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).
Nomenclatural
Notes
Type
specimens: dead or alive?
(1) A. Wakeham-Dawson
and Solene Morris
Secretariat, International Commission
on Zoological Nomenclature, c/o The Natural
History Museum, Cromwell Road, London
SW7 5BD, U.K.
Philip Tubbs
16 New Road, Ham, Richmond, Surrey
TW10 7HY, U.K.
It is a widespread
misunderstanding that an animal species
cannot be given a scientific name until
a specimen has been killed and preserved
as the name-bearing type specimen for that
taxon. An example of this misunderstanding
was published in The Daily Telegraph
magazine Weekend (London; 17 November
2001). The leading article by Sandy Mitchell
claimed that it had been necessary for
a scientist (Julia Robinson Dean) to return
to Indonesia to kill a rare bird before
she could name it and thereby allow it
to be added to a list of protected species.
A letter outlining the error was sent in
response to the magazine article by the
then Executive Secretary of the Commission,
Philip Tubbs. However, the letter was not
published.
The Code does not require a museum
specimen as type material or that the naming
process requires an anatomically detailed
description to be made on the basis of
such material. However, every new name
must ‘be accompanied by a description
or definition that states in words characters
that are purported to differentiate the
taxon’
(Article 13.1.1 of the Code), and since
January 2000 the specimen (holotype) or
specimens (syntypes) on which the name
is based must be explicitly stated (Article
11.6.4.1).
In the case of the Indonesian bird,
a description based on notes from the scientist’s
notebook, or even the picture and description
that appeared in the newspaper article,
would have been sufficient to make the
name available. The holotype or syntypes
remain the specimens of which the photographs
were taken and the descriptions made, even
if they are allowed to return alive to
their natural habitat and are never seen
again. The holotype (or syntype) is not
the picture of the specimen (see Articles
72.5.6 and 73.1.4). Similarly, when a new
species is described and named on the basis
of DNA sequences, the specimen from which
these were taken remains the holotype (or
syntypes in the case of a series of specimens
from which samples are taken). For example,
a new species of Somalian shrike was named
from a living specimen that was released
after samples had been taken for DNA analysis
(see Smith, Arctander, Fjeldsa
& Amir, 1991; Hughes, 1992).
There are good reasons why a dead
specimen cannot be required for formal
naming. Capture, killing and export may
be illegal, unethical or impossible (e.g.
capture of a new taxon of fish seen from
a deep-sea submersible may not be practical)
and absence of a museum specimen to act
as holotype does not prevent the naming
process. Many thousands of names would
be invalid if dead type specimens were
mandatory. For example, many of the species
named by Linnaeus were not based on any
cited type material, and name-bearing specimens
have never been fixed for many well-known
species.
The misconception that a dead holotype
specimen is mandatory under the Code has
perhaps arisen from the wording used in
relation to designation of new species
in early editions of the International
Code of Zoological Nomenclature (1961,
1964) and the Règles Internationales
de la Nomenclature Zoologique (1905)
that preceded them. This misunderstanding
has been compounded in textbooks on taxonomy.
However, preserved specimens have
never been a mandatory requirement, although
they have been (and still are) recommended.
In 1926, the Règles were
translated into English and published as
the International Rules of Zoological Nomenclature
in the Proceedings of the Biological
Society of Washington, 39:
75-104. In this document, Recommendation
B (pp. 7R-8R) on Articles 1-3 recommended ‘that
in published descriptions of a new species
or of a new subspecies, only one specimen
should be designated as type. The specimen
itself should be labelled type’.
Recommendation B was re-presented in the
form of Article 72(a) of the First and
Second Editions (1961 & 1964) of the
Code (p. 75 in both editions). This stated
that ‘the type of each taxon of the
species-group is a single specimen’.
In Articles 72 of the Third Edition (1985,
p. 139) it is explained that ‘the
term “type”
forms part of many compound terms used
by taxonomists to distinguish between particular
kinds of specimens’. Some of these
terms do not refer to name-bearing types.
The wording of the Règles and
First and Second Editions of the Code (1961 & 1964)
might have been held to imply that a holotype
could only be designated when a dead specimen
was to hand. The Third Edition (1985) did
not state that this was not the case, but
Article 73(a)(iv) stated that ‘designation
of an illustration of a single specimen
as a holotype is to be treated as designation
of the specimen illustrated; the fact that
the specimen cannot be traced does not
of itself invalidate the designation’.
This clearly indicated that a preserved
specimen was not a mandatory requirement
of the Code. Eligibility for name-bearing
type status was stated in Article 72(c).
