NOVITATES - American Museum of Natural History
NOVITATES - American Museum of Natural History
NOVITATES - American Museum of Natural History
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AMEIRI[CAN MUSEUM<br />
<strong>NOVITATES</strong><br />
PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY<br />
CITY OF NEW YORK APRIL 2, 1951 NUMBER 1496<br />
A CLASSIFICATION OF RECENT BIRDS<br />
BY ERNST MAYR AND DEAN AMADON<br />
During the course <strong>of</strong> incorporating the Rothschild Collection <strong>of</strong><br />
birds with the general collection <strong>of</strong> the <strong>American</strong> <strong>Museum</strong> <strong>of</strong><br />
<strong>Natural</strong> <strong>History</strong>, an attempt was made to arrive at a natural<br />
arrangement for each family or other unit. This <strong>of</strong>ten led to<br />
rather detailed studies or to intensive efforts to determine the<br />
correct position <strong>of</strong> difficult genera. A number <strong>of</strong> publications<br />
growing from these studies are included in the bibliography (see<br />
titles by Amadon, Chapin, Delacour, Mayr, Vaurie, and Zimmer).<br />
They relate primarily to Old World families not yet included<br />
in Peters' "Check-list" for which no authoritative list<br />
exists comparable to Hellmayr's for the New World.<br />
The principal purpose <strong>of</strong> this paper is to give these findings more<br />
general expression. We have <strong>of</strong> course incorporated the work <strong>of</strong><br />
others whenever known to us and have included the non-passerine<br />
groups, although few changes are made from the now wellestablished<br />
sequence <strong>of</strong> Wetmore (1934, followed by Peters).<br />
Indeed we have throughout attempted to make no changes from<br />
the established sequence except when they are clearly indicated<br />
by recent evidence. Occasion is taken to give a corrected count<br />
<strong>of</strong> species in each family <strong>of</strong> birds; such a count proved a useful<br />
feature <strong>of</strong> a previous paper by the senior author (Mayr, 1946).<br />
As a result <strong>of</strong> various discoveries and recent revisions the total<br />
number <strong>of</strong> species in the present list is 8590 as compared with<br />
8616 in the previous one. The change within five years amounts<br />
to less than one-half <strong>of</strong> one per cent. Because <strong>of</strong> the large number<br />
<strong>of</strong> insular forms <strong>of</strong> doubtful status, the number <strong>of</strong> species <strong>of</strong> birds<br />
will always remain an estimate. The final figure may vary by<br />
several hundreds either way, depending on the point <strong>of</strong> view <strong>of</strong> the<br />
enumerator. The five "species" <strong>of</strong> Todus or the three <strong>of</strong> Ryn-
2 AME3RICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
chops, for example, might be considered races just as have the<br />
former "species" <strong>of</strong> Anhinga. Further study <strong>of</strong> continental<br />
forms, on the other hand, <strong>of</strong>ten gives clear-cut answers as to the<br />
racial or specific status <strong>of</strong> forms previously <strong>of</strong> dubious status.<br />
The result <strong>of</strong> the two recent counts indicates, however, that the<br />
final figure will be within 2 per cent <strong>of</strong> 8600. For all practical<br />
purposes this figure will be satisfactory as a very close approach<br />
to the actual number <strong>of</strong> species <strong>of</strong> living birds.<br />
We would like to express our appreciation to Mr. James L.<br />
Peters for providing revised figures for some <strong>of</strong> the families <strong>of</strong><br />
South <strong>American</strong> mesomyodian song birds and to Dr. R. C.<br />
Murphy for up-to-date figures <strong>of</strong> the number <strong>of</strong> species in some <strong>of</strong><br />
the sea bird families. We have pr<strong>of</strong>ited greatly from a painstaking<br />
examination <strong>of</strong> our manuscript by Dr. J. Van Tyne.<br />
Two important aims <strong>of</strong> any classification are to keep related<br />
groups as close together as possible and to put ancestral groups<br />
first, derived groups later. In a list that does not include fossil<br />
forms, the second <strong>of</strong> these principles can have only limited expression.<br />
Specialization is <strong>of</strong>ten a clue to relationship and phylogenetic<br />
sequence, but de-specialization may occur and much<br />
phylogenetically meaningless specialization exists. Cotingas, for<br />
example, appear more specialized than most <strong>of</strong> the Oscines, but<br />
the structure <strong>of</strong> the syrinx and tarsus, as well as other evidence,<br />
indicates that the Oscines are a later, more advanced group than<br />
the cotingas.<br />
There is no particular difficulty in incorporating the above aims<br />
in a classification so long as a single, non-branching sequence is<br />
involved. Difficulties arise in a branching system, for it is impossible<br />
to pursue all avenues <strong>of</strong> descent simultaneously in a<br />
linear sequence <strong>of</strong> names. Inevitably one must follow each<br />
branch in succession to its most recent end twig and then go back<br />
to the main stem and repeat the process.<br />
The problem <strong>of</strong> the relationship <strong>of</strong> the avian orders is an old one<br />
and one that will probably never be solved satisfactorily. One<br />
point that is frequently overlooked is that all living birds are exceedingly<br />
specialized in different directions and that this specialization<br />
had its beginning in the remote past. Perhaps the most<br />
specialized <strong>of</strong> known birds is Hesperornis, which lived in the Cretaceous.<br />
The Eocene Diatrymca is perhaps more specialized than<br />
any bird that lives today. The connections between the living<br />
orders are lost in antiquity, and their analysis is further obscured<br />
by much convergent evolution <strong>of</strong> habitus type.
1951 A CLASSIFICATION OF RECENT BIRDS 3<br />
Predators, for example, evolve a hooked bill and sharp claws.<br />
It is now generally accepted that hawks and owls are not closely<br />
related, but Hudson (1948) supplies evidence that even the hawks<br />
may be a polyphyletic group. Hesperornis (Hesperornithes), the<br />
loons (Gaviae), and the grebes (Podicipedes) are <strong>of</strong>ten cited as<br />
forming a group <strong>of</strong> three related orders in view <strong>of</strong> the structure <strong>of</strong><br />
their rear limbs, but as Stolpe (1935) has shown, the anatomical<br />
similarities that have been cited as evidence for relationship are<br />
actually functional convergences. A better known example <strong>of</strong><br />
convergence is that <strong>of</strong> the "Ratites," a group which consists <strong>of</strong> at<br />
least five unrelated groups <strong>of</strong> birds, which have become flightless<br />
secondarily and attained large size. Many other cases <strong>of</strong> convergent<br />
evolution have been unmasked recently among the passerine<br />
birds, e.g., the "shrikes," "titmice," "finches," "warblers,"<br />
and others (see below). In view <strong>of</strong> the frequency <strong>of</strong> convergence<br />
among birds, it seems mandatory that some <strong>of</strong> the other heterogeneous-appearing<br />
orders <strong>of</strong> birds (e.g., the Gressores, Grues, and<br />
Laro-Limicolae) be reexamined to evaluate the authenticity <strong>of</strong><br />
relationship <strong>of</strong> the included families and suborders. Some <strong>of</strong><br />
them (e.g., the Thinocoridae, Turnicidae, and Mesoenatidae)<br />
have, <strong>of</strong> course, long been a puzzle to taxonomists.<br />
The other extreme is provided by such groups as the Sphenisci<br />
and Tubinares, which appear exceedingly different but are clearly<br />
related, as first pointed out by Furbringer and later confirmed by<br />
Simpson (1946). The relationship <strong>of</strong> the kiwis to the moas, and,<br />
if confirmed, the relationship <strong>of</strong> the turacos to the gallinaceous<br />
birds would also be illustrations <strong>of</strong> highly dissimilar but related<br />
groups.<br />
The existing orders <strong>of</strong> birds are thus the terminal twigs <strong>of</strong> an<br />
exceedingly ramified phylogenetic tree, and it is rather immaterial<br />
in what exact sequence many <strong>of</strong> the orders are listed. Portmann<br />
(1938) is convinced that the gallinaceous birds are the most primitive<br />
<strong>of</strong> living birds. Stresemann (1927-1934) also placed them<br />
near the beginning, yet Wetmore lists them as the nineteenth<br />
order. In view <strong>of</strong> our scanty knowledge <strong>of</strong> relationships and the<br />
uneven rate <strong>of</strong> specializing trends, it is <strong>of</strong>ten impossible to decide<br />
which <strong>of</strong> two orders is the more primitive or ancestral.<br />
The use <strong>of</strong> the ending "-formes" for ordinal names has met<br />
with considerable opposition among zoologists who are reluctant<br />
to see such a venerable name as, for example, "Lepidoptera" replaced<br />
by "Papilioniformes." We have followed Stresemann in
4 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
using the older ordinal names, but when those used by Wetmore<br />
have a different root they are given in parentheses.<br />
RATITES<br />
The problem <strong>of</strong> ratite phylogeny continues to receive much<br />
attention. The present consensus is that the main groups <strong>of</strong> these<br />
birds are <strong>of</strong> independent origin. Stresemann's suggestion that<br />
the kiwis (Apterygidae) be placed in the same order as the moas<br />
(Dinornithidae) is followed. McDowell (1948) may well be correct<br />
in believing the rheas to be allied to the tinamous, but we<br />
doubt whether they should be placed in the same order, pending<br />
further study.<br />
In the literature one finds records <strong>of</strong> moas and kiwis from<br />
Australia and <strong>of</strong> elephant birds from Africa. In our opinion all <strong>of</strong><br />
these records are based on such fragmentary material as to be<br />
subject to doubt. Oliver (1949) has recently written a volume<br />
called "The moas <strong>of</strong> New Zealand and Australia." The supposed<br />
Australian moa, "Dinornis" queenslandiae DeVis, is based on a<br />
femur head from "Post Tertiary" deposits. Two other giant<br />
extinct ratites, Dromornis and Genyornis, are known from the<br />
Pleistocene <strong>of</strong> Australia. Both are assigned to the Dromaeidae.<br />
Careful comparison <strong>of</strong> the published figures <strong>of</strong> the femur <strong>of</strong><br />
"Dinornis" queenslandiae, made by McDowell at the suggestion<br />
<strong>of</strong> the authors, led him to the conclusion that the development <strong>of</strong><br />
the trochanter is so different that it seems impossible that queenslandiae<br />
belongs in the Dinornithidae. There is no reason to<br />
suppose it could not represent a distinct ratite type allied to the<br />
emus rather than to the moas. According to McDowell (verbal<br />
communication) the supposed records <strong>of</strong> kiwis from Australia<br />
(Metapteryx) are also erroneous.<br />
The presence <strong>of</strong> relatives <strong>of</strong> Aepyornis in fossil remains from<br />
