MORE natural grouping of species with a better understanding
of their affinities expressed in a simpler taxonomy has been one
of our principal objects for many years. Among the most popular
groups of birds, the waterfowl, as the Anatidae are known, have
perhaps been more arbitrarily classified than any other. Because of
the general interest attached to these birds, we have thought that it
might be useful to revise the group and to state our views on the
relationships within it. Delacour (1933, 1936, 1938) has already
published several papers on the subject. But since their appearance
our knowledge has advanced considerably, and the present paper is
a corrected, expanded, and up-to-date version, in English, of these
earlier articles.
For over 20 years Delacour maintained in the park of the Chateau
de Cl[res, in Normandy, the greatest collection of live waterfowl ever
gathered. All existing species of swans, geese, tree ducks, and shel-
drakes were represented in it; and of all the other ducks, only 26
species were missing. They lived under conditions approaching those
of the wild state, and consequently they bred freely and displayed
their natural behavior, including their courtship. In addition,
we have observed many of the rarer exotic species in their natural
habitat, and we have extensively studied museum series at the
American Museum and elsewhere.
We also have benefited by the work of many authors, ornitholo-
gists, sportsmen, and breeders, particularly by the excellent pioneer
studies of Dr. O. Heinroth (1910; 1911; and with M. Heinroth, 1928).
For many years, Delacour has exchanged views, notes, and specimens
with Dr. K. Lorenz, of Vienna, on the subject of the display and
affinities of the Anatidae, with a view to later joint publication. The
files kept at Cl[res were destroyed by a fire in 1939. We know that
Dr. Lorenz has since published a paper on the subject, but this is
unfortunately not yet available to us (Lorenz, 1941). It will be
interesting to compare his conclusions with ours.
The classification of ducks which has been accepted up to the
present is more than 50 years old. In spite of criticism by a number
4 THE WILSON BULLETIN March 1945
Vol. 57, No. 1
of recent authors, it has been more or less followed in all recent
works, such as Phillips' "A Natural History of the Ducks" (1922-26);
Peters' "Check-List of Birds of the World" (1931); and the fourth
edition of "The A.O.U. Check-List of North American Birds" (1931).
In fact, Salvadori's classification in the "Catalogue of Birds in the
British Museum" (Vol. 27, 1895) is in some ways more acceptable
than several later ones. All these systems have the weakness of being
based exclusively on a small selection of morphological characters,
primarily on the shape of the bill and feet. Nothing could be more
misleading, for the form of bill or feet is entirely functional and
undoubtedly often recently acquired, representing merely a secondary
adaptation that is repeated in widely separate groups. It is useful
in distinguishing species but has certainly no deeper phylogenetic
significance. Non-adaptive morphological characters are far more
useful taxonomically. The most important of these in the duck
family are: pattern of tarsus (whether scutellate or reticulate in
front), a very fundamental character in the family; plumage pattern
in both adults and young, the downy young of most of the nine
main groups in the family having a very characteristic pattern;
presence or absence of a double annual molt; posture, general body
proportions, length of neck, and shape of head, all of which show
characteristic differences among the nine main groups; characteristics
of the internal anatomy, especially the structure and shape of the
syrinx and trachea (as Heinroth has repeatedly pointed out). Simi-
larly, biological characters--almost entirely ignored by the currently
adopted systems of classification--are of paramount importance to
the classifier, for habits and behavior are deeply rooted and are
usually the product of very ancient evolution. In the duck family
the main points are pair formation, displays, nesting, and feeding
habits. To be satisfactory and reliable, any system must be based
on the greatest possible number of known characters, and an over-
valuation of a few primarily functional characters has led to great
confusion in the taxonomy of the Anatidae.
Several branches, for example, the pochard group, the goldeneye-
merganser-scoter group, and the stiff-tailed duck group, have de-
veloped into divers par excellence, and are structurally rather similar
to one another. However, their non-adaptive characters, such as
the general proportions of the body, the color pattern of the downy
young, the structure of the syrinx, and the courtship performances,
are sufficiently different among the three groups to suggest that
the three are not at all closely related.
A further instance is that of the so-called geese. In addition to
the typical geese of the Anser-Branta group, there are a number of
We refer to his detailed account (O. and M. Heinroth, 1928:226-229). The taxonomic
advantage of this structure lies in the fact that its shape is not easily modified by any peculiar
adaptations of a given species. It tends to be phylogenetically conservative.
J. I)elacour THE FAMILY ANATIDAE 5
Ernst Mayr
"goose-like" genera such as the Cape Barren Goose (Cereopsis), the
Pied Goose (Anseranas), the Maned Goose (Chenonetta), the South
American"geese" (Chlophaga), the Egyptian Goose (Alopochen), and
the group commonly known as sheldrakes ("Casarca" and Tadorna),
all of which are characterized by rather large size and long legs,
many by grazing habits. They are the "ungulates" of the duck
family. Again the evidence is rather strong that the goose-like
features were acquired independently by the several groups. This
adaptability poses a problem to the classifier of the duck family
which by no means has been solved entirely. However, even though
the position of certain species and genera is still uncertain, the study
of live specimens and the consideration of previously neglected
morphological characters have shed much light on the relationship
of the birds included in this family.