In addition, the introduction to the Third
Edition of the Code (1985) stated (p. xvi)
that ‘although the principle [of
name-bearing types] is fundamental, it
is still not obligatory for name-bearing
types to be designated for new species-group
taxa although the Code recommends the practice
and provides procedures by which the name-bearing
type of any species-group taxon can be
discovered and fixed’.
The introduction to the Fourth (current)
Edition of the Code (1999) states (p. xxvii)
that ‘when the name-bearing type
of a species group taxon proposed after
1999 consists of a preserved specimen or
specimens, the proposer is required to
include a statement naming the collection
in which the name-bearing type is or will
be deposited’. From this statement,
it is clear that a dead type specimen is
not essential under the Code. However,
it is desirable that this should be stated
directly, rather than just by implication,
in future editions of the Code to prevent
nomenclature and taxonomy from being wrongly
discredited in situations of biological
conservation sensitivity or where modern
techniques (e.g. blood sampling for molecular
analysis etc.) are a viable alternative
to killing specimens.
In the future, it may be possible
to describe all species solely on the results
of molecular analysis techniques from blood
or other samples taken from living animals.
For the time being, it is still desirable
to have preserved specimens at hand to
allow a full description of new taxa to
be made, and for re-examination of those
specimens at a later date.
In the case of marine organisms,
there are some old nominal species that
were based on animals only seen in the
water. As no specimens were actually obtained
these have not been considered
‘taxonomically sound’ (William
Perrin, personal communication) even though
these names remain available under the
Code. The following note by Drs Dalebout
and Scott Baker on the description of a
new whale species illustrates the value
of having preserved specimens. The use
of morphological comparison and DNA analysis
techniques allowed the determination and
description of a new animal taxon, which
would have been impossible in the absence
of preserved material.
References
Hughes, A.L. 1992.
Avian species described on the basis of
DNA only. Trends in Ecology and Evolution, 7:
2-3.
Smith, E.F.G., Arctander, P., Fjeldsa, J. & Amir,
O.G. 1991. A new species of shrike (Laniidae: Laniarius)
from Somalia, verified by DNA sequence data from the only
known individual. Ibis, 133: 227-235.
(2) Merel L. Dalebout
and C. Scott Baker
School of Biological Sciences, University
of Auckland, Private Bag 92019, Auckland,
New Zealand
Beaked whales (ZIPHIIDAE)
are among the least known of mammals (Wilson,
1992). Twelve new cetacean species have
been described in the last 100 years, of
which seven were beaked whales, primarily
of the genus Mesoplodon. This
total does not include M. bahamondi Reyes,
Van Waerebeek, Cardenas
& Yanez, 1995, a species now recognized
as synonymous with M. traversii (Gray,
1874) (van Helden et al., 2002). Given
this synonymy, the most recently described
beaked whale species was M. peruvianus Reyes,
Mead & Van Waerebeek, 1991.
Sightings of beaked whales at sea
are generally rare due to their elusive
habits and preference for deep oceanic
waters. Several species have yet to be
seen alive and the distinctiveness of others
has been questioned. Species of beaked
whales are comparatively undifferentiated
in external morphology. Species identification
is based primarily on features of cranial
morphology and, especially for the most
species-rich genus Mesoplodon,
on the size, shape and position of the
teeth in the lower jaw. All beaked whale
species (except the monotypic Tasmacetus)
have a highly reduced dentition, retaining
only one or two pairs of teeth in the lower
jaw. In genera with a single pair of teeth,
such as Mesoplodon, the teeth
develop and erupt from the gum only in
adult males. Females and juveniles are
effectively toothless. These teeth are
not used for feeding. Instead, based on
observations of scarring patterns on stranded
animals, males use these tusk-like teeth
as weapons in intra-specific combat with
other males (see Heyning, 1984). Due to
the often small number of known specimens,
pronounced sexual dimorphism and wide geographic
distribution (all oceans except the high
Arctic), the potential for the misidentification
of beaked whales based on morphological
features is considerable, even for experts.
In the mid to late 1970s, four beaked
whales (an adult male, an adult female
and two calves) were stranded within 50
miles of each other along the southern
coast of California. These animals were
identified as Mesoplodon hectori (Hector's
beaked whale) from morphology, the first
and only records of this species from the
Northern Hemisphere (Mead, 1981). Three
of the specimens were collected for the
Smithsonian Institution National Museum
of Natural History, while the fourth was
collected for the Los Angeles County Museum
of Natural History.
In 1997, a database of mitochondrial
(mt) DNA control region sequences was compiled
to assist in beaked whale species identification
(Henshaw et al., 1997; Dalebout et al.,
1998). All specimens in this reference
database were validated through examination
by experts in cetacean morphology and the
collection of diagnostic skeletal material
or photographic records following the recommen¬dations
of Dizon et al. (2000). A sequence from
one of the California specimens was included
in the database but was found to differ
from specimens of Southern Hemisphere M.
hectori and all other species in the database
at that time (Dalebout et al., 1998).