Africa is based on even less satisfactory fragments and should not<br />
be accepted without much more evidence. We doubt if this will<br />
ever be forthcoming.<br />
It is a curious fact that the male assumes all the duties <strong>of</strong> incubation<br />
in all the living ratites except sometimes in the ostrich<br />
but including the tinamous. Probably this is only a remarkable<br />
coincidence.<br />
SPHENISCI; TUBINARES (PROCELLARIIFORMES)<br />
As already noted, the penguins are related to the petrels and
1951 A CLASSIFICATION OF RECENT EIRDS .5<br />
less closely to the Steganopodes. Simpson (1946) has shown that<br />
the Miocene penguins were osteologically much like those <strong>of</strong> today.<br />
The metatarsals are less fused in the modern forms, an interesting<br />
example <strong>of</strong> "reversal <strong>of</strong> evolution" and "de-specialization."<br />
LOONS: GAVIAE; GREBES: PODICIPEDES (COLYMBIFORMES)<br />
The controversy as to whether the name Colymbus Linnaeus,<br />
1758, applies to a loon or to a grebe is perhaps best resolved, as<br />
suggested in a paper by Dr. F. Salomonsen presented at the Xth<br />
International Ornithological Congress, by declaring the name<br />
indeterminable. The names Gavia will then be used for the loons<br />
and Podiceps for the typical grebes.<br />
As already indicated, Stolpe (1935) concluded that there is little<br />
reason to believe that the loons and grebes are related to the Cretaceous<br />
Hesperornis (cf. Howard, 1950) or, for that matter, to each<br />
other. Since, however, the grebes have been thought to be remote<br />
allies <strong>of</strong> the petrels, and since McDowell (oral communication)<br />
thinks that the loons may be a specialized <strong>of</strong>fshoot <strong>of</strong> petrel stock,<br />
it is possible that the grebes and loons have some distant or indirect<br />
relationship. It is permissible, therefore, to continue to keep<br />
them near each other in a classification.<br />
STEGANOPODES (PELECANIFORMES)<br />
Lanham (1947) has assembled anatomical data to show that the<br />
tropic birds (Phaithontidae) and frigate birds (Fregatidae) share<br />
a number <strong>of</strong> apparently primitive characters with the Tubinares.<br />
These characters are <strong>of</strong>ten lacking in the third main group <strong>of</strong> the<br />
Steganopodes, which includes the pelicans, cormorants, anhingas,<br />
and gannets. Phaethon is certainly further removed from Pelecanus<br />
and allies than is Fregata.<br />
The resemblances between the tropic birds and the gulls and<br />
terns have <strong>of</strong>ten been pointed out, but until evidence <strong>of</strong> true<br />
affinity is brought forward they must be attributed to convergence.<br />
G. Timmermann (MS) has found certain <strong>of</strong> the Mallophaga <strong>of</strong><br />
these two groups to be related, but it is possible that this is a result<br />
<strong>of</strong> secondary transfer <strong>of</strong> parasites in birds <strong>of</strong> similar size and<br />
habits. The case may be comparable to the occurrence on the skua<br />
(Catharacta) <strong>of</strong> an "endemic" species <strong>of</strong> Mallophaga <strong>of</strong> a genus<br />
otherwise restricted to the Tubinares (Hopkins, 1942, p. 100).<br />
Anhinga is so much like Phalacrocorax that it would seem to
6 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
require no more than subfamily status. Stresemann did not give<br />
it even that.<br />
ACCIPITRES OR FALCONES<br />
The diurnal birds <strong>of</strong> prey are a highly differentiated group,<br />
thought by some to be related to the Gressores, and more distantly<br />
to the Steganopodes. Hudson (1948) and others have<br />
written on the classification <strong>of</strong> this order in recent years. The<br />
<strong>American</strong> vultures (Cathartidae) are very distinct and may not be<br />
related to the other Falcones. Perhaps they are representatives<br />
<strong>of</strong> some ancient <strong>American</strong> radiation which may even include<br />
some or all <strong>of</strong> such families as the Anhimidae, Cracidae, and Tinamidae<br />
(McDowell, verbal communication). The occurrence <strong>of</strong> a<br />
cathartid, Pleseocathartes, in the upper Eocene <strong>of</strong> France requires<br />
confirmation, in the opinion <strong>of</strong> Dr. A. Wetmore who has examined<br />
the specimens (verbal communication).<br />
The African secretary bird, Sagittarius, resembles the gruiform<br />
Cariamidae <strong>of</strong> South America. Some would transfer the Sagittariidae<br />
to the Grues, others the Cariamidae to the Accipitres.<br />
Actually, the resemblance may be parallelism. Still, Sagittarius<br />
may not belong to the Accipitres.<br />
Certain similarities in the pterylography and in the plantar<br />
tendons <strong>of</strong> the Pandionidae and Cathartidae (Compton, 1938)<br />
are apparently <strong>of</strong> no phylogenetic significance (Hudson, 1948).<br />
GRESSORES (CICONIIFORMES)<br />
The whale-head or shoe-bill stork, Balaeniceps, has <strong>of</strong>ten been<br />
placed in a monotypic family. Bohm (1930) thought its anatomy<br />
like that <strong>of</strong> a true stork, and Dr. James P. Chapin tells us that he<br />
believes it well placed in the Ciconiidae. It has the bill-rattling<br />
habit <strong>of</strong> storks. The hammerhead, Scopus, on the other hand, is<br />
more distinct and may be left in a separate family. Both genera<br />
share some characters with the Ardeidae.<br />
The boat-billed heron, Cochlearius, appears to be a typical<br />
heron in all except its curious bill. It is even possible to point<br />
to the night herons as being probably its nearest relatives. We<br />
think this genus may properly be placed in the Ardeidae, just as<br />
the spoonbills, Platalea, are placed in the Threskiornithidae.<br />
The trematodes <strong>of</strong> the Ciconiidae and Ardeidae are quite different<br />
(Szidat, 1942), but possibly study <strong>of</strong> those <strong>of</strong> some <strong>of</strong> the<br />
other families <strong>of</strong> Gressores will bridge this gap.
1951 A CLASSIFICATION OF RECENT BIRDS 7<br />
PHOENICOPTERI<br />
Flamingos resemble the Anseres in some ways, though many <strong>of</strong><br />
the characters involved, particularly the bill, are highly adaptive.<br />
The Mallophaga <strong>of</strong> the two groups are much alike (Hopkins, 1949),<br />
while those <strong>of</strong> the Phoenicopteri and Gressores (with which the<br />
flamingos are usually placed) are very different.<br />
The flamingos have, however, many anatomical similarities<br />
with storks and ibises. Stresemann, after enumerating these<br />
similarities, stated that the resemblance <strong>of</strong> the flamingos to the<br />
waterfowl is superficial. In view <strong>of</strong> this conflicting evidence it<br />
seems best to place the flamingos as a separate order between the<br />
Anseres and Gressores. They may be related to both.<br />
ANSERES<br />
The waterfowl are a very distinct group but may be placed between<br />
the flamingos and gallinaceous birds. The two South<br />
<strong>American</strong> families Anhimidae (Anseres) and Cracidae (Galli)<br />
may be distantly related (Delacour, 1949). Delacour and Mayr<br />
(1945) wrote on the classification <strong>of</strong> the Anatidae, which they<br />
divided into two subfamilies and five tribes. Several genera are<br />
<strong>of</strong> doubtful allocation, and one in particular, the magpie-goose,<br />
Anseranas, deserves to be placed in a sixth trbe (von Boetticher,<br />
1943).<br />
GALLI<br />
The gallinaceous birds are one <strong>of</strong> the primitive and basic orders.<br />
It is highly unlikely that they are related to the Tinami. We follow<br />
Delacour (1951) in considering the grouse and guineafowls<br />
as only subfamilies <strong>of</strong> the Phasianidae.<br />
Stresemann placed the hoatzin, Opisthocomus, in a separate<br />
order, which, however, he considered to be "einen sehr nahen<br />
Verwandten der Galli." Some <strong>of</strong> the peculiarities <strong>of</strong> this bird,<br />
such as the large crop and the well-developed wing claws <strong>of</strong> the<br />
chick, are specializations for its peculiar diet and habitat and may<br />
not be primitive. Pycraft thought Opisthocomus to have significant<br />
resemblances to the genus Centropus (Cuculidae) and to the<br />
turacos (Musophagidae), as well as to the Galli. Stresemann<br />
considered the similarity to cuckoos and turacos as superficial.<br />
For the time being, it seems best to retain the Opisthocomidae<br />
in the Galli as a suborder. In the opinion <strong>of</strong> McDowell the hoatzin<br />
has a large number <strong>of</strong> important resemblances to the Cracidae<br />
and Anhimidae.
8 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
CUCULI<br />
There is a difference <strong>of</strong> opinion, first as to whether or not the<br />
turacos (Musophagidae) should be associated with the Galli, and<br />
second whether or not the cuckoos (Cuculidae) are related to the<br />
turacos. The Mallophaga <strong>of</strong> the turacos are said to be strongly<br />
indicative <strong>of</strong> relationship to the Galli, though Hopkins (1949)<br />
thinks this might be the result <strong>of</strong> accidental transfer in a dust<br />
bath. Pycraft and Furbringer found morphological resemblances<br />
between turacos and the Galli, though Stresemann and others<br />
considered these superficial. We think, however, it is best to<br />
place the turacos tentatively near the Galli.<br />
The turacos and cuckoos agree in most features <strong>of</strong> basic anatomy.<br />
Moreover, the cuckoos <strong>of</strong> the genus Coua <strong>of</strong> Madagafcar<br />
<strong>of</strong>ten resemble turacos in body form, presence <strong>of</strong> bare areas on<br />
face, coloration including even purplish or reddish tinge on tail,<br />
and general configuration <strong>of</strong> bill and nostrils. While they lack<br />
the full crest <strong>of</strong> the turacos, there is an incipient one. Turacos<br />
always perch with two toes back and two forward, though in<br />
death this functional zygodactyly is not always indicated by the<br />
position <strong>of</strong> the toes. Moreau (1938) has emphasized developmental<br />
differences between turacos and cuckoos, but the tendency<br />
towards peculiar nesting habits in the latter lessens the significance<br />
<strong>of</strong> this. It is entirely possible that the Musophagidae are somewhat<br />
primitive relatives <strong>of</strong> the Cuculidae, so we tentatively follow<br />
convention in associating the two families in the same order.<br />
GRUES<br />
This order has no obvious ties with the orders preceding it,<br />
though various links with the Falcones, Galli, or even the Gressores<br />
have been advanced at one time or another. The large number <strong>of</strong><br />
relict families <strong>of</strong> a genus or two each in the Grues marks it as a<br />
declining and ancient order, as does its fossil history. The phorlioracids<br />
<strong>of</strong> South America and possibly the Eocene Diatryma are<br />
among the extinct relatives <strong>of</strong> the Grues.<br />
LARO-LIMICOLAE (CHARADRIIFORMES)<br />
This diversified order may be connected with the Grues through<br />
one or all <strong>of</strong> the Burhinidae, Jacanidae, and Thinocoridae.<br />
Several <strong>of</strong> the shore bird families currently recognized seem to<br />
require no more than subfamily status (cf. Stresemann).
