General
Articles and Nomenclatural Notes with
the following titles were published
on 30 September 2006 in Volume 63,
Part 3 of the Bulletin of Zoological
Nomenclature
Copies
of these General Articles and Nomenclatural
Notes can be obtained free of charge
from the Executive Secretary, The International
Commission on Zoological Nomenclature,
c/o The Natural History Museum, Cromwell
Road, London SW7 5BD, U.K. (e-mail: iczn@nhm.ac.uk).
Mammal Species
of the World. A
taxonomic and geographic reference. 2005. Don E. Wilson & DeeAnn
M. Reeder (Eds.). Ed. 3, 2 vols., 2142 pp.
Johns Hopkins University Press, Baltimore.ISBN
0–8018–8221–4
A nomenclatural review
Anthea Gentry
Littlewood, Copyhold
Lane, Cuckfield, Haywards Heath, West Sussex,
RH17 5EB, U.K. (e-mail:
alantgentry@aol.com)
Publication
of the new edition of Mammal
Species of the World is a considerable
achievement and is most welcome. Once again
it is edited by Don E. Wilson and DeeAnn
M. Reeder. The previous edition was published
in 1993 and since then there have been significant
changes. The work is now presented in two
volumes, the near doubling in size resulting
from the inclusion of accepted subspecies,
English vernacular names for each recognised
species, authorship and dates for all synonyms,
and a large increase in the number of bibliographic
references (which include publications up
to the end of 2003 and some works then in
press). There has also been an increase from
4629 recognised species in 1993 to 5416 in
the current edition. Volume 1 of the work
(pp. i-xxxv, 1–743)
contains the list of 26 specialist contributors,
preface, acknowledgments, introduction, list
of museum abbreviations and all the orders
except rodents; volume 2, which is larger,
contains the rodents (pp. 745–1600),
references (pp. 1601–1944),
index of scientific names (pp. 1945–2115)
and index of English vernacular names (pp.
2115–2142).
The work covers extant and recently
extinct species (those probably alive during
the past 500 years). Information now given
for each species includes the type locality,
distribution, a complete list of synonyms
including those for currently recognised
subspecies, status in the IUCN Red
list of threatened animals (2003), CITES appendices
(2004) and the U.S. Endangered Species Act
(2004), and a record of ICZN rulings. The
work reflects the numerous taxonomic changes
since the previous edition at both the species
and higher levels (the 26 orders in the previous
edition have now increased to 29).
During my university studies I specialised
in mammalogy and subsequently
worked as a mammal palaeontologist. While
employed in the ICZN Secretariat (1987–2002)
I researched and published a number of mammal
nomenclatural applications, sometimes co-authored
with colleagues, relating to both Recent
and fossil taxa. I also processed other mammal
cases submitted by fellow workers. I was
interested to see how names approved by the
Commission had been treated in the new Mammal
Species of the World.
A number of applications submitted
to the Commission sought to set aside a hitherto
overlooked type species designation in order
to conserve the current usage of one or more
generic names. Such applications included
Case 3058 (BZN 56: 136–141,
June 1999) on the fur seal names Arctocephalus F.
Cuvier, 1826 and Callorhinus Gray,
1859, and the sea lion names Otaria Péron,
1816 and Eumetopias Gill, 1866,
and Case 3121 (BZN 56: 255–261,
December 1999) on the rodent names Holochilus Brandt,
1835, Proechimys J.A. Allen, 1899
and Trinomys Thomas,
1921. The Commission approved these applications
in Opinion 1962 (BZN 57:
193–195, September 2000) and Opinion
1984 (BZN 58: 245–246,
September 2001) respectively. The generic names
have been included in the new edition of
Mammal Species of the
World, although the
compilers of the group which includes Holochilus state
that they have followed the application,
apparently unaware of the definitive Commission
ruling.
