How many species are there within the genus Acipenser?

Abstract

In their paper in this volume Bemis et al. (1997) ask: ‘How many valid species of Acipenser should we recognize-’ Although a partial answer to this question is presented in their Table 5, we discovered in the course
of preparing this volume that sonic additional commentary is needed. In fact, there are two questions: (1) how many species
should be recognized? and (2) what scientific names should be used for some of the species? The sympatric distributions of
most species of sturgeons set the stage for much confusion about species boundaries, but the situation is actually much more
complicated. Confusion about the number of species of sturgeons living within the same basin can result from the often close
morphological and meristic similarities of certain species of Acipenser, particularly during juvenile period. Moreover, westill have remarkably inadequate knowledge of the comparative anatomy of
thespeciesof Acipener: no modern study has ever attempted a comprehensive examination of all species, and it is impossible to rely on literature
for the sorts of comparisons that must be made (for more on this general problem, see Grande & Bemis 1991,1997). Most classical
descriptions and comparativean atomical studies relieduponsmall sample sizes. Voucher specimens of large sturgeons areespecially
rare in mosthistorical collections, and type specimens (if available at all) are seldom prepared in ways that are suitable
for making detailed anatomical comparisons (e.g., many skins are simply overstuffed with straw, so that all internal structures
are lost). Intraspecific morphological and meristic polymorphisms occur in all species of acipenserids, and in most cases
we have very poor knowledge of differences that develop during ontogeny, particularly changes in such features as the shape
of the rostrum (Bemis et al. 1997). Another problem is the ease of hybridization between different species of sturgeons (reviewed
in Birstein et al. 1997 this volume). In many of these cases, it is not easy to discriminate between parental species and
the hybrids.

Full text

Please indicate author’s corrections in blue, setting errors in red
000000 EBFI vol 48 pag 156 ORD.NO 00000.0
Portraits of a juvenile Huso huso 23 cm TL from the Ryal Ontario Museum collection (given originally as Caspian Sea fish to Montreal
Expo 1967) above the head of Acipenser schrenckii 81 cm TL from the Amur River stock held at the Propa-Gen International, Komadi,
Hungary. Originals by Paul Vecsei, 1996.

Please indicate author’s corrections in blue, setting errors in red
115067 EBFI ART.NO 1655 (182) ORD.NO 231386.VP
Environmental Biology of Fishes 48: 157–163, 1997.
1997 Kluwer Academic Publishers. Printed in the Netherlands.
How many species are there within the genus Acipenser?
Vadim J. Birstein1& William E. Bemis2
1The Sturgeon Society, 331 West 57th Street, Suite 159, New York, NY 10019, U.S.A.
2Department of Biology and Graduate Program in Organismic and Evolutionary Biology, University of Mas-
sachusetts, Amherst, MA 01003, U.S.A.
Received 23.4.1996 Accepted 17.5.1996