.1951 A CLASSIFICATION OF RECENT BIRDS 9<br />
COLUMBAE; PSITTACI<br />
The Columbae may be tentatively placed near the Laro-Limicolae,<br />
but certain resemblances between pigeons and game birds<br />
may eventually prove to be <strong>of</strong> significance. It is unlikely that<br />
the sand grouse (Pteroclidae) are grouse-like except in superficial<br />
adaptations.<br />
The Psittaci are a strongly differentiated group. Resemblance<br />
to the Accipitres is probably mere convergence, and relationship<br />
to the Cuculi, championed by Gadow, must be rather distant at<br />
best. McDowell has found similarities between the humeri <strong>of</strong><br />
parrots and those <strong>of</strong> pigeons.<br />
CAPRIMULGI; STRIGES<br />
Hudson (1937) found a. great difference between the thigh<br />
muscles <strong>of</strong> owls and those <strong>of</strong> Caprimulgidae, but perhaps examination<br />
<strong>of</strong> other families, such as the Aegothelidae, would bridge the<br />
gap. Certainly there are strong reasons to believe the two orders<br />
to be related, but the resemblance <strong>of</strong> goatsuckers to swifts is<br />
evidently superficial.<br />
CORACIAE<br />
The Madagascar cuckoo roller, Leptosomus, and the ground<br />
rollers have been placed in the same family by Sclater (1924-<br />
1930) and by Wetmore. Yet the ground rollers appear to be<br />
much closer to the true rollers than to Leptosomus, though some<br />
may prefer to recognize three families.<br />
MACROCHIRES (APODIFORMES); COLII<br />
Whether the swifts and hummingbirds are actually related<br />
to each other or not is an open question, but Furbringer found<br />
much evidence that they are. He considered both to be rather<br />
closely allied to the Passeres. Since, however, the Pici seem to be<br />
even closer to the Passeres, the sequence Macrochires, Pici, Passeres<br />
is suggested. The very isolated Colii are closest to the<br />
Macrochires, according to Furbringer, although there is some<br />
indication <strong>of</strong> relationship with the Coraciae, particularly the<br />
Alcedinidae.<br />
PERCHING BIRDS: PASSERES<br />
The subdivision <strong>of</strong> the Passeres on the basis <strong>of</strong> the muscles <strong>of</strong><br />
the syrinx was made by Johannes Muller as early as 1847. About
10 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
1880 Garrod and Forbes published several papers establishing the<br />
position <strong>of</strong> such families as the broadbills and philepittas. These<br />
and other important references on the Order Passeriformes are<br />
given by Ridgway (1901, pp. 2-23; 1907, pp. 328-332) and by<br />
Stresemann (1927-1934, pp. 843-850). The following brief summary<br />
mentions only the more important or controversial points.<br />
SUBOSCINES<br />
The broadbills differ from other Passeres and agree with some<br />
non-Passeres by having a vinculum or band connecting the deep<br />
muscles <strong>of</strong> the toes. They have 15 cervical vertebrae as compared<br />
to 14 in most other Passeres, as well as other distinctions.<br />
It thus seems justifiable to follow those authorities who place the<br />
Eurylaimidae in a separate suborder. Their resemblances to the<br />
Pittidae or even the Cotingidae have led some to give them lesser<br />
rank.<br />
The remaining Passeres may be divided into two groups, as<br />
follows:<br />
A. Mesomyodes or Clamatores: Those in which the intrinsic<br />
muscles (if any) <strong>of</strong> the syrinx are attached to one <strong>of</strong> the ends or to<br />
the middle <strong>of</strong> the bronchial half rings (the Eurylaimidae agree with<br />
this group).<br />
B. Acromyodes: Those in which these muscles are attached to<br />
both ends <strong>of</strong> the bronchial half rings.<br />
Group A, Mesomyodes or Clamatores, is further divisible into<br />
two main groups: (1) those in which the syrinx is tracheobronchial<br />
(Haploophonae), and (2) those in which it is entirely<br />
tracheal (Tracheophonae). Group 1 contains both Old World<br />
and New World families. The former are the Pittidae (Asia,<br />
Australo-Papua, and Africa), Philepittidae (Madagascar), and<br />
Xenicidael (New Zealand). Recent study has shown that the<br />
genus Neodrepanis <strong>of</strong> Madagascar is a philepittid (Amadon, in<br />
press). These three families, through not very closely related,<br />
are very likely more nearly allied to one another than to the<br />
Neotropical members <strong>of</strong> this group.<br />
The remaining families <strong>of</strong> the "Haploophonae" are exclusively<br />
or predominantly (Tyrannidae) Neotropical, viz., Tyrannidae (in<br />
which we include Oxyruncus as a subfamily), Pipridae, Cotingidae,<br />
1 Since the name Xenicidae has prior usage, as for example in the "Catalogue <strong>of</strong><br />
birds" <strong>of</strong> the British <strong>Museum</strong>, it is unnecessary to replace it by the more cumbersome<br />
Acanthisittidae merely because the latter is based on an older generic name.
1951 A CLASSIFICATION OF RECENT BIRDS 11<br />
and Phytotomidae. The Pipridae and Cotingidae differ from<br />
other passerines in that the femoral, not the ischiatic, artery is<br />
the principal one <strong>of</strong> the thigh. This is probably a functional<br />
character reflecting the relative development <strong>of</strong> the thigh muscles.<br />
Hence we do not think it necessary to make a separate family for<br />
the cock-<strong>of</strong>-the-rock, Rup'icola, because it has the ischiatic artery<br />
predominant. This aberrancy may be correlated with its terrestrial<br />
dances.<br />
Garrod united the manakins and cotingas in a single family.<br />
Further study may show that this procedure, with subfamily rank<br />
for the two groups, is justified. Of the genera currently assigned<br />
to the Pipridae, Schiffornis appears somewhat like a cotinga.<br />
Calyptura <strong>of</strong> the Cotingidae has been placed by some authors in<br />
the Pipridae. Some <strong>of</strong> the genera <strong>of</strong> both <strong>of</strong> these families are<br />
very much like some genera <strong>of</strong> the Tyrannidae. This is true <strong>of</strong><br />
Tyranneutes and Sapayoa <strong>of</strong> the Pipridae. Xenopsaris was placed<br />
in the Cotingidae by Ridgway, in the Tyrannidae by Hellmayr.<br />
The Cotingidae recall in some ways the Eurylaimidae, even to<br />
the construction <strong>of</strong> large, bag-like nests by the broadbills and by<br />
some cotingas. Pycraft was especially impressed by these similarities,<br />
and he thought the two groups might some day be reduced<br />
to subfamilies That the relationship is this close seems more than<br />
doubtful, but further study is needed. The peculiar genus Phytotomca<br />
(plant-cutters) somewhat resembles the cotingas, though the<br />
ischiatic is the principal artery <strong>of</strong> the thigh. It must be retained<br />
as a monotypic family pending further study.<br />
The second division <strong>of</strong> group A is peculiar in that the syrinx is<br />
entirely tracheal. There are four families <strong>of</strong> "Tracheophonae,"<br />
all Neotropical, the Rhinocryptidae, Conopophagidae, Formicariidae,<br />
and Dendrocolaptidae (including "Furnariidae").<br />
The usual subdivision <strong>of</strong> the Tracheophonae into these four<br />
families is not particularly satisfactory.. We follow the earlier<br />
authors who left the ovenbirds (Furnarius and allies) with the<br />
closely related woodhewers (Dendrocolaptes and allies), but the<br />
two groups may be retained as separate subfamilies. This family,<br />
Dendrocolaptidae, differs from the remaining tracheophones by<br />
having two rather than one pair <strong>of</strong> tracheo-bronchial muscles.<br />
The group with one pair <strong>of</strong> tracheo-bronchial muscles is usually<br />
divided into a large family, the antbirds (Formicariidae), supposedly<br />
characterized by a two-notched metasternum, and two<br />
smaller families (Rhinocryptidae, Conopophagidae) in which the
12 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496-i<br />
metasternum is four-notched. Dr. J. T. Zimmer has called to our<br />
attention that the late Waldron Miller (unpublished) found that<br />
the genus Melanopareia, comprising otherwise typical formicarians,<br />
has a four-notched sternum. It is possible that further<br />
study will show that the Rhinocryptidae cannot be separated<br />
satisfactorily from the Formicariidae. The former may prove to:<br />
be a polyphyletic group composed <strong>of</strong> specialized ant birds. The<br />
genus Conopophaga lacks intrinsic syrinx muscles altogether and<br />
has an exaspidean tarsal envelope. Whether these characters are<br />
constant enough to warrant full family status for this genus or<br />
not must remain with the future.<br />
The Xenicidae <strong>of</strong> New Zealand resemble some <strong>of</strong> the Tracheophonae,<br />
particularly the genus Conopophaga. Both are longlegged,<br />
short-tailed denizens <strong>of</strong> the undergrowth, and both<br />
happen to have a white supraocular stripe. We do not think the<br />
parallelism is any more indicative <strong>of</strong> close relationship than, for<br />
example, that between such formicariids as Graliaria and such<br />
thrushes as Amalocichla or Zeledonia. The similarity <strong>of</strong> the<br />
Australian scrub bird, Atrichornis, to some <strong>of</strong> the Rhinocryptidae,<br />
e.g., Liosceles or Scytalopus, is, to our minds, equally a matter <strong>of</strong><br />
adaptation to similar habitats.<br />
In regard to group B above (those passeres in which the syrinx<br />
muscles are attached to both ends <strong>of</strong> the bronchial half rings),<br />
it contains, in addition to the true song birds, two primitive<br />
Australian families, the Atrichornithidae and Menuridae. Each<br />
<strong>of</strong> the latter contains but a single genus. Although very dissimilar<br />
in appearance, they share a number <strong>of</strong> anatomical peculiarities<br />
and are probably nearer to each other than either is to<br />
any other group. Although the attachment <strong>of</strong> the syrinx muscles<br />
in these two Australian families is similar to that <strong>of</strong> the Oscines,<br />
only two or three pairs <strong>of</strong> such muscles are present, as compared to<br />
five to seven pairs in the Oscines proper.<br />
Probably further study <strong>of</strong> some <strong>of</strong> the larger <strong>of</strong> the groups mentioned<br />
above, e.g., the tyrant flycatchers, will reveal that they<br />
should be divided into subfamilies or tribes equivalent to those<br />
used below for some <strong>of</strong> the families <strong>of</strong> song birds.<br />
The various suboscine perching birds give every appearance <strong>of</strong><br />
being in the process <strong>of</strong> replacement by the Oscines. Many <strong>of</strong> the<br />
families <strong>of</strong> the former group have a relict distribution in Madagascar,<br />
New Zealand, Australia, or South America. The last<br />
continent was <strong>of</strong> course isolated for a long time. It has few well-
1951 A CLASSIFICATION OF RECENT BIRDS 13'<br />
differentiated families <strong>of</strong> indigenous song birds, although the subfamilies<br />
<strong>of</strong> the tanager-pyrrhuloxine finch assemblage are richly<br />