To investigate this anomaly, DNA
was extracted from cartilage and tooth
material from the remaining three California
specimens described by Mead (1981). Phylogenetic
comparisons of mtDNA control region and
cytochrome b sequences from these specimens
to a now complete reference database including
all 20 recognized beaked whale species
(Dalebout, 2002; see also www.dna-surveillance.auckland.ac.nz)
confirmed that all four are of the same
species, yet do not represent M. hectori or
any other known ziphiid species. A fifth
specimen, a calf stranded at Monterey in
1997 and initially identified as a neonate Ziphius
cavirostris (Cuvier's beaked whale)
from external morphology, is also grouped
with these anomalous California specimens
in phylogenetic analyses. These analyses
provided strong evidence that these five
specimens represent a previously undescribed
species of beaked whale (Dalebout et al.,
2002). This conclusion was confirmed through
phylogenetic analysis of nuclear DNA sequence
data (Dalebout, 2002) and supported by
re-examination of morphological features
(Dalebout et al., 2002).
A formal description of this new
species including details of diagnostic
molecular and morphological features was
given by Dalebout et al. (2002). This species,
like M. hectori, is a small beaked
whale, approximately 4 m in length, with
a relatively short rostrum (beak/upper
jaw). Both species have a single pair of
triangular teeth set at the apex of the
mandible, but there are subtle differences
in position and angle of inclination. Of
the four specimens stranded in California
in the 1970s, the adult female and one
of the calves share the same mtDNA haplotype
(the mitochondrial genome is inherited
only through the maternal line). These
specimens were found a week apart and are
probably a mother and her offspring. There
are no confirmed observations of this species
at sea and little is known of its ecology.
We assume that like many other beaked whales,
these animals eat mainly pelagic squid.
The adult male bore a number of white,
linear scars on its postcranial flanks,
probably inflicted by the teeth of conspecific
males. Although the stranding pattern of
the five specimens known to date is suggestive
of an eastern North Pacific distribution,
there are too few records to date to draw
any bounds on this. We have named this
new species Mesoplodon perrini (Perrin's
beaked whale) in tribute to the American
cetologist, William F. Perrin, of the U.S.
National Marine Fisheries Service South
West Fisheries Science Center (La Jolla,
California) for his role in the collection
of two of the known specimens of this species
and his ongoing contribution to marine
mammal science and conservation.
References
Dalebout, M.L. 2002. Species identity,
genetic diversity and molecular systematic
relationships among the Ziphiidae (beaked
whales). PhD thesis, School of Biological
Sciences, University of Auckland, Auckland,
New Zealand.
Dalebout, M.L., Mead, J.G., Baker, C.S., Baker, AX &
van Helden, A.L. 2002. A new species of beaked
whale Mesoplodon perrini sp. n. (Cetacea: Ziphiidae)
discovered through phylogenetic analyses of mitochondrial
DNA sequences. Marine Mammal Science, 18:
577-b08.
Dalebout, M.L., van Helden, A., Van Waerebeek, K. &
Baker, C.S. 1998. Molecular genetic identification
of southern hemisphere beaked whales (Cetacea: Ziphiidae). Molecular
Ecology, 7: 687-694.
Dizon, A., Baker, C.S., Cipriano, F., Lento, G.,
Palsboll, P. & Reeves, R. 2000. Molecular
genetic identification of' whales, dolphins and porpoises:
proceedings of a workshop on the forensic use of molecular
techniques to identify wildlife products in the marketplace.
NOAA Technical Memorandum NMF5 NOAA-TM-NMFS-SWFSC-286, La
Jolla, California.
Henshaw, M.D., LeDuc, R.G., Chivers, S.J. & Dizon,
A.E. 1997. Identification of beaked whales (family
Ziphiidae) using mtDNA sequences. Marine Mammal Science, 13:
487-495.
Heyning, J.E. 1984. Functional morphology
involved in intraspecific fighting of the beaked whale, Mesoplodon
carlhubbsi. Canadian Journal of Zoology, 62:
1645-1654.
Mead, J.G. 1981. First records of Mesoplodon
hectori (Ziphiidae) from the Northern Hemisphere and
a description of the adult male. Journal of Mammalogy, 62:
430-432.
Reyes, J.C., Mead, J.G. & Van Waerebeek, K. 1991.
A new species of beaked whale Mesoplodon peruvianus sp.
n. (Cetacea: Ziphiidae) from Peru. Marine Mammal Science, 7:
1-24.
Reyes, J.C., Van Waerebeek, K., Cardenas, J.C. & Yanez,
J.L. 1995. Mesoplodon bahamondi sp. n.