Key words: Acipenser baerii, A. brevirostrum, A. dabryanus, A. fulvescens, A. gueldenstaedtii, A. medirostris,
A. mikadoi, A. naccarii, A. nudiventris, A. oxyrinchus, A. persicus, A. ruthenus, A. schrenckii, A. sinensis, A.
stellatus, A. sturio, A. transmontanus, Huso huso, H. dauricus
In their paper in this volume Bemis et al. (1997) ask: ‘How many valid species of Acipenser should we recog-
nize?’ Although a partial answer to this question is presented in their Table 5, we discovered in the course of
preparing this volume that some additional commentary is needed. In fact, there are two questions: (1) how
many species should be recognized? and (2) what scientific names should be used for some of the species? The
sympatric distributions of most species of sturgeons set the stage for much confusion about species bounda-
ries, but the situation is actually much more complicated. Confusion about the number of species of sturgeons
living within the same basin can result from the often close morphological and meristic similarities of certain
species of Acipenser, particularly during juvenile period. Moreover, we still have remarkably inadequate
knowledge of the comparative anatomy of the species of Acipenser: no modern study has ever attempted a
comprehensive examination of all species, and it is impossible to rely on literature for the sorts of comparisons
that must be made (for more on this general problem, see Grande & Bemis 1991, 1997). Most classical descrip-
tions and comparative anatomical studies relied upon small sample sizes. Voucher specimens of large stur-
geons are especially rare in most historical collections, and type specimens (if available at all) are seldom
prepared in ways that are suitable for making detailed anatomical comparisons (e.g., many skins are simply
overstuffed with straw, so that all internal structures are lost). Intraspecific morphological and meristic poly-
morphisms occur in all species of acipenserids, and in most cases we have very poor knowledge of differences
that develop during ontogeny, particularly changes in such features as the shape of the rostrum (Bemis et al.
1997). Another problem is the ease of hybridization between different species of sturgeons (reviewed in
Birstein et al. 1997 this volume). In many of these cases, it is not easy to discriminate between parental species
and the hybrids.
Two opposite tendencies appeared in the literature
on the genus Acipenser. (1) Recognizably different
species have been considered to be the same spe-
cies. This situation is illustrated below by two spe-
cies pairs, A. gueldenstaedtii and A. persicus and A.
medirostris and A. mikadoi. (2) Some authors ele-
vated many forms to the rank of species. For in-
stance, Duméril (1870) described six subgenera of
Acipenserwith more than 30 species of Acipenserin
five of them (he considered Huso as the sixth sub-
genus of Acipenser). Most of the species described
by Duméril (1870) have long since been recognized
as conspecific with other well-known species.
We still do not know the number of species of
Acipenser, and may never know it because of over-
fishing and habitat destruction in Europe and Asia,