developed. An adaptive radiation <strong>of</strong> the wrens, mimids, and<br />
other song birds <strong>of</strong> South America might reduce the Suboscines <strong>of</strong><br />
that continent to the subordinate position they occupy in the Old<br />
World.<br />
OSCINES OR TRUE SONG BIRDS<br />
Of the approximately 8600 species <strong>of</strong> living birds, 5100 belong to<br />
the Passeres, and about 4000, to the Suborder Oscines. The difficulty<br />
in finding good anatomical characters that besets the avian<br />
taxonomist at every level is particularly acute in this suborder.<br />
As' a group song birds are the most advanced, successful, and apparently<br />
latest to evolve <strong>of</strong> the entire Class Aves. They have developed<br />
an infinite variety <strong>of</strong> types, and many annectant aand in-,<br />
termediate forms are still in existence. The "phylogenetic tree"<br />
<strong>of</strong> -the group, if it could be drawn accurately, would probably<br />
resemble a great flat-topped "umbrella" tree.<br />
Added to the above difficulties is a large amount <strong>of</strong> parallelism,<br />
<strong>of</strong>ten in different continents. For example, the <strong>American</strong> orioles<br />
and' blackbirds (Icteridae) are the ecological homologues <strong>of</strong> the<br />
Old World starlings (Sturnidae). So slight are the anatomical<br />
distinctions in this suborder that these two groups were once'<br />
placed in the same family by competent anatomists, though they<br />
are probably quite unrelated.<br />
The object <strong>of</strong> the following discussion <strong>of</strong> the families <strong>of</strong> Oscines<br />
is not to present entirely novel findings. Rather it is to summarize<br />
the results <strong>of</strong> investigations <strong>of</strong> numerous workers and particularly<br />
the results <strong>of</strong> field workers who have <strong>of</strong>ten produced better clues<br />
to relationship than the anatomists. Furthermore, it is intended<br />
to expose our abysmal ignorance <strong>of</strong> the relationships <strong>of</strong> most songbird<br />
families. In spite <strong>of</strong> their basic uniformity, there are certain<br />
minor anatomical differences as pointed out by Stonor (1937) and<br />
Beecher (MS). It would seem high time to investigate previously<br />
unexplored anatomical systems (as Beecher has for the jawmuscles)<br />
in an attempt to shed new light on oscinine classification.<br />
The shape <strong>of</strong> the bill has been used with too much confidence<br />
in the past as a reliable basis for classification. Actually the bill<br />
is the most plastic <strong>of</strong> all the organs <strong>of</strong> a bird, and evidence is<br />
mounting daily that shrike-like bills, warbler-like bills, finch-like<br />
bills, and flycatcher-like bills have been acquired repeatedly and
14 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
polyphyletically by various unrelated groups <strong>of</strong> Oscines. The investigation<br />
<strong>of</strong> the true relationship <strong>of</strong> these groups will be a rewarding<br />
task.<br />
Habits are <strong>of</strong>ten a better clue to relationships than structure.<br />
Nest-building habits, for instance, are diagnostic for the subfamilies<br />
<strong>of</strong> Paridae, and similarity in nests was a clue to Mayr and<br />
Bond (1943) in allying Ptyonoprogne with Hirundo rather than<br />
with Riparia.<br />
Only two or three oscinine families are reasonably well defined.<br />
The larks (Alaudidae) differ from the others by having the back <strong>of</strong><br />
the tarsus rounded and broken up into plates. They also lack the<br />
bony pessulus at the junction <strong>of</strong> the bronchi in the syrinx. The<br />
Hirundinidae have the bronchial rings more or less complete; in<br />
other Oscines they are "half rings," with a membrane across the<br />
inner face. But, from here on, consistent family characters are extremely<br />
hard to find.<br />
These difficulties have led some anatomists to divide the Oscines<br />
into only three or four families. This serves to keep passerine<br />
families equivalent to those <strong>of</strong> other orders, but does not help in<br />
classifying the many obviously monophyletic groups <strong>of</strong> song birds<br />
that must be designated somehow, whether we call them families,<br />
subfamilies, tribes, or what not. Certain groups clearly connected<br />
by linking forms may be reduced to the level <strong>of</strong> subfamilies as has<br />
been done for the thrush-babbler-flycatcher-warbler assemblage.<br />
This will, however, inevitably lead to the use <strong>of</strong> "tribes" or other<br />
intermediate categories, as in Delacour's (1946) revision <strong>of</strong> the<br />
Timaliinae. For this reason we have retained, for the time being,<br />
family status for a number <strong>of</strong> groups which it may prove expedient<br />
to reduce to subfamilies later.<br />
The arrangement <strong>of</strong> the passerine families in a linear sequence<br />
is a task involving many compromises. At best, one can seek<br />
only to keep related families together. The classification <strong>of</strong><br />
Stejneger, followed in considerable measure by that <strong>of</strong> Wetmore,<br />
places much weight on the relative reduction <strong>of</strong> the tenth (outer)<br />
primary. This trend towards specialization is <strong>of</strong> some importance,<br />
especially since all, or almost all, <strong>of</strong> the Suboscines have a long<br />
tenth primary, longer than in any oscinine. On the other hand,<br />
reduction <strong>of</strong> the tenth primary is certainly subject to reversal and<br />
is a highly adaptive, plastic, and polyphyletic character. For<br />
example, the genus Urocynchramus <strong>of</strong> the fringilline finches has a<br />
relatively long tenth primary, though all other finches lack one.
1951 A CLASSIFICATION OF RECENT BIRDS 15<br />
Other families in which the tenth primary may be present or absent<br />
from genus to genus are the Bombycillidae (sensu lato), Dicaeidae,<br />
and Sturnidae. The seemingly primitive Alaudidae have<br />
the tenth primary quite reduced or even vestigial in a few genera.<br />
Some highly specialized families, such as the Nectariniidae, have<br />
a moderately developed tenth primary. It certainly does not follow<br />
that the nine-primaried families are necessarily the "highest"<br />
song birds. Other characters denoting specialization, such as a<br />
booted condition <strong>of</strong> the tarsus, are also <strong>of</strong> variable significance.<br />
A number <strong>of</strong> authorities, including W. K. Parker, Macgillivray,<br />
Sharpe, and Alfred Newton (1893-1896, pp. 117-120), thought<br />
that the Corvidae are, all in all, the most advanced and "highest"<br />
type <strong>of</strong> bird. Studies <strong>of</strong> the avian brain by Portmann (1947)<br />
have tended to show that Corvus ranks above other birds in brain<br />
development. As Dr. J. P. Chapin has pointed out to us, however,<br />
were similar attention given to certain icterids, sturnids,<br />
ploceids, and perhaps others, they might prove just as "advanced."<br />
Nevertheless, we have thought it best to follow the school that<br />
would terminate the oscinine series with the Corvidae and allied<br />
families. It is the sequence adopted in Sharpe's "Handlist"<br />
and in many ornithological reference works throughout the world.<br />
It will be all too evident from this discussion that no linear<br />
sequence for the Oscines can be adequate and that various lines <strong>of</strong><br />
specialization must <strong>of</strong>ten be followed to their conclusion and<br />
then a fresh start be made.<br />
Unlike the Suboscines, which have their richest present representation<br />
in South America, the Oscines have flourished and differentiated<br />
chiefly in the Old World tropics and Australia. In these<br />
areas literally dozens <strong>of</strong> genera present such a problem that they<br />
have been shifted from family to family or assigned to "scrap<br />
baskets" such as the Prionopidae, Timaliidae, etc. We hope that<br />
the following discussion <strong>of</strong> these difficult groups, <strong>of</strong>ten with a<br />
listing <strong>of</strong> the genera involved, will prove useful.<br />
ALAUDIDAE<br />
The larks are perhaps best left at the beginning <strong>of</strong> the oscinine<br />
series because <strong>of</strong> the reticulate posterior surface <strong>of</strong> the tarsus and<br />
the absence <strong>of</strong> an ossified pessulus in the syrinx.<br />
HIRUNDINIDAE<br />
The swallows are specialized, but since they are well dif-
16 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
ferentiated from all other Oscines they may be left near the<br />
beginning <strong>of</strong> the sequence, for the time being at least, pending<br />
definite pro<strong>of</strong> <strong>of</strong> affinity to other families. The African flycatcher,<br />
Artnnyias, is very swallow-like and shows how swallows may<br />
have evolved from perching insect catchers. This is probably not<br />
an indication <strong>of</strong> true relationship. For a discussion <strong>of</strong> the genera<br />
<strong>of</strong> swallows, see Mayr and Bond (1943).<br />
The curious African river-martin, Pseudochelidon, is very doubtfully<br />
a member <strong>of</strong> the swallow family, though Lowe (1938) placed<br />
it there. On the other hand, there is still less reason to place it in<br />
the Artamidae, which have completely different nesting habits,<br />
possess powder downs lacking in Pseudochelidon, and are absent<br />
from the Ethiopian region. We have examined the syrinx <strong>of</strong><br />
Pseudochelidon from plentiful material supplied by Dr. James P.<br />
Chapin. The bronchial rings are all half rings, with a large internal<br />
membrane running the length <strong>of</strong> the bronchial tubes.<br />
Likewise, the bronchidesmus connecting the bronchii seems to be<br />
present, as in most song birds, whereas in swallows the bronchial<br />
rings tend to be complete and the bronchidesmus absent, though<br />
the only hirundinids available to us were so poorly preserved as to<br />
make it somewhat difficult to confirm these details.<br />
We leave Pseudochelidon as a subfamily <strong>of</strong> the Hirundinidae,<br />
but probably it belongs in a separate family, possibly near the<br />
Prionopidae, to the members <strong>of</strong> which the bill and texture <strong>of</strong> the<br />
plumage bear some resemblances.<br />
BULBULS AND ALLIES<br />
CAMPEPHAGIDAE<br />
Of the vast group <strong>of</strong> Old World insect eaters the cuckoo-shrikes<br />
are perhaps among the more primitive. In most <strong>of</strong> them the<br />
shafts <strong>of</strong> the dense feathers-<strong>of</strong> the back are stiffened and enlarged,<br />
but this is absent or but weakly indicated in the minivets, Pericrocotus,<br />
and in the genus Chlamydochera <strong>of</strong> Borneo. The recent<br />
transfer <strong>of</strong> the genera Tephrodornis and Hemipus to the Campephagidae<br />
by Delacour and others is thus not necessarily invalidated<br />
by the absence <strong>of</strong> such modified shafts in these genera.<br />
In habits and nidification these two genera resemble cuckooshrikes,<br />
but it is quite possible that they are only "shrike-billed<br />
flycatchers."