Other applications sought to ratify the
current spelling of a name by amending the
originally published name. These included Case
3018 (BZN 56: 262–265,
December 1999) for the deer name Mazama
gouazoubira (Fischer,
1814), Case 3033 (BZN 57:
36–38, March
2000) for the rodent name Glirulus
japonicus (Schinz,
1845), and Case 3004 (BZN 55:
165–168,
September 1998 and 57: 228–231, December
2000) for the primate family-group names LORISIDAE
Gray, 1821, GALAGIDAE Gray,
1825 and INDRIIDAE Burnett,
1828. These applications were approved in Opinion
1985 (BZN 58: 247, September
2001), Opinion 1978 (BZN 58:
159–160,
June 2001) and Opinion 1995 (BZN 59:
65–67,
March 2002) respectively; the corrected spellings
have been incorporated in the new checklist.
A number of applications sought to stabilise
the usage of names by a ruling on their authorship
and date. These included Case 3022 (BZN 58:
41–52, March 2001) for the publication
Catalogue des mammifères
du Muséum National d’Histoire
Naturelle by É. Geoffroy Saint-Hilaire
(1803) which included the mole name Scalopus and
22 specific names in several orders; Case 3178
(BZN 58: 126–132, June
2001) for the artiodactyl generic and family-group
names Hippotragus and HIPPOTRAGINAE
published by Sundevall (1845); and Case 2928
(BZN 51: 135–146, June
1994) for the generic names Philander (marsupial),
Pteropus (bat), Glis, Cuniculus and Hydrochoerus (rodents),
Meles, Lutra and Hyaena (carnivores), Tapirus (perissodactyl),
Tragulus and Giraffa (artiodactyls)
published by Brisson (1762). The Commission
took action in Opinion 2005 (BZN 59:
153–154,
June 2002), Opinion 2030 (BZN 60:
90–91,
March 2003) and Opinion 1894 (BZN 55:
64–71,
March 1998); the new edition of Mammal
Species of the World now lists all the
names with the appropriate authorship and date.
The ruling on Brisson’s (1762) names
has been welcomed by some contributors but
not by others. The entry for Cuniculus records ‘The
generic name of this taxon has been debated
. . . , but the nomenclatural instability of
this genus was resolved by the ruling by the
International Commission on Zoological Nomenclature
(1998) for the conservation of Cuniculus Brisson,
1762’.
The entry for GLIRIDAE,
however, notes ‘the unfortunate ruling
by the International Commission on Zoological
Nomenclature (1998) to conserve Glis Brisson’ and
thereby ignores all the usage of the name and
the support received, particularly from Europe-based
zoologists (a full discussion on the usage
of Glis was given in BZN 52:
90–91,
March 1995). The long discussion on the identity
and use of the name Tragulus by authors
subsequent to Brisson (1762) is mostly irrelevant.
Two applications, both related to bats,
sought to ratify the majority usage of an earlier
specific name where there were taxonomic difficulties.
Case 3073 (BZN 56: 182–186,
September 1999) proposed to establish Pipistrellus
pygmaeus (Leach, 1825) as the name for
a cryptic species confused with P.
pipistrellus (Schreber,
1774) until 1993. The Commission approved the
application in Opinion 2028 (BZN 60:
85–87,
March 2003) and, in Mammal
Species of the World,
P. mediterraneus Cabrera, 1904, the
name preferred to P.
pygmaeus by some workers,
is listed among the synonyms. Case 3095 (BZN
56: 250–254, December
1999) proposed confirmation of the majority
usage of Mystacina tuberculata Gray,
1843 and
Chalinobus tuberculatus (J.R. Forster,
1844). The Commission approved this application
in Opinion 1994 (BZN 59: 63–64,
March 2002) and in Mammal
Species of the World both
names have been adopted with M.
velutina Hutton,
1872, the name preferred by a few workers in
place of M. tuberculata, listed as a synonym.
For many domesticated animals, especially
artiodactyls, separate names exist for a wild
species and its supposed domestic derivative.