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115067 EBFI ART.NO 1655 (182) ORD.NO 231386.VP
158
1In the literature on genetics, molecular phylogenetics and sys-
tematics, the taxonomic unit subspecies is often preserved
(Avise 1994, Mallet 1995). Avise & Ball (1990) and Avise (1994, p
253) suggested that we recognize ‘by the evidence of concordant
phylogenetic partitions at multiple independent genetic attri-
butes’. ‘When phylogenetic concordance is exhibited across ge-
netic characters solely because of extrinsic barriers to reproduc-
tion, subspecies status is suggested’. It is evident that according
to these terminology, populations of the same species of stur-
geon in disjunct sea basins (e.g., Caspian and Black seas), could
be considered as subspecies.
which have quickly eliminated sturgeons from cer-
tain river basins (see discussions in this volume by
Bacalbas¸a-Dobrovici 1997, Khodorevskaya et al.
1997, Krykhtin & Svirskii 1997, Wei et al. 1997).
Therefore, we probably have already lost forever
the opportunity to study some species of Acipenser.
In the meantime, it is clear that genetic and mole-
cular phylogenetic approaches are increasingly cru-
cial for the recognition of sturgeon species and their
relationships (for discussion, see Birstein et al. 1997
this volume).
In Eurasia, the genus Acipenseris centered upon
three main basins: (1) the Black Sea and Sea of
Azov, (2) Caspian Sea, and (3) the Aral Sea. Each of
three main species of Acipenser, A. gueldenstaedtii
Brandt, 1833, A. stellatus Pallas, 1771, and A. nudi-
ventris Lovetsky, 1828 were described as having
subspecies or forms in these basins (see Berg 1948,
Shubina et al. 1989, Sokolov & Vasilev 1989a, Vla-
senko et al. 1989a). If we follow the view on nomen-
clature of species discussed by Holcˇík & Jedlicˇka
(1994), then the concept of subspecies and trinomial
nomenclature is inefficient. Therefore, we consider
all intraspecies forms and subspecies of A. guelden-
staedtii, A. stellatus, and A. nudiventris invalid until
detailed molecular and morphological studies of
different forms within these species can be per-
formed.1The same is true for A. ruthenus Linnaeus,
1758, for which a few intraspecies forms were de-
scribed by different authors (see Berg 1948, Soko-
lov & Vasilev 1989b).
An example helps to illustrate the taxonomic
frustration of sturgeon biologists. Acipenser persi-
cus was described as a valid species by Borodin in
1897 (Borodin 1897, 1926), but it was later consid-
ered to be a subspecies (Berg 1934), and, still later,
again regarded as a valid species (see Vlasenko et al.
1989b, Birstein & Bemis 1997 this volume, for dis-
cussion). Moreover, Artyukhin & Zarkua (1986) de-
scribed two subspecies within A. persicus: the popu-
lation inhabiting the Caspian Sea they named as A.
persicus persicus Borodin, 1897, and the population
inhabiting the Black Sea, as A. persicus colchicus
Marti, 1940. Although some Russian authors follow
this nomenclature (Pavlov et al. 1994), additional
support from genetic and molecular data is desirable.
The validity of some Asian species and subspecies
of Acipenser is questionable. For example, Ruban
(1997 this volume) reviewed and presented new data
on the Siberian sturgeon, A. baerii Brandt, 1869,
which has an extremely wide range. Ruban’s new
work supports the traditionally recognized subspe-
cies (A.b. baerii, A.b. baicalensis and A.b. stenor-
rhynchus,e.g., Sokolov & Vasiliev 1989c). No genet-
ic study on the subspecies of A. baerii is yet avail-
able.
The three far eastern Asian species, A. schrencki
Brand, 1869 of the Amur River, and A. dabryanus
Duméril, 1868, and A. sinensis Gray, 1834 of the
Yangtze River are certainly valid (see Krykhtin &
Svirskii 1997, Wei et al. 1997, Zhuang et al. 1997, all
this volume). Chinese sturgeon, A. sinensis, from the
Pearl River differ morphologically from those of the
Yangtze River, but whether this difference warrants
separate species status is not clear (Wei et al. 1997).
The nomenclature and species status of the so-
called ‘green sturgeon’ and ‘Sakhalin sturgeon’ of
the Pacific Northwest of America and northeastern
Pacific in Asia has been particularly confusing.
Ayres (1854) described the American green stur-
geon, A. medirostris. Nearly 40 years later, Hilgen-
dorf (1892) described an Asian species caught in the
northern waters of Japan as A. mikadoi, and
Schmidt (1904) soon thereafter referred a sturgeon
caught in the Aniwa Bay of Sakhalin Island to A.
mikadoi. However, Berg (1911, 1948) considered this
Sakhalin sturgeon to be conspecific with the Amer-
ican green sturgeon, A. medirostris. Schmidt (1950)
eventually reconsidered his 1904 view, and named
Sakhalin sturgeon as a subspecies of A. medirostris,
A. medirostris mikadoi (Schmidt, 1950). Therefore,
three names coexisted in the literature for the Sak-
halin sturgeon: A. mikadoi (Okada & Matsubara