1951 A CLASSIFICATION OF RECENT BIRDS 17<br />
IRENIDAE (AEGITHINIDAE)<br />
The leafbirds were formerly associated with the Pyenonotidae<br />
but seem not very close. They may, however, be left in the<br />
general vicinity <strong>of</strong> the cuckoo-shrikes and bulbuls. Since Oberholser<br />
(1917b) proposed a family for the genus Irena, this family<br />
should be called Irenidae, not Aegithinidae. The genera <strong>of</strong> this<br />
family are A ethorhynchus, Aegithina, Chloropsis, and Irena.<br />
PYCNONOTIDAE<br />
Bulbuls are among-the more primitive appearing <strong>of</strong> Old World<br />
song birds. They may be related to the Campephagidae, but a<br />
few, such as Nicator, resemble some <strong>of</strong> the bush shrikes (Laniidae:<br />
Malaconotinae), though perhaps only superficially. Delacour<br />
(1943) has recently revised the family. We would leave the<br />
African genus Neolestes in the Pycnonotidae (Chapin, 1921).<br />
A few other aberrant genera sometimes associated with the<br />
bulbuls are best removed. Hypergerus <strong>of</strong> west Africa is more apt<br />
to be a giant sylviid. Apalopteron <strong>of</strong> the Bonin Islands differs<br />
markedly from bulbuls, particularly by reason <strong>of</strong> its very long<br />
legs. It may well be an aberrant, insular sylviid or timaliid.<br />
We have tentatively placed it in the Timaliinae, near Siva.<br />
The peculiar Tylas eduardi <strong>of</strong> Madagascar may, in the opinion <strong>of</strong><br />
Chapin, be a member <strong>of</strong> the Oriolidae. Oberholser (1917a) set<br />
up a separate family for this genus, but this seems scarcely advisable<br />
pending further knowledge <strong>of</strong> its anatomy. Other<br />
Madagascan genera <strong>of</strong> difficult assignment are dealt with by Delacour<br />
in his revisions <strong>of</strong> the bush warblers (1942), bulbuls (1943),<br />
and babblers (1946). The genus Malia <strong>of</strong> Celebes is hard to place.<br />
It may, as Delacour has suggested, be a bulbul adapted for terrestrial<br />
life.<br />
PRIMITIVE INSECT EATERS<br />
MUSCICAPIDAE<br />
The group embraced by the above descriptive term has sometimes<br />
been called the "Old World insect eaters," but it is impossible<br />
to separate the New World wrens and thrashers from it, while<br />
even the predominantly Old World warblers and babblers have<br />
a few New World representatives, and the thrushes are well represented<br />
there. The phrase "insect eaters" is not, <strong>of</strong> course, to be<br />
taken too literally, since many <strong>of</strong> the thrushes, in particular, feed<br />
to a considerable extent on earthworms, snails, and fruit.
18 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
The satisfactory association <strong>of</strong> many components <strong>of</strong> this group<br />
into families is virtually impossible. This was recognized by<br />
Hartert and many later authorities who have listed the Old World<br />
flycatchers and warblers, as well as the babblers and thrushes, as<br />
subfamilies <strong>of</strong> the Muscicapidae. The Mimidae and Troglodytidae<br />
seem to deserve similar treatment, some <strong>of</strong> the West Indian genera,<br />
in particular, being very thrush-like, while others seem intermediate<br />
between wrens and mockers.<br />
The Turdinae include a number <strong>of</strong> aberrant genera that probably<br />
deserve the status <strong>of</strong> separate tribes or may even be wrongly<br />
placed with this subfamily. By no means all <strong>of</strong> the tropical<br />
thrushes have a "booted" tarsus. Among these difficult genera<br />
are Henicurus and Cochoa in the Old World and Zeledonia in the<br />
New World. The last-named genus was thought by Sharpe and<br />
by Pycraft (1905) to be an aberrant thrush, distantly related to<br />
such genera as Catharus. Erythropygia, sometimes placed with the<br />
Sylviinae, seems correctly assigned to the Turdinae (Chapin).<br />
Our listing <strong>of</strong> tribes is only tentative.<br />
The babblers or Timaliinae have long been a "scrap basket"<br />
for genera that did not fit well into the Turdinae or Sylviinae.<br />
Delacour's (1946) revision brought a semblance <strong>of</strong> order into this<br />
group, but even so he was obliged to set aside a section for "aberrant<br />
genera." To the five tribes that he set up, a sixth, the<br />
Picathartini, including only the west African genus Picathartes,<br />
is to be added (Delacour and Amadon, in press). This genus has<br />
usually been placed in the Corvidae, where it surely does not belong.<br />
We agree with Delacour in placing the wren-tit, Chamaea, <strong>of</strong><br />
California in the tribe Chaemaeini <strong>of</strong> the Timaliinae. This bird<br />
appears to be very closely allied to the genus Moupinia <strong>of</strong> China.<br />
A number <strong>of</strong> genera <strong>of</strong> the Australo-Papuan region (Eupetes,<br />
Cindosoma, Orthonyx, Psophodes, Ifrita, Androphobus, and Melampitta)<br />
do not fit well in the Timaliinae but may for the time<br />
being be left as a tribe, the Cinclosomatini, <strong>of</strong> that subfamily.<br />
Some may consider them as nearer to the Turdinae. These seven<br />
genera probably do not comprise a homogeneous group; Psophodes,<br />
as noted elsewhere, may belong to the Pachycephalini.<br />
Cinclorhamphus, sometimes associated with this group, we tentatively<br />
place in the Malurinae.<br />
The subfamily Sylviinae contains a large number <strong>of</strong> genera and<br />
species difficult to divide satisfactorily into major subgroups
1951 A CLASSIFICATION OF RECENT BIRDS 19<br />
(with the exception <strong>of</strong> the <strong>American</strong> gnatcatchers and allies which<br />
may be considered a tribe). A few genera, such as Megalurulus<br />
<strong>of</strong> New Caledonia, might be placed in either the Sylviinae or<br />
Timaliinae.<br />
Another most peculiar "warbler" is Lamprolia victoria <strong>of</strong> Fiji;<br />
further study may show that it has nothing to do with this family<br />
(Mayr, 1945, p. 137).<br />
Regulus agrees so well with some species <strong>of</strong> Phylloscopus as<br />
to leave little room for doubt that it is a sylviid adapted to boreal<br />
coniferous forests. More doubt exists as regards the central<br />
Asiatic Lophobasileus and Leptopoceile. These little birds suggest<br />
Regulus, but might be related to titmice, notably Aegithalos.<br />
We follow custom in leaving them near Regulus and Phylloscopus.<br />
Their mossy ground nest favors this assignment.<br />
In Australia, New Guinea, Polynesia, and New Zealand there<br />
is a group <strong>of</strong> some 85 species <strong>of</strong> warblers which appears to be better<br />
differentiated than either the kinglets (Regulus) or the gnatcatchers<br />
(Polioptila and allies) and which deserves subfamily<br />
rank under the above name. It includes the birds grouped in the<br />
Acanthizidae in the 1926 "Checklist <strong>of</strong> Australian birds" <strong>of</strong> the<br />
Royal Australasian Ornithologists Union and comprise the following<br />
genera:<br />
Malurus and its closest allies: Clytomyias (New Guinea only),<br />
Chenorhamphus (New Guinea only), Todopsis (New Guinea only),<br />
Malurus, and Stipiturus. The following are less closely allied to<br />
Malurus: Dasyornis, Amytornis, Aphelocephala, Ephthianura<br />
(including A shbyia), Smicrornis, Acanthiza, Pyrrholaema, Finschia<br />
(New Zealand only), Mohoua (New Zealand only), Pycnoptilus,<br />
Calamanthus, Eremiornis, Hylacola, Chthonicola, Gerygone, Hapolorhynchus<br />
(New Zealand only), Sericornis, Vitia, Origmella, and<br />
Crateroscelis. Cinclorhamphus also may belong here.<br />
The Muscicapinae are in need <strong>of</strong> a generic revision, but we have<br />
recognized as tribes the well-defined fantails <strong>of</strong> the genus Rhipidura<br />
(see Oliver, 1945, p. 143) and also the monarch-paradise flycatcher<br />
group (Monarcha, Hypothymis, Terpsiphone, and allies).<br />
Of the Muscicapini proper, some <strong>of</strong> the African genera, such<br />
as Fraseria, have a great resemblance to thrushes. Stizorhina,<br />
formerly placed with the flycatchers, is considered by Chapin to<br />
be a thrush, very close to Neocossyphus.<br />
The shrike-billed flycatchers (Pachycephalini) <strong>of</strong> the Australian<br />
region are <strong>of</strong>ten quite different from the average flycatchers in
20 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
appearance and manner <strong>of</strong> feeding but intergrade completely with<br />
them through such genera as Pachyrephalopsis and Heteromyias.<br />
Concerning the genera <strong>of</strong> the tribe Pachycephalini, Coracornis<br />
and Hylocitrea <strong>of</strong> Celebes and Rhagalogus <strong>of</strong> New Guinea are not<br />
far from Pachycephala proper. Eulacestoma <strong>of</strong> New Guinea, set<br />
apart by its large wattles, is intermediate in specialization <strong>of</strong> the<br />
bill between Pachycephala and Falcunculus. The peculiar<br />
Pachycare, which may not be a flycatcher at all, is best left near<br />
Falcunculus.<br />
Another line leads to rather large, coarse-billed but otherwise<br />
unspecialized forms, from Pachycephala through Oreoica and<br />
Colluricincla (including Myiolestes) to Pitohui. Turnagra, the<br />
so-called New Zealand thrush, probably belongs here rather<br />
than in the Turdinae. Its skull is very different from that <strong>of</strong><br />
Turdus (Oliver, 1945, p. 148). The Australian genus Psophodes,<br />
considered by Delacour to be an aberrant babbler, may belong<br />
near Oreoica in the Pachycephalini.<br />
CINCLINAE<br />
The dippers agree with the Turdinae in having a booted tarsus<br />
and (to some extent) in the mottled immature plumage <strong>of</strong> some<br />
<strong>of</strong> the Old World dippers, characters emphasized by Stejneger<br />
(1905). Other writers have been more impressed by their resemblance<br />
to wrens (Troglodytinae). The classification depends<br />
somewhat on the decision as to whether Cinclus is <strong>of</strong> New or Old<br />
World origin, but the distributional evidence is again somewhat<br />
equivocal. In either case, we do not feel that the characters <strong>of</strong><br />
Cinclus, consisting chiefly <strong>of</strong> adaptations for its aquatic accomplishments,<br />
merit more than subfamily rank.<br />
PRUNELLIDAE<br />
The hedge sparrows resemble buntings (Emberizinae) in appearance<br />
and posture and also by living in part upon seeds and by<br />
having a true crop. On the other hand, the bill is more or less<br />
thrush-like, and the tenth primary is not vestigial as in the buntings.<br />
