artedi
[1738: gen. 40 [80], syn. 68, spec. 77 [40], Perca dorso monopterygio capite ...], Gronovius [1754: 41,
n. 94, idem]; type locality: “in Europae lacubus”)
Family Cottidae
(sculpins)
Cottus altaicus Kashchenko, 1899
Cottus poecilopus altaicus Kashchenko, 1899: 151 (type locality: Russia: Altai: Sema River at Cherga/
Ryblushka, a settlement, close to Cherga, on Rybnuskka stream, Katun system/Katun River at Nizhnii
Uimon)
Cottus dzungaricus Kottelat, 2006

Cottus sibiricus altaicus Li & Ho in Li, Tai, Chang, Ma & Ho, 1966: 49, fig. 2 (type locality: China: Altai,
northern Sinkiang; junior primary homonym of Cottus poecilopus altaicus Kashchenko, 1899: 151)
Cottus dzungaricus Kottelat, 2006 (see above) (replacement name for Cottus sibiricus altaicus Li & Ho in
Li, Tai, Chang, Ma & Ho, 1966: 49)
90
Cottus poecilopus Heckel, 1837
Cottus poecilopus Heckel, 1837: 145, pl. 8 figs. 1-2 (type locality: Slovakia: a hill stream [probably Cerveny;
Kottelat, 1997: 169] of the Carpathes [Vysoké Tatry], near Grossschlagendorf [Vel’ky Slavkov] near
Käsmark [Kezmarok], Upper Hungary [now Slovakia], Vistula basin)
Cottus sibiricus Warpachowski, 1889

Cottus sibiricus Warpachowski, 1889a: 12 (type locality: Russia: River Yenisei at Minusinsk and Abokak
[Abakan] [Berg, 1949: 1148 also mentions River Irtysh off Ust-Kamenogorsk, but without evidence])
Appendix 1
91
Appendix 2: Figures
Appendix 21
Appendix
Lethenteron reissneri Image: M. Kottelat
Acipenser baerii Source: Berg, 1911 Acipenser schrenckii Source: Berg, 1911
Hucho taimen juvenile Image: M. Kottelat
Hucho taimen Image: Z. Hogan
Brachymystax sp. juvenile Image: M. Kottelat
Brachymystax lenok Image: J. Schöffmann
93
Brachymystax cf. tumensis Image: J. Schöffmann Coregonus chadary Source: Berg, 1932
Coregonus migratorius Source: Berg, 1932 Coregonus pidschian Source: Berg, 1932
Thymallus cf. arcticus Image: M. Kottelat Thymallus baicalensis Image: J. Schöffmann
Thymallus brevirostris Image: J. Schöffmann Thymallus grubii Image: J. Schöffmann
Thymallus nigrescens Image: M. Kottelat Thymallus sp. 1 Image: Erdenebat M.
94
Esox lucius Image: M. Kottelat
Esox reichertii Image: Erdenebat M.
Appendix 21
Appendix
Acheilognathus asmussii Image: Erdenebat M. Carassius carassius (?) Image: M. Kottelat
Chanodichthys erythropterus Image: M. Kottelat
Carassius gibelio Image: Erdenebat M.
Chanodichthys mongolicus Source: Berg, 1932 Culter alburnus Source: Berg, 1932
95
Eupallasella percnurus Image: M. Kottelat
Cyprinus rubrofuscus (?) Image: M. Kottelat
Gobio acutipinnatus Image: M. Kottelat
Gnathopogon strigatus Source: Berg, 1914
Gobio cynocephalus Image: M. Kottelat Gobio soldatovi Source: Berg, 1911
Gobio tenuicorpus Source: Mori, 1934 Gobio sp. Onon Source: Nikolski, 1956
Hemibarbus labeo Source: Berg, 1914 Hemibarbus maculatus Image: M. Kottelat
96
Hemiculter leucisculus Image: M. Kottelat Hemiculter varpachovskii Image: M. Kottelat
Ladislavia taczanowskii Image: M. Kottelat
Appendix 21
Leuciscus baicalensis Image: Erdenebat M.
Appendix
Leuciscus dzungaricus Image: M. Kottelat Leuciscus idus Image: M. Kottelat
Leuciscus waleckii Image: M. Kottelat

Microphysogobio anudarini Image: M. Kottelat
Oreoleuciscus angusticephalus Image: M. Kottelat Oreoleuciscus dsapchynensis Image: M. Kottelat
97
Oreoleuciscus humilis Image: M. Kottelat

Oreoleuciscus potanini Image: M. Kottelat
Phoxinus cf. phoxinus Image: M. Kottelat Phoxinus ujmonensis Image: M. Kottelat
Pseudaspius leptocephalus Image: Erdenebat M. Pseudorasbora parva female Image: M. Kottelat
Pseudorasbora parva male Image: M. Kottelat Rhodeus sericeus Source: Berg, 1932
Rhynchocypris czekanowskii Image: M. Kottelat Rhynchocypris lagowskii Image: M. Kottelat
98
Rutilus rutilus Image: M. Kottelat
Sarcocheilichthys soldatovi Image: M. Kottelat
Saurogobio dabryi Image: M. Kottelat
Appendix 21
Squalidus chankaensis Image: M. Kottelat
Appendix
Barbatula compressirostris Image: M. Kottelat
Tinca tinca Image: M. Kottelat
Barbatula dgebuadzei Image: M. Kottelat Barbatula toni Image: M. Kottelat
Barbatula sp. Tuul Image: M. Kottelat Barbatula sp. Egiin Image: M. Kottelat
99
Barbatula altayensis Image: M. Kottelat