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2Since the description of the species, the name A. oxyrinchus has
changed a few times. Mitchill described this species in 1815 under
the name A. oxyrinchus (Mitchill, 1815). Later, the name was
changed to A. oxyrhynchus and an incorrect date of publication
(1814) began to be cited widely (e.g., Vladykov & Greely 1963).
Also, A. oxyrinchus desotoi was first described under the name
A. oxyrhynchus de sotoi (Vladykov 1955). In this volume we fol-
low Smith & Clugston (1997) and use the names A. o. oxyrinchus
and A. oxyrinchus desotoi.
1938, Matsubara 1955), A. medirostris (Berg 1948,
Andriyashev & Panin 1953, Masuda et al. 1984,
Houston 1988, Artyukhin & Andronov 1990, Pavlov
et al. 1994), and A. medirostris mikadoi (Lindberg &
Legeza 1965, Shilin 1995). Recently Birstein (Bir-
stein et al. 1993, Birstein 1993) noted the difference
in ploidy between the Sakhalin sturgeon and Amer-
ican green sturgeon, and suggested that they should
be considered different species, A. mikadoi Hilgen-
dorf, 1892, and A. medirostris Ayres, 1854, respec-
tively. Molecular data on three mitochondrial genes
presented in this volume (Birstein & DeSalle 1997)
also show great differences between these two spe-
cies. Other molecular data obtained show a close ge-
netic relationship of A. medirostris to another
American Pacific sturgeon species, A. transmonta-
nus (Brown et al. 1996, Birstein et al. 1997). There-
fore, A. mikadoi and A. medirostris should be con-
sidered as morphologically similar, but genetically
different, species. The Sakhalin sturgeon inhabits
the Sea of Japan up to the Korean Peninsula and wa-
ters to the north from Hokkaido Island (Berg 1948,
Lindberg & Legeza 1965). It occurs in the mouths of
small rivers of the Asian far east and Korean Penin-
sula, as well as the Amur River, and rivers of the
Sakhalin Island. Now it spawns in the Tumnin (Dat-
ta) River in the Russian far east (Artuykhin & An-
dronov 1990), and historically it also spawned in the
Ishikari and Teshio rivers of Hokkaido Island (Oka-
da 1955). Acipenser medirostris ranges from the Gulf
of Alaska to southern California (Houston 1988),
with three known spawning rivers: the Sacramento
and Klamath rivers in California and the Rogue Riv-
er in Oregon (Moyle et al. 1994).
Two other species of sturgeons are usually men-
tioned in descriptions of the fish fauna of Japan, A.
kikuchii Jordan & Snyder, 1901, and A. multiscuta-
tus Tanaka, 1908 (Okada 1959–1960, Masuda et al.
1984, Rochard et al. 1991). Only one specimen of A.
kikuchiiis known (Jordan & Snyder 1901, 1906), and
this species was re-identified as A. sinensis (Take-
uchi 1979). Only a few specimens of A. multiscutatus
were described (Tanaka 1908, Fowler 1941, Matsub-
ara 1955). It seems that these specimens are morph-
ologically similar to A. schrenckii (Lindberg & Le-
geza 1956) and are probably conspecific with A.
schrenckii. It is most improbable that a sturgeon spe-
cies could be restricted only to Japan and not inhab-
iting Asian continental waters (Artyukhin & An-
dronov 1990). There are no new reports on the catch
of A. multiscutatus in Japanese literature (see a
compilation of data in Honma 1988) since the re-
view of Okada (1959-1960). Therefore, A. multiscu-
tatus is most probably a synonym of A. schrencki.
It is easy to distinguish the second Pacific North
American species, A. transmontanus Richardson,
1836, the freshwater North American A. fulvescens
Rafinesque, 1817, and one of the two Atlantic North
American sturgeons, A. brevirostrum Le Sueur,
1818 (Vladykov & Greeley 1963, Scott & Crossman
1973, Lee et al. 1980). Molecular data on the struc-
ture of the control region of mtDNA not only sup-
ported close relationships of two Pacific North
American sturgeon species, A. medirostris and A.
transmontanus, but also showed a significant genet-
ic difference between these species, A. fulvescens,
and the second Atlantic North American species,
A. oxyrinchus (Brown et al. 1996).
American and the European Atlantic sturgeon
were long considered to be one species, A. sturio
Linnaeus, 1758. In this older terminology, the
American Atlantic sturgeon was regarded as sub-
species A. sturio oxyrinchus, with the European At-
lantic sturgeon being known as A. sturio sturio (see
Smith 1891, Vladykov & Greeley 1963). Magnin &
Beaulieu (1963) suggested elevation of these sub-
species to species ranks, with the European form re-
taining the name A. sturio Linnaeus, 1758, and
American form named A. oxyrinchus Mitchill, 1815.
Two subspecies, the Atlantic sturgeon, A.o. oxyrin-
chus, and the Gulf coast sturgeon, A.o. desotoi,
were described within A. oxyrinchus (Vladykov
1955, Vladykov & Greeley 1963).2These two sub-
species of A. oxyrinchus are morphologically simi-
lar, with the most significant known difference be-