The Heinroths (1924-1926, p. 34) suggest that Prunella may be<br />
a rather primitive genus related both to thrushes and finches.<br />
Since the Emberizinae are presumably <strong>of</strong> <strong>American</strong> origin and
1951 A CLASSIFICATION OF RECENT BIRDS 21<br />
Prunella is, at least at present, Palearctic, it is possible that the<br />
resemblance <strong>of</strong> the hedge sparrows to the buntings is parallelism.<br />
MOTACILLIDAE<br />
No one believes any longer that the pipits or "tit-larks" are related<br />
to the true larks, yet they are left in that vicinity in most<br />
classifications. The completely nine-primaried wing is correlated<br />
with their rapid sweeping flight and is merely paralleled by the<br />
tendency towards loss <strong>of</strong> the tenth primary in the Alaudidae.<br />
Probably the pipits are modified descendants <strong>of</strong> some group <strong>of</strong><br />
turdids or sylviids, and they are best placed following this group <strong>of</strong><br />
families.<br />
SHRIKES AND ALLIES<br />
This group appears possibly to represent somewhat specialized<br />
<strong>of</strong>fshoots <strong>of</strong> the preceding large group <strong>of</strong> insect eaters. In the<br />
pachycephaline flycatchers <strong>of</strong> the Australian region, the intermediate<br />
steps leading from the Muscicapinae are unmistakable.<br />
The shrikes and their allies may have evolved similarly, but the<br />
intermediates are absent or can no longer be determined with<br />
confidence.<br />
LANIIDAE<br />
The Laniidae <strong>of</strong> early authors included distantly related genera<br />
that had only one feature in common, a hooked, Lanius-like bill.<br />
This functional adaptation has apparently arisen independently<br />
many times in birds with similar feeding habits. Various genera<br />
assigned to the Laniidae at one time or another have since been<br />
placed with the Pachycephalinae, Campephagidae (Tephrodornis,<br />
Hemipus), Vireonidae, Cracticidae, Vangidae, and Prionopidae.<br />
The African bush shrikes (subfamily Malaconotinae) do not<br />
seem to be true shrikes either, but may be left in the Laniidae<br />
until more is known <strong>of</strong> their affinities. Of the 10 genera listed<br />
by Sclater (1924-1930, pp.- 615-638) we would recognize only the<br />
following six, synonymizing those placed in parentheses: Malaconotus<br />
(Chiorophoneus, Telophorus), Rhodophoneus, Tchagra<br />
(Antichromus), Laniarius, Dryoscopus (Chaunonotus), Lanioturdus.<br />
As suggesting the possible relationships <strong>of</strong> this subfamily, one<br />
may note a similarity <strong>of</strong> Malaconotus to certain bulbuls (Nicator).<br />
Lanioturdus (which may not be a bush shrike itself) along with<br />
Nilaus (formerly considered a bush shrike but assigned by Mayr
22 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
[19431 to the Muscicapinae) suggests that the Malaconotinae in<br />
Africa may be allied to the Muscicapinae, notably to such Ethiopian<br />
genera as Batis and Platysteira, in somewhat the same way as<br />
are the Pachycephalinae in the Australian region.<br />
In the Laniinae proper we leave only Lanius and the long-tailed<br />
African shrikes, Urolestes and Corvinella.<br />
PRIONOPIDAE<br />
The prionopids proper are reduced to three African genera<br />
(Prionops, Sigmodus, and Eurocephalus) after the removal <strong>of</strong><br />
various unrelated genera (Mayr, 1943). There is a certain resemblance<br />
between the Prionopidae and some genera <strong>of</strong> African<br />
flycatchers and bush shrikes (Malaconotinae). It is therefore<br />
probable that this family should be associated with the primitive<br />
insect eaters and with the Laniidae.<br />
On geographical premises it seems unlikely that the peculiar<br />
Pityriasis gymnocephala <strong>of</strong> Borneo (one <strong>of</strong> the strangest members<br />
<strong>of</strong> all the Oscines) is related to the Prionopidae. It was for this<br />
reason that Mayr (op. cit.) suggested that it be removed from that<br />
family. This bird does not seem to fit well anywhere else, however,<br />
and agrees with Prionops, and more particularly with Sigmodus<br />
scopifrons, in proportions, wing pattern, and other details.<br />
Moreover, one sees in these African Prionopidae a tendency towards<br />
specialization <strong>of</strong> the head feathers and presence <strong>of</strong> wattles<br />
that might be the forerunner <strong>of</strong> the bristle-like head feathers that<br />
suggested the name "Pityriasis" for the Borneo genus. Yet T. H.<br />
Harrisson (in litt.) is impressed by its "mynah-like" behavior.<br />
VANGIDAE<br />
The vangas are <strong>of</strong> somewhat uncertain affinities but may be<br />
related to the other shrike-like birds, particularly the Prionopidae.<br />
The Vangidae have become so diverse in the isolation provided by<br />
Madagascar that it is difficult to decide what the ancestor <strong>of</strong> the<br />
group was like.<br />
The Australian "magpies" (Cracticidae) resemble shrikes and,<br />
even more, Pityriasis in the shape <strong>of</strong> the bill, but we think they<br />
are more likely to be related to the Australian corvid-like families,<br />
including the Grallinidae, Paradisaeidae, etc., and place them<br />
there.
1951 A CLASSIFICATION OF RECENT BIRDS 23<br />
ARTAMIDAE<br />
The family contains but the one genus, Artamus (unless the<br />
African Pseudochelidon eventually is shown to belong here). The<br />
relationships <strong>of</strong> Artamus are doubtful, but there is a certain resemblance<br />
to some <strong>of</strong> the Vangidae and even to the Bombycillidae.<br />
It may be tentatively left in the vicinity <strong>of</strong> these families.<br />
BOMBYCILLIDAE<br />
The reasons for treating the <strong>American</strong> silky flycatchers, the<br />
West Indian palm chat, Dulus, and the Persian genus Hypocolius<br />
as subfamilies <strong>of</strong> the Bombycillidae have been given by Arvey<br />
(1949) and Delacour and Amadon (1949). The broader question<br />
<strong>of</strong> the relationships <strong>of</strong> the family are much more difficult. As<br />
pointed out to us by Chapin, Hypocolius shares with Eurocephalus,<br />
and to a lesser extent with Prionops, the peculiar feature <strong>of</strong> having<br />
the normally unbroken plates comprising the rear half <strong>of</strong> the<br />
oscinine tarsus divided weakly into a number <strong>of</strong> shields or scutes.<br />
This might mean that the resemblance <strong>of</strong> Hypocolius to other<br />
bombycillids is superficial. Another possibility is that the<br />
Bombycillidae belong in the general vicinity <strong>of</strong> the Prionopidae<br />
and related families. We here tentatively follow the latter alternative.<br />
CREEPERS, NUTHATCHES, AND TITMICE<br />
These birds are all primarily <strong>of</strong> Old World origin and may have<br />
evolved from generalized insect eaters.<br />
CERTHIIDAE<br />
The genus Certhia has had a long history <strong>of</strong> tree creeping, as<br />
shown by the spiny tail and specialized tail molt wherein the two<br />
central feathers are retained until all the others are replaced, thus<br />
giving continuous support. This adaptation occurs elsewhere<br />
only in the Picidae. All the other "creepers" sometimes placed<br />
in the Certhiidae are more like nuthatches, but we leave the<br />
Certhiidae near the Sittidae, should there be any relationship.<br />
SITTIDAE<br />
This family, as we conceive it, is admittedly something <strong>of</strong> a<br />
"scrap basket." We tentatively suggest the following subfamilies:<br />
SALPORNINAE: Salpornis, Rhabdornis, Climacteris, and Tichodroma.<br />
The members <strong>of</strong> these genera all have a rather long, de-
24 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
curved bill. Rhabdornis (Philippines) and Climacteris (Australia)<br />
appear to be related, but whether or not they are allied to Salpornis<br />
(India, Africa) is a moot question. The wall creeper<br />
(Tichodroma) resembles in some ways the nuthatches (Sitta) and<br />
may be a specialized derivative <strong>of</strong> some such bird as the rock<br />
nuthatch (Sitta tephronota). The Heinroths (1924-1926) point<br />
out a number <strong>of</strong> differences between the two genera, and the true<br />
position <strong>of</strong> Tichodroma remains doubtful.<br />
SITTINAE: Sitta, Neositta, and Daphoenositta. The Australo-<br />
Papuan tree-runners (Neositta, Daphoenositta) resemble in many<br />
ways some <strong>of</strong> the nuthatches such as Sittafrontalis. This may be<br />
misleading, since the open nests <strong>of</strong> the various species <strong>of</strong> Neositta<br />
are not at all like those <strong>of</strong> the members <strong>of</strong> Sitta. Perhaps Neositta<br />
is related to Climacteris.<br />
HYPOSITTINAE: The coral-billed "nuthatch" <strong>of</strong> Madagascar is<br />
another puzzling bird. It suggests some <strong>of</strong> the nuthatches, but<br />
the proportions are those <strong>of</strong> the true creepers (Certhia), which it<br />
resembles in habits. The tail feathers are unmodified. Rand<br />
(1936a, 1936b) has written on the habits and position <strong>of</strong> Hypositta,<br />
Neositta, and Daphoenositta.<br />
PARIDAE<br />
The tits appear to be a polyphyletic group <strong>of</strong> genera placed together<br />
on the basis <strong>of</strong> superficial resemblance due to arboreal<br />
habits and a-fine, more or less conical bill. Three groups can be<br />
distinguished:<br />
1. The true titmice. This consists <strong>of</strong> the genus Parus and<br />
the closely related Melanochlora (sultan tits). Sylviparus is a<br />
dubious member <strong>of</strong> this group.<br />
There is a remarkable resemblance between some <strong>of</strong> the jays,<br />
particularly the boreal Perisoreus, and the genus Parus. This<br />
similarity does not extend to the skull or the internal anatomy,<br />
and Ridgway (1904, pp. 253-254) stated that even the external<br />
features "when closely examined, show many points <strong>of</strong> difference."<br />
It is probable, therefore, that the like manner in which jays,<br />
titmice, and even nuthatches hold nuts or seeds in their feet while<br />
opening them or store them away in crevices for future use has no<br />
great phylogenetic significance. Kramer (1930) has published<br />
data on the occurrence <strong>of</strong> such habits in various groups <strong>of</strong> birds.<br />
A parallel case is the habit <strong>of</strong> the true shrikes (Lanius) and <strong>of</strong><br />
the unrelated Australian "shrikes" (Cracticus) <strong>of</strong> impaling and<br />
dismembering small birds and rodents.