Very often the ‘domestic’ name
predates the ‘wild’ one and, although
a majority of writers have preferred to use
the ‘wild’ name for the wild species,
there has been some confusion about the application
of names and the significance to be read into
one usage rather than another. In Case 3010
(BZN 53: 28–37, March
1996) Juliet Clutton-Brock, Colin Groves and
I proposed that, where there were traditionally
separate names for a wild species and its domestic
derivative, the majority usage for the wild
species of the first available name based on
a wild population should be stabilised. This
affected 15 mammals and in these the name of
the domesticate had been established by Linnaeus
(1758, 1766) and a few other authors and was
earlier or contemporary with the name of the
wild ancestor. The name for the domesticate
had been applied by a few authors to the wild
species, thereby increasing the confusion mentioned
above.
Over the next six years our application
received 28 out of 33 comments and five brief
notes in favour of the proposals with considerable
support from workers in zoology, archaeozoology,
palaeontology, conservation, ecology, ethology
and endangered species management. A few commentators
were not in favour but this seemed to be because
they had misunderstood the intention of the
application: they assumed either that earlier
names based on domestic forms were going to
be discarded or that two alternative names
would be adopted as valid for the wild species.
We explained in published replies that neither
assumption was correct.
In March 2003 the Commission approved
the proposals (Opinion 2027, BZN 60:
81–84)
and 15 names for wild mammal progenitor species
were fixed as those based on wild populations.
Names based on domestic forms can now only
refer to domesticates.
Despite Opinion 2027 the minority and
now incorrect treatment of the names for wild
and domestic forms in Mammal
Species of the World has remained unaltered
from the 1993 edition. In the chapters on the
Perissodactyla and the Artiodactyla, compiled
by Peter Grubb, names based on domestic forms
have actually been used for wild species. Thus
the Linnaean names Equus
asinus and E. caballus,
based on domestic forms, are used for both
the wild and domestic ass and the horse respectively
with the wild species’ names
E. africanus Heuglin & Fitzinger,
1866 and E. ferus Boddaert, 1785 cited
as synonyms. Similarly, Bos
taurus Linnaeus,
1758,
based on domestic cattle, is used for the extinct
wild aurochs in preference to the almost universally
used B. primigenius Bojanus, 1827.
The Linnaean names Ovis aries and
Capra hircus are still given as the
wild progenitors of sheep and goats, which
usually appear as O.
orientalis Gmelin, 1774
and C. aegagrus Erxleben, 1777. This
erroneous nomenclature has not been followed
in other chapters: in the Carnivora Felis
catus Linnaeus,
1758, the domestic cat, is treated as distinct
from F. silvestris Schreber, 1777,
the wild cat, and in the Rodentia Cavia
porcellus Linnaeus,
1758 is restricted to the domestic form of
the guinea pig.
In using the Linnaean names Equus
asinus, E.
caballus, Camelus bactrianus
(domestic Bactrian camel), Bos
bubalis (domestic
water buffalo) and B.
grunniens (domestic yak)
for wild species, Grubb notes the Commission
ruling and states that it has not been demonstrated
that most authors have used the names based
on wild taxa for the species. On the contrary,
the application (para. 7) cited 15 recent reference
works in which the names based on wild populations
have been used for the wild species, and noted
that there were numerous publications in all
fields which had adopted them. Moreover, many
published comments supported our proposals,
as noted above.
In adopting the names Equus
asinus and E. caballus,
Grubb states that Opinion 2027 did not explicitly
specify which name was to be assigned to the
whole species by those who consider wild and
domestic populations to be conspecific. Again,
in adopting the names Camelus
bactrianus, Bos
frontalis (domestic gaur), B.
bubalis, B.