Lefua costata Image: M. Kottelat
Triplophysa gundriseri Image: M. Kottelat Triplophysa sp. Tuul Image: M. Kottelat
Cobitis melanoleuca Image: M. Kottelat Cobitis melanoleuca Image: M. Kottelat
Misgurnus mohoity Image: M. Kottelat
Silurus asotus Image: M. Kottelat
Lota lota Image: M. Kottelat
Perca fluviatilis Image: M. Kottelat
100
Cottus szanaga Image: M. Kottelat Mesocottus haitej Source: Berg, 1909
Leocottus kesslerii Source: Berg, 1932
Appendix 21
Perccottus glenii Source: Berg, 1932
Appendix
Coregonus peled Source: Berg, 1932 Coregonus sardinella Source: Berg, 1932
Ctenopharyngodon idella Image: M. Kottelat Hypophthalmichthys molitrix Image: M. Kottelat
Silurus soldatovi Source: Nikolski, 1954
101
Addendum
Addendum
hile this report was going to press, funding became available for a brief period of fieldwork,
W unfortunately too late to have the results included here. The present addendum is written in
Khovd in the middle of my field trip and mentions a few raw observations on the material
obtained. It was decided to include the photographs, although the identifications of several
samples are very tentative, made in the field, without access to any literature. Further, a number of species
apparently new to science have been observed.
Oreoleuciscus dsapchynensis
Material obtained in Airag Lake agrees with my hypothesis that it is a valid species.
Oreoleuciscus humilis
Material collected at several localities, including at the type locality (Ulaangom), suggests that several
discrete species are confused under this name. Material from Baydrag River seems to include two species
in sympatry.
Oreoleuciscus angusticephalus
Material from Lakes Airag and Khyargas does not seem to be conspecific.
Barbatula (?) compressirostris

A species quite similar to the figure in the original description was observed at many localities in the
Khovd River drainage. It agrees with the original description of B. golubtsovi, treated as a synonym of B.
compressirostris (see above).
Carassius carassius
Presence in Bulgan River is confirmed.
Barbatula altayensis
Presence in Bulgan River is confirmed. At least one other species of Barbatula, still unidentified, is also
observed in Bulgan River.
Barbatula dgebuadzei
Observed only in Baydrag River.
Thymallus, Phoxinus and Cottus were not observed in Bulgan River.
103
Environment and Social Development
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Fishes of Mongolia
A check-list of the fishes
known to occur in Mongolia with
comments on systematics and nomenclature
MAURICE KOTTELAT

Fishes of Mongolia

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Carassius carassius (Linnaeus, 1758)

INVALID NAMES VALID NAME

Rostrogobio amurensis misidentiﬁed Microphysogobio anudarini

Family Petromyzontidae habitats. It is not known from the Yenisei drainage

Coregonus migratorius Distribution. In Mongolia: native, migratory in

Thymallus nigrescens The nominal species T. arcticus dentatus described

Cyprinus rubrofuscus used to produce hybrids with other species of the

Gobio acutipinnatus Distribution. In Mongolia: Bulgan River. Outside

Gobio tenuicorpus Gobio sp. Onon

Ladislavia taczanowskii The diﬀerence in the number of branched anal-ﬁn

Chen, 1998: 68), who treated it as a species of Microphysogobio anudarini

Remarks on systematics. Naseka & Bogutskaya Triplophysa strauchii is a species originally

Misgurnus mohoity Silurus asotus

Distribution. In Mongolia: rivers and lakes of

Mesocottus haitej Distribution. In Mongolia: Lake Buir; in Russia,

Accounts of Species Recorded from Adjacent Areas

Irtysh (Ertix) drainage

Some priority areas should be surveyed:

Okada, Y. 1959–60. Studies on the freshwater Pallas, P. S. 1814. Zoographia rosso-asiatica,

Coregonus clupeoides La Cepède, 1803

Coregonus lavaretus (Linnaeus, 1758)

Coregonus pollan (Thompson, 1835)

Coregonus sardinella (Pallas, 1814)

Prosopium cylindraceum (Pennant, 1784)

Thymallus brevipinnis Svetovidov, 1931

Thymallus burejensis Antonov, 2004

Thymallus mertensii Valenciennes, 1848

Thymallus pallasii Valenciennes, 1848

Thymallus signifer (Richardson, 1823)

Aspiopsis merzbacheri Zugmayer, 1912

Barbus barbus (Linnaeus, 1758)

Genghis Howes, 1984

Gobio macrocephalus Mori, 1930

Gobio tungussicus Borisov, 1928

Leuciscus chuanchicus (Kessler, 1876)

Luxilus cornutus (Mitchill, 1817)

Microphysogobio amurensis (Taranetz, 1937)

Microphysogobio kiatingensis (Wu, 1930)

Microphysogobio tungtingensis (Nichols, 1926)

Microphysogobio yaluensis (Mori, 1928)

Phoxinus phoxinus (Linnaeus, 1758)

Rhynchocypris oxycephalus (Sauvage & Dabry, 1874)

Rhynchocypris steindachneri (Sauvage, 1883)

Sarcocheilichthys nigripinnis (Günther, 1873)

Sarcocheilichthys czerskii (Berg, 1914)

Barbatula altayensis Zhu, 1992

Triplophysa stolickai (Steindachner, 1866)

Triplophysa strauchii (Kessler. 1874)

Iksookimia choii (Kim & Son, 1984)

Misgurnus anguillicaudatus (Cantor, 1842)

Cottus dzungaricus Kottelat, 2006

Cottus sibiricus Warpachowski, 1889

Hucho taimen juvenile Image: M. Kottelat

Hucho taimen Image: Z. Hogan

Brachymystax sp. juvenile Image: M. Kottelat