1951 A CLASSIFICATION OF RECENT BIRDS 25<br />
2. Remiz-Auriparus group. These birds may be distantly related<br />
to the Dicaeidae (flowerpeckers). They are characterized<br />
by the peculiarly shaped bill and the bag-like nest they construct.<br />
Cephalopyrus may belong to this group, though it is a hole nester.<br />
3. Aegithalos-Psaltriparus-Psaltria. In their complete juvenal<br />
molt (Stresemann, 1923), cranial characters, and nest structure<br />
(Lucas, 1890), the long-tailed tits and bush tits differ radically<br />
from the Paridae. The young hatch naked, while young <strong>of</strong> the<br />
true titmice are partly downy. They share several <strong>of</strong> these<br />
characters with Panurus and Paradoxornis and are perhaps nothing<br />
but an <strong>of</strong>fshoot <strong>of</strong> the Timaliinae. The fact that the parrotbills<br />
make a cup-like nest, while the nest <strong>of</strong> the present group is<br />
bag-like, is not, however, favorable to this suggestion.<br />
It is now universally agreed that Panurus is a close relative <strong>of</strong><br />
Paradoxornis. Delacour (1946) places it, together with Chamaea,<br />
in the tribe Chamaeini <strong>of</strong> the Timaliinae, along with the other<br />
"parrot-bills."<br />
OLD WORLD NECTAR EATERS<br />
DICAEIDAE<br />
The primitive fruit-eating dicaeid genus Melanocharis <strong>of</strong> New<br />
Guinea has a great resemblance to certain genera <strong>of</strong> Meliphagidae<br />
(Timeliopsis, Oedistoma). Since the latter have a highly specialized<br />
tongue and that <strong>of</strong> Melanocharis is unspecialized (Mayr and<br />
Amadon, 19!47), this resemblance may not indicate affinity.<br />
Another moot affinity <strong>of</strong> the flowerpeckers is with the birds<br />
placed by us in the subfamily Remizinae <strong>of</strong> the Paridae. The<br />
genus Cephalopyrus <strong>of</strong> the Himalayas could be an intermediate.<br />
Finally, one may mention, as possibly repaying study, the resemblance<br />
<strong>of</strong> certain flowerpeckers to the Bombycillidae in appearance,<br />
fondness for mistletoe berries, gregarious habits,<br />
and other details <strong>of</strong> behavior.<br />
NECTARINIIDAE<br />
The sunbirds, a very compact group (Delacour, 1944), are evidently<br />
relatives <strong>of</strong> the Dicaeidae, highly specialized for nectar<br />
feeding. The tongue is quite similar in sunbirds (Scharnke, 1932)<br />
and in the nectar-feeding kinds <strong>of</strong> flowerpeckers, and both build<br />
a similar pensile nest. Though more specialized as regards plumage<br />
and modification <strong>of</strong> the tongue, sunbirds do not show the<br />
progressive loss <strong>of</strong> the tenth primary culminating in the nine-
26 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
primaried condition <strong>of</strong> such dicaeid genera as Pardalotus and<br />
many species <strong>of</strong> Dicaeum.<br />
The Dicaeidae are best represented in Australia, Papua, the<br />
East Indies, and the Philippines, and do not reach Africa; almost<br />
the reverse is true <strong>of</strong> the Nectariniidae.<br />
The curious African genera, Pholidornis and Parmoptila, formerly<br />
associatedwith the Dicaeidae orNectariniidae, are considered<br />
by Chapin (1917) to be aberrant, thin-billed Ploceidae, with<br />
which we agree.<br />
The African species Hylia prasina has usually been placed in<br />
the Sylviinae and may be an aberrant member <strong>of</strong> that group.<br />
Bates (1930, p. 447) figured the hyoid bones <strong>of</strong> this species, which<br />
he found to be flattened like those <strong>of</strong> a sunbird. The tenth primary<br />
is rather larger than in typical sunbirds. He placed Hylia<br />
in a separate family but thought it was allied to the Nectariniidae.<br />
Serle (1949, p. 212), among others, found its habits and call notes<br />
to suggest those <strong>of</strong> a sunbird, though the song is more musical.<br />
The nest is more or less globular, not much like that <strong>of</strong> a sunbird.<br />
Hylia may be a primitive sunbird. Chapin considers this possible<br />
but not established.<br />
MELIPHAGIDAE<br />
The numerous members <strong>of</strong> this Australo-Papuan family have,<br />
with one or two exceptions, a highly modified suctorial tongue,<br />
unlike that <strong>of</strong> flowerpeckers or sunbirds and surely <strong>of</strong> independent<br />
evolution. Otherwise they are rather unspecialized Oscines, except<br />
for the frequent presence <strong>of</strong> ornamental tufts <strong>of</strong> yellow<br />
feathers or <strong>of</strong> bare areas or wattles around the eyes. As noted<br />
above, they may be distantly related to the Dicaeidae.<br />
The curious nectar-feeding sugar bird, Promerops cafer <strong>of</strong><br />
South Africa, has a stomach and tongue like those <strong>of</strong> the Australian<br />
meliphagids (Scharnke, 1932). Its external appearance is also<br />
similar. It seems definitely not to be a sunbird (Nectariniidae).<br />
Promerops feeds on flowers <strong>of</strong> trees <strong>of</strong> the genus Protea, plants also<br />
well represented in Australia. Salomonsen (1933) advanced the<br />
theory that Promerops was once much more widespread (along<br />
with Protea) and that both were gradually restricted to South<br />
Africa. The Meliphagidae are, however, so strictly Australian<br />
in distribution that it is difficult to believe that one genus wandered<br />
so widely while all the others failed to make their way even<br />
to Asia, much less to Africa. Also, many Meliphagidae are not
1951 A CLASSIFICATION OF RECENT BIRDS 27<br />
specialized as to food plants. The possibility <strong>of</strong> some South<br />
African-Australian transfer remains, <strong>of</strong> course, highly speculative.<br />
We leave Promerops as a subfamily <strong>of</strong> the Meliphagidae, but we<br />
feel that its similarity may, after all, be parallelism.<br />
ZOSTEROPIDAE<br />
The relationships <strong>of</strong> the Zosteropidae remain to be discovered.<br />
Since some <strong>of</strong> them are somewhat specialized for feeding on<br />
nectar, they may continue to be left in the vicinity <strong>of</strong> the Meliphagidae<br />
and Dicaeidae. The nine-primaried wing indicates a<br />
considerable degree <strong>of</strong> specialization.<br />
VIREOS, TANAGERS, FINCHES, AND ALLIES<br />
The Vireonidae may be the most primitive family <strong>of</strong> this assemblage.<br />
Its members <strong>of</strong>ten have a functional tenth primary,<br />
and they are otherwise rather unspecialized, usually insectivorous<br />
birds. The shrike-billed vireos, Vireolanius, and pepper-shrikes,<br />
Cyclarhis, have the longest tenth primaries and may well be the<br />
most primitive <strong>of</strong> the vireos, despite their large size and bright<br />
colors. It seems best to place these genera in the Vireonidae and<br />
not to set up monotypic families for them (Zimmer, 1942).<br />
The wood warblers (Parulinae) are perhaps most like the vireos.<br />
The warblers in turn are not far from the Coerebinae, while some<br />
<strong>of</strong> the latter have always been admitted to be very difficult to<br />
separate from many tanagers (Thraupinae). All <strong>of</strong> these groups<br />
(except the vireos) may best be considered subfamilies <strong>of</strong> the<br />
Thraupidae. Additional subfamilies are the plush-capped finch,<br />
Catamblyrhynchus, which is not aberrant enough to require a<br />
family, and the pyrrhuloxias and cardinal grosbeaks (Pyrrhuloxiinae).<br />
The latter, as is well known, are very difficult to separate<br />
from the tanagers. Such a genus as Saltator provides a<br />
transition. Richmondena seems to be congeneric with Pyrrhuloxia,<br />
so the subfamily takes its name from the latter genus.<br />
The swallow-tanager, Tersina, has a very peculiar palate (Lucas,<br />
1895). Its habit <strong>of</strong> nesting in holes in trees is also remarkable.<br />
It is best to leave it in a separate family for the present.<br />
Whether the Pyrrhuloxiinae are closely related to the buntings<br />
and <strong>American</strong> sparrows (Emberizinae) or merely parallel them will<br />
remain uncertain until careful studies have been made <strong>of</strong> the<br />
numerous, chiefly Neotropical, genera. Until such time, it is
28 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
best to keep the buntings as a subfamily <strong>of</strong> the Fringillidae.<br />
The only other subfamily <strong>of</strong> this family will be the Fringillinae<br />
which includes the chaffinches, linnets, evening grosbeaks, and<br />
others. It is the only subfamily <strong>of</strong> the entire assemblage <strong>of</strong><br />
families better represented in the Old World than in the New,<br />
but even this group is presumably <strong>of</strong> <strong>American</strong> origin. If the<br />
Ploceidae should be found to be related to the finches, which we<br />
doubt, the center <strong>of</strong> origin <strong>of</strong> the Fringillinae and perhaps other<br />
subfamilies must be viewed in a different light.<br />
Sushkin (1924) separated the chaffinches, Fringilla, in a subfamily<br />
distinct from the goldfinches and their relatives ("Carduelinae").<br />
He gave no reason for this separation, and we think it unnecessary.<br />
A few puzzling genera.may be mentioned. The extinct, hugebilled<br />
Chaunoproctus <strong>of</strong> the Bonin Islands is, in our opinion, related<br />
to Carpodacus, not to the Pyrrhuloxiinae (pair in Leiden<br />
examined in 1950). Neospiza concolor <strong>of</strong> San Tome is now recognized<br />
to be a finch, not a weaverbird, and is to be placed in the<br />
Fringillinae next to Poliospiza (fide Chapin). The two species <strong>of</strong><br />
Nesospiza from the Tristan da Cunha group seem to belong to<br />
the Fringillinae, perhaps near Serinus, although Lowe and others<br />
are correct in believing that the Gough Island finch, Rowettia, is<br />
not related to Nesospiza but was derived from the South <strong>American</strong><br />