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115067 EBFI ART.NO 1655 (182) ORD.NO 231386.VP
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3Such decisions to use the originally published spellings of
names, regardless of subsequent practices, are based on the In-
ternational Code of Zoological Nomenclature (Ride et al. 1985).
For a specific explanation of rules, see Chapters 31 and 33 of the
International Code of Zoological Nomenclature. 1985, 3rd ed.
International Trust for Zoological Nomenclature, London.
ing the length of the spleen (in A.o. oxyrinchus the
spleen is statistically smaller than it is in A.o. de
sotoi, Wooley 1985). Molecular data are more infor-
mative for the discrimination between subspecies.
Comparison of the control region of mtDNA se-
quences of both subspecies showed three fixed nu-
cleotide changes in that region (Ong et al. 1996).
Bowen & Avise (1990) suggested that there is gene-
tic structuring among A. oxyrinchus from various
drainages of the North American Atlantic coast.
Recently, analyses of the control regions of mtDNA
supported this hypothesis: Atlantic sturgeon pop-
ulations in the Saint Lawrence and Saint John rivers
(Canada), the Hudson River (U.S.A.), and rivers of
Georgia (U.S A.) are genetically distinct (Waldman
et al. 1996a,b).
Unpublished results of Birstein & DeSalle on the
sequences of three more genes of mtDNA (cyto-
chrome b, 12S rRNA, and 16S rRNA) also show a
genetic difference between the two subspecies of A.
oxyrinchus (one fixed nucleotide change in cyto-
chrome bgene). The analysis of these genes demon-
strated that the European A. sturio is the only stur-
geon species closely related to A. oxyrinchus. More-
over, it appeared that there is a significant genetic
differentiation within A. sturio. Birstein & DeSalle
studied samples from two specimens of A. sturio
caught in the Gironde estuary system (Dorgonne
and Garonne rivers) and in the North Sea. The ge-
netic difference between two individuals of A. stu-
rio (6 nucleotide changes in the region of cyto-
chrome banalyzed) was even more than the differ-
ence between subspecies of A. oxyrinchus (one
change). These data seem to support the difference
in some meristic characters between specimens
from the Baltic Sea, from one side, and specimens
from the Atlantic Ocean, Mediterranean and Black
seas, from the other (Marti 1939, Magnin 1963, Ni-
nua 1976, Holcˇík et al. 1989). Because A. sturio has
almost disappeared in the wild (Holcˇík et al. 1989),
more work should be done in museum collections
on the comparison of specimens from different
populations. This is especially important in terms of
recovery projects for this species (Hochleithner
1995, Williot et al. 1997, this volume).
The last species in the genus Acipenser is the
Adriatic sturgeon, A. naccarii Bonaparte, 1836. It is
restricted to the Adriatic only and resembles A.
gueldenstaedtii in meristic characters (Tortonese
1989).
Since Berg (1904), Huso huso Brandt, 1869 and
H. dauricus Georgi, 1775 were considered as repre-
sentatives of a distinct genus Huso, not Acipenseras
they were usually considered in the 19th century (al-
so see Findeis 1997, this volume). Results of recent
molecular studies, however (see Birstein et al. 1997
this volume) showed that the two species of Huso
do not form a separate monophyletic group, but are
inserted among species of Acipenser. This result re-
activates the old discussion on the validity of the ge-
nus Huso. In the absence of detailed work on this
problem, it makes sense for now to regard Husoas a
genus based on morphological and anatomical data
(Findeis 1997 this volume). Also, a few subspecies
were described within H. huso (reviewed in Piro-
govskii et al. 1989). For instance, some authors still
consider the Sea of Azov population of H. huso as
Huso huso maeoticus Salnikov & Myatskii, 1934
(Pavlov et al. 1994). Until genetic differences can be
shown in combination with morphology, we recom-
mend the name H. huso for the Mediterranean,
Black, Azov, and Caspian sea populations of belu-
ga.
In conclusion, we recognize 17 valid extant spe-
cies within Acipenser. For the moment, we accept
that two species (A. baerii and A. oxyrinchus) con-
tain subspecies. Further genetic and molecular
studies will generate new data for correction of our
contemporary knowledge about some of the spe-
cies, including A. sturio.
A final note regarding the names of sturgeon spe-
cies concerns the need to return to the originally
published spellings for names of genera and spe-
cies3. In addition to two recent clarifications on the
correct spelling of species names for Siberian (A.
baerii, see Ruban 1997, this volume) and American
Atlantic sturgeon (A. oxyrinchus, see Gilbert 1992),
we note the following correct spelling for two other

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115067 EBFI ART.NO 1655 (182) ORD.NO 231386.VP
161
species of Acipenser. The scientific name of the
Russian sturgeon should be spelled Acipenser guel-
denstaedtii Brandt, 1833, and the scientific name of
the Amur River sturgeon should be spelled Acipen-
ser schrenckii Brandt, 1869.
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