This might be an appropriate place to state again our views on
the subject of zoological nomenclature. We have always stood for
the strict application of the law of priority, but according to the rules
and opinions of the International Commission. These provide for
corrections in evident cases of misprints, of lapsus calami, and of
errors in transcription. There is sometimes a certain difficulty in
determining the validity of the evidence for such mistakes, but
moderate degrees of common sense and classical scholarship are
usually sufficient to enable a zoologist to make up his mind. To
retain the original spelling of a name, however wrong it evidently is,
constitutes a retrograde solution too easy and too uncritical. It is
a great pity that both the A.O.U. and the B.O.U. committees on
nomenclature have recently chosen to follow such a course. We are
absolutely opposed to it, now as in the past, 2 and consequently we
correct all misprints, lapsus calami, and errors in transcription.
Also, according to the same rules, the endings of the adjectival
species names should agree with the gender of the genus, and Greek
endings should not be latinized. Furthermore, we conserve long-used
names, unless the necessity for a change is unequivocally established.
We believe in large genera, since it is the function of the generic
name to express relationship (as an aid to the memory), not distinct-
ness, which is expressed by the species name. Even Peters, who is
certainly not a splitter, recognizes in the family of Anatidae 62 genera
for 167 species (an average of 2.7 species per genus), and 42 (70
per cent) of his genera are monotypic. The A.O.U. Check-List goes
even further. Such nomenclature comes dangerously close to being
mononomial. The modern broadening of the species concept (Mayr,
1942:102-122) necessitates a corresponding adjustment of the genus
limits. In the classification here presented we recognize 40 genera
for 144 species (3.6 species per genus). It is interesting to find that
a number of the vernacular names for the waterfowl--swans, scoters,
See Delacour, 1931, L'Oisau, n. s. 1:438-440.
March 1945
6 THE WILSON BULLETIN Vo. s7. o. i
eiders, mergansers--delimit natural groups more accurately than the
generic names currently used by taxonomists. It has been our en-
deavor to bring the generic nomenclature of the duck family back
to an expression of these natural groups. The proponents of generic
splitting forget that if morphological difference is acknowledged as
an inevitable generic criterion, sooner or later nearly every species
will deserve a genus of its own. Generic subdivision carried to extremes
not only places an unbearable burden on the memory of the taxonom-
ist, but also completely obliterates the difference between the weak
and the really distinct genera. The differences separating Anser,
lVhilacte, and Chen; Arias, Nettion, and Dafila; Aix and Dendronessa;
or Somateria, Arctonetta and lVolysticta, are certainly very slight com-
pared with the differences separating Anser, Cygnus, and Coscoroba;
or Chlophaga, Alopochen, and Tadorna; or Arias, Malacorhynchus,
Tachyeres, and Stictonetta. Since no category above the genus can be
expressed in the scientific name, the splitter has no way of making a
distinction between "weak" and "good" genera. We consider this
another strong argument in favor of recognizing only pronounced
genera. (Mayr, 1942:280-291.)
A NEW CLASSIICATIOBI O THE ABIATIDAE
The new classification of the duck family that we propose attempts
to do two things: to arrange the species in related groups and in a
natural sequence, and to adjust the nomenclature of species and
genera to progressive concepts of these categories.
Following the popular classification of this family, the first
taxonomists divided the waterfowl into: swans, geese, ducks, and
mergansers. As more and more was learned about the anatomy as
well as about the habits of members of the family, it was realized
that this simple division was unsatisfactory. For example, Linnaeus
included in the duck genus Arias such widely divergent species as
the river ducks of the mallard and teal type, the diving ducks of the
scaup-pochard group ("Nyroca" = Aythya), the diving ducks of the
goldeneye-scoter-eider group (Mergini), the tree ducks (Dendrocygna),
and the sheldrakes (Tadorna). Although subsequent classifiers recog-
nized some of these subdivisions, they were guided in their reclassi-
fication mainly by the shape of the bill or by the presence or absence
of the diving habit.
All the ducks, geese, and swans, including even the most aberrant
species, are so much alike in their basic structure and habits that
there can be no doubt that those modern authors are right who in-
clude all waterfowl in a single family, the Anatidae. Within this
family a number of groups of genera can be recognized, but they are
clearly arranged in two main groups, which we admit as two sub-
families:
J. Delacour THE FAMILY ANATIDAE 7
Ernst Mayr
(1) Anserinae. This subfamily includes the swans, geese, and the
whistling ducks ("tree" ducks). The attributes of the group are a
"goose-like" posture and body shape (with a long neck); a tarsus
reticulated in front; a single annual molt; absence of sexual dimor-
phisin in plumage, voice, and structure of the syrinx. Displays are
simple and are similar in the two sexes.
(2) Anatinae. This subfamily includes the rest of the Anatidae.
The attributes of the group are a tarsus that is scutellated in front
(with a few exceptions); a double annual molt; sexual dimorphism
in plumage (frequent), in voice and structure of syrinx (usual).
Displays are usually elaborate and different in the two sexes.
Within each subfamily further subdivisions are recognizable. We
use the term tribes (with the ending -ini) for such groups of genera,
following a custom that is widespread in entomology. The reasons
for the recognition as well as for the delimitation of these tribes
will be found in the following discussion. The phylogenetic relation-
ships within the duck family are diagrammed in Figure 1.
Mcr ini
Merganlrini
/ini Oxy rrini
Ay!
AnaMni
Tdornini
Indrocygnini
Ans
An$1/2rina1/2
. Cairinini / ,,'
r J / ,
Anatina1/2
Anatida1/2
I
Anhimae
Phonicopteri .--/
Figure 1. Diagram of the theoretical relationships of the subfamilies and tribes
of the Anatidae.