grunniens, Ovis
aries and Capra
aegagrus, Grubb
states that it might still be valid for those
who consider the wild and domestic forms to
be conspecific to employ the senior (domestic)
name for the name of the species. In support
of his statements Grubb cites a 1997 comment
by Walter Bock, six years before the Opinion
was published. It has to be pointed out that
my colleagues and I replied to Bock in the
same issue of the Bulletin (BZN 54(2),
June 1997) and then published two further comments
(BZN 58: 233–234, September
2001 and 59: 48–50, March 2002). We set
out the situation in the usage of names for
wild species with domestic derivatives, how
this might be seen to differ from a strict
interpretation of the Code, and the consequences
that would result from Commission approval
of the proposals. Our intentions regarding
the names for wild and domestic forms, both
when treated as separate species (two names)
and when included in one species (one name),
were stated in all three comments, and in both
the second and third comments we made
clear that ‘Approval of our proposals
by the Commission will merely ratify the
current nomenclatural situation: names based
on wild populations will continue to be used
for wild species and will include those for
domestic forms if these are considered conspecific’.
Similar comments were made by Corbet (BZN 53:
193), Kitchener (BZN 53: 194)
and Uerpmann (BZN 58: 231).
Following all this, the outcome was that Opinion
2027 does state explicitly (BZN 60:
83): ‘The
names listed [pp. 81–82]
in the ruling above, which are the first available
names in use based on wild populations, apply
to wild species and include those for their
domestic derivatives if these are not distinguishable’.
Grubb himself has admitted that Ovis
aries and Bos
frontalis, the names for
domestic sheep and domestic gaur, have not
been used for wild species and, more seriously,
that all wild taxa recorded as endangered in
IUCN (2003) and CITES (2004) publications have
been listed under the names for the wild ancestral
species.
Another consideration reveals an unappreciated
benefit of the Commission ruling which will
eliminate much future confusion. Following
publication of Opinion 2027, Gentry, Clutton-Brock & Groves
(2004) set out the history and consequences
of the ruling. We noted, citing recent papers,
that genetic analyses have demonstrated that
there are two or more lineages in several domestic
animals (including cattle, water buffalo, sheep,
goat, pig, horse, llama and alpaca) derived
from two or more wild ancestral species or
subspecies from different geographical areas.
There is also genetic evidence of hybridisation
of domestic animals with wild species and other
domestic forms during domestication. Still
more recent publications (Dobney & Larson,
2006, and cited references; Zeder, Bradley,
Emshwiller & Smith,
2006) have shown that the history of domestication
is considerably more complex than originally
thought and that the majority of modern domestic
animals have multiple ancestors, often in different
species. This makes it unsafe to assume a one-to-one
equivalence between domesticates and single
ancestors and to give it a formal expression
in their names.
Under Article 17.2 of the Code the availability
of specific names for animals is not affected
by hybrid origins, so Gentry et al. (2004)
acted legitimately in recommending that names
based on domestic forms be adopted for the
domestic derivatives. Under Article 23.8, however,
a specific name for an animal later found to
be hybrid must not be used as the valid name
for either, or any, of the parental species
even if it is older than all other available
names for them. In adopting names based on
hybrid domestic animals for wild progenitor
species Grubb has ignored this Article of the
Code.
To my knowledge the Code and Commission
rulings have not hitherto been flouted on such
a large scale and on such unsafe grounds. Applications
submitted to the Commission to resolve nomenclatural
difficulties are made on behalf of the zoological
community as a whole. Publication of cases
in the BZN and on the ICZN website includes
an invitation to zoologists to comment or make
alternative suggestions, and a two-thirds majority
of those Commissioners voting is needed for
approval of proposals. Decisions of the Commission
are thus arrived at democratically and, in
the interests of universality, clarity and
stability, need to be followed.
References
Dobney,
K. & Larson,
G. 2006. Genetics and animal domestication:
new windows on an elusive process. Journal
of Zoology, 269: 261–271.
Gentry, A., Clutton-Brock,
J. & Groves,
C.P. 2004. The naming of wild animal
species and their domestic derivatives. Journal
of Archaeological Science, 31: 645–651.
Zeder, M.A., Bradley,
D.G., Emshwiller, E. & Smith,
B.D. (Eds.). 2006. Documenting
domestication: new genetic and archaeological
paradigms. 375
pp., 40 illustrations, 120 figs.
University of California Press, Berkeley.