(Cetacea, Ziphiidae), a living beaked whale from the Juan
Fernandez Archipelago, Chile. Boletim de Museo Nacional
de Mistoria Natural, Chile, 45: 31-44.
van Helden, A.L., Baker, AX, Dalebout, M.L., Reyes,
J.C., Van Waerebeek, K. & Baker, C.S. 2002.
Resurrection of Mesoplodon traversii (Gray, 1874),
senior synonym of M. bahamondi Reyes, Van Waerebeek,
Cardenas and Yanez, 1995 (Cetacea: Ziphiidae). Marine
Mammal Science, 18: 609-621.
Wilson, E.O. 1992. Diversity of Life.
Harvard University Press, Cambridge, Massachusetts.
Acaulona peruviana Townsend,
1913 (Insecta, Diptera): application
of Article 75.8 of the Code
Ronaldo Toma
Museu de Zoologia de Universidade de São
Paulo, caixa postal 42594, São Paulo
04299-970, Brazil. (rtkuna@zipmail.com.br)
In 1913, Townsend
(p. 93) described a species of parasitic
fly (family TACHINIDAE) and named it Acaulona
peruviana. His description was based
on two reared specimens (a male and a female),
from San Jacinto, Chira valley, Piura Department,
Peru. They emerged as adults on 29 October
1912, having been collected by E.W. Rust
from adults of the cotton stainer bug Dysdercus
ruficollis (Linnaeus, 1764) (Hemiptera,
PYRRHOCORIDAE). The syntypes were deposited
in the United States National Museum, Washington
D.C. (U.S.N.M). Townsend (1913) reported
that Acaulona peruviana was comparatively
rare and that he had collected only five
specimens in the course of three years.
The capture data are as follows:
(1) one female, Somate, Rio Chira, 18 November
1910, on flower of Telanthera sp.;
(2) one male on foliage, Chapaira, Rio
Piura valley, 21 May 1911;
(3) two females, Cañada de Samán,
Chira valley, 14 February 1912, on flowers
of Philibertella flava;
(4) one female, Sullana, Chira valley,
17 February 1912, on foliage.
In 1950 Sabrosky (pp. 369-370) stated
that the cotton stainer parasite, Acaulona
peruviana, had not been formally described,
but that the name had been established
in connection with the full-page figure
published by Townsend (1928, p. 7, fig.
3). Sabrosky (1950) redescribed the species
from Townsend’s figure and designated
a neotype, an allotype, and seventeen neoparatypes.
Four of these had the same data as the
material listed by Townsend (1913).
Sabrosky (1951, p. 210), after being
alerted by Dr Claude Dupuis to his oversight
of the original description of Acaulona
peruviana, acknowledged that he had
made a mistake in redescribing the species.
However, as he had been unable to find
the syntypes of Acaulona peruviana deposited
by Townsend in the U.S.N.M., Sabrosky (1951)
assumed that they were lost and stated
that his neotype designation was still
valid.
In 1989, the two supposedly lost
original specimens on which the description
of Acaulona peruviana was based
were rediscovered in the U.S.N.M. According
to Article 75.8 of the Code: ‘if,
after the designation of a neotype, the
name-bearing type of the nominal species-group
taxon that was presumed lost is found still
to exist, on publication of that discovery
the rediscovered material again becomes
the name-bearing type and the neotype is
set aside’. As a result, Sabrosky’s
(1950) neotype designation is no longer
valid and herewith I designate the male
syntype specimen numbered U.S.N.M. 19477
as the lectotype of the nominal species Acaulona
peruviana Townsend, 1913. The taxonomic
reason underlying this lectotype designation
is that the female and (to a lesser extent)
male genitalia of species in the genus Acaulona Wulp,
1888 are very similar and it is only possible
to differentiate Acaulona peruviana from
others species of the genus Acaulona by
the morphology of the male genitalia.
The lectotype is a male fly in good
condition with the left wing separated
from the thorax and glued on a paper support
(Figure 1). It is from San Jacinto, Chira
valley, Piura Department, Peru, and was
collected by E.W. Rust. The paralectotype
is the former syntype specimen U.S.N.M.
19477 (the same number as the lectotype).
It is a female in good condition, but without
the fore left leg and middle right leg.
It has the same data as the lectotype.
Acknowledgements
I thank Dr Ubirajara
R. Martins de Souza (Museu de Zoologia
da Universidade de São Paulo) and
Dr Andrew Wakeham-Dawson (Executive Secretary,
International Commission on Zoological
Nomenclature) for reading the manuscript,
Dr Norman Woodley for the loan of specimens
and Dan Hansen for providing the bibliography.
This work was supported by a grant from
Fundação de Amparo à Pesquisa
do Estado de São Paulo.