genus Melanodera. From the latter it is scarcely separable<br />
generically.<br />
In the "Catalogue <strong>of</strong> birds <strong>of</strong> the Americas, part 11" Hellmayr<br />
(1938) lists no fewer than 28 genera in the Fringillinae and "Carduelinae."<br />
Of these only the following seem properly to belong<br />
there: Fringilla, Coccothraustes, Hesperiphona, Pyrrhula, Carpodacus,<br />
Pinicola, Leucosticte, Chloris, Carduelis, Acanthis,<br />
Loximitris, Spinus, and Loxia. The extraneous genera included<br />
by Hellmayr belong for the most part to the Pyrrhuloxiinae and<br />
perhaps, in a few cases, to the Emberizinae. This includes a few<br />
genera, such as Sicalis, which resemble goldfinches, Spinus, but<br />
apparently only superficially. In the Old World there has been<br />
less trouble in delimiting the Fringillinae, with the exception <strong>of</strong> the<br />
genus Montifringilla, which resembles Leucosticte but is actually a<br />
member <strong>of</strong> the Ploceidae.<br />
The Hawaiian honeycreepers (Drepaniidae) while closely related<br />
to the families discussed above are rather difficult to align<br />
with any one <strong>of</strong> them (Amadon, 1950b). For this reason we leave
1951 A CLASSIFICATION OF RECENT BIRDS 29<br />
them as a family. An insectivorous and nectarivorous rather<br />
than frugivorous or granivorous type <strong>of</strong> bird seems most primitive<br />
in this family.<br />
The Icteridae are a better differentiated and perhaps a somewhat<br />
older group than most <strong>of</strong> the other nine-primaried passerines.<br />
There is a remarkable amount <strong>of</strong> parallelism between the Icteridae<br />
as compared with the Ploceidae and Sturnidae, which might indicate<br />
distant relationship.<br />
WEAVERBIRDS, STARLINGS, AND ASSOCIATED FAMILIES<br />
Among the more important papers on the Ploceidae are those<br />
<strong>of</strong> Chapin (1917), Sushkin (1927), and Delacour (1943 and earlier).<br />
It is quite possible that the family is composite. Of the more<br />
typical weavers, the genera Bubalornis and Dinemellia seem the<br />
most primitive. They contain species that are rather large,<br />
coarse, heavy-billed birds suggestive <strong>of</strong> starlings (von Boetticher,<br />
1931), and presumably may represent the stock ancestral to<br />
Ploceus and its relatives. On the other hand, the Estrildinae may<br />
well have evolved from thin-billed, small birds similar to the living<br />
species <strong>of</strong> Pholidornis, Parmoptila, and Nigrita. That Pholidornis<br />
is a weaver is still open to question, but Chapin considers<br />
it probable.<br />
The parasitic Viduinae resemble their hosts, the Estrildinae,<br />
even to the mouth spots <strong>of</strong> the nestlings. Some consider this<br />
mimicry, as indeed it may be in part, but Chapin believes the two<br />
subfamilies to be closely allied, if, indeed, subfamily separation is<br />
necessary. The other parasitic weaver, Anomalospiza imberbis,<br />
belongs to the Ploceinae. It is parasitic on warblers, not upon<br />
other weavers. We think it unnecessary to recognize subfamilies<br />
for Plocepasser (Passerinae) or Sporopipes (Ploceinae).<br />
STURNIDAE<br />
The genera <strong>of</strong> the Sturnidae were revised by Amadon (1943).'<br />
1 A correction to that paper may be given here. Speculipastor bicolor Reichenow<br />
was placed in the genus Sprec and renamed bicoloratus to avoid conflict with Spreo bicolor<br />
Gmelin. Dr. H. von Boetticher has brought to my attention that Reichenow<br />
later (1914, p. 356) made his own genus Speculipastor a synonym <strong>of</strong> Spreo and used the<br />
name speculiferus for the species he had named bicolor. This was done in such a cryptic<br />
manner that the name was missed in all subsequent general lists. The name speculiferus<br />
Reichenow will stand, however, with bicoloratus and bicolor as synonyms, so<br />
long as this species is placed in the genus Spreo.-D. A.
30 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
Since then Chapin (1948) has shown that Neocichla belongs to this<br />
family.<br />
The primitive buffalo-weavers (Bubalornithinae) bear a certain<br />
resemblance to some <strong>of</strong> the starlings (von Boetticher, 1931),<br />
and the two families may be related, although this cannot be<br />
taken as definitely established. Some <strong>of</strong> the Papuan glossy<br />
starlings <strong>of</strong> the genus Aplonis nest in colonies and build long,<br />
hanging nests similar to those <strong>of</strong> such species as Ploceus philippinus.<br />
Both Passer domesticus (Ploceidae) and Acridotheres<br />
tristis (Sturnidae) lay their eggs either in holes or in domed tree<br />
nests, according to local conditions. Whether this similarity in<br />
nesting habits is indicative <strong>of</strong> relationship or not is another question.<br />
The bill <strong>of</strong> the African ox-bird, Buphagus, and <strong>of</strong> the Celebesian<br />
starling, Scissirostrum dubium, are slightly like that <strong>of</strong> some<br />
weavers, but this is pure convergence. Scissirostrum is a specialized,<br />
not a primitive, starling as regards bill form and function<br />
(Amadon, 1943).<br />
The long-extinct Fregilupus varius <strong>of</strong> the Mascarene Islands is<br />
not unlike some <strong>of</strong> the Vangidae in color pattern. Recent comparison<br />
<strong>of</strong> mounted specimens <strong>of</strong> Fregilupus in the museums at Leiden<br />
and at Paris convinced the junior author that it is correctly<br />
assigned to the Sturnidae. The outermost primary is much reduced<br />
as in most Sturnidae, not long as in the Vangidae. Presumably<br />
the same is true <strong>of</strong> Necropsar leguati, another long-extinct<br />
Mascarene species.<br />
ORIOLIDAE; DICRURIDAE<br />
The orioles and drongos parallel in degree <strong>of</strong> specialization some<br />
<strong>of</strong> the starlings, though they are not necessarily related to them.<br />
Indeed, it may be that both families are better placed in the vicinity<br />
<strong>of</strong> the Campephagidae and Irenidae, but more evidence is needed.<br />
For a revision <strong>of</strong> the Dicruridae, see Vaurie (1949).<br />
As stated above, we tentatively place the genus Tylas <strong>of</strong> Madagascar<br />
in the Oriolidae.<br />
CROWS AND AUSTRALIAN CORVID-LIKE FAMILIES<br />
CORVIDAE<br />
For a revision <strong>of</strong> the genera, see Amadon (1944). Lowe (1949)<br />
has since established a monotypic family for Zavattariornis, but<br />
this Abyssinian genus seems to be a generalized corvid. Our rea-
1951 A CLASSIFICATION OF RECENT BIRDS 31<br />
sons for following the earlier authorities who placed the crows at<br />
the apex <strong>of</strong> the Class Aves were mentioned earlier.<br />
The jays are the most primitive <strong>of</strong> the Corvidae. From them<br />
evolved first the magpies and finally Corvus, a genus that contains<br />
what are in many ways the most adaptable and highly evolved <strong>of</strong><br />
all birds.<br />
The Australian families associated with the crows resemble<br />
the Corvidae and equal them in degree <strong>of</strong> specialization but may<br />
well be <strong>of</strong> independent evolution. They all have a tarsus that is<br />
weakly scutellated or even booted, a point <strong>of</strong> difference from the<br />
true corvids.<br />
GRALLINIDAE<br />
In this family we unite the three genera <strong>of</strong> so-called Australian<br />
mud-nest builders, Grallina, Corcorax, and Struthidea. The last<br />
two give many indications <strong>of</strong> relationship despite the difference in<br />
the bill (Amadon, 1944). Grallina (including the Papuan Pomareopsis)<br />
is a very peculiar genus, but the fact that it builds the same<br />
bowl-shaped mud nest as the other two suggests that it may,<br />
after all, be related to them. A preliminary study by the junior<br />
author, including comparison <strong>of</strong> the skulls <strong>of</strong> all three genera, has<br />
not brought to light any basic differences (Amadon, 1950a).<br />
Partly to avoid too many monotypic families, we place all three<br />
genera in the same family but keep Grallina in a separate subfamily.<br />
Should this course prove correct, the variation in the bill<br />
<strong>of</strong> these genera would parallel that in the three genera <strong>of</strong> the New<br />
Zealand Callaeidae. Indeed, the two families have much in common.<br />
CALLAEIDAE<br />
Stonor (1942) showed that Callaeas, Philesturnus, and Neomorpha<br />
[= Heteralochac] belong in a single group. He accepted<br />
Garrod's (1872) supposed evidence for the affinity <strong>of</strong> Neomorpha<br />
to the Sturnidae, but perusal <strong>of</strong> that paper does not indicate that<br />
Garrod himself meant to imply any close ties. At that time Garrod<br />
included the Icteridae in the Sturnidae, and he apparently<br />
found as much resemblance in the icterid as in the true sturnid<br />
skull to that <strong>of</strong> the Huia. The chief point <strong>of</strong> resemblance consisted<br />
<strong>of</strong> facets for the attachment <strong>of</strong> the large digastric muscles,<br />
which, as Lowe (1938) and Lorenz (1949) have shown, are an<br />
adaptation for operating a long bill that can be opened forcibly
32 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 149W.<br />
against resistance. We do not think the Calleidae and Sturnidae<br />
are allied.<br />
CRACTICIDAE<br />
The bell magpies or piping crows <strong>of</strong> the Australian region, as<br />
mentioned earlier, resemble the Laniidae and associated families,<br />
but in our opinion their real relationship is probably with the<br />
Australian and New Zealand corvid-like families. The Cracticidae<br />
agree with these families by having the tarsus almost booted.<br />
Pycraft thought the Cracticidae to be very near to the bowerbirds<br />
and birds <strong>of</strong> paradise. There is a striking resemblance in color<br />
pattern between Grallina and some species <strong>of</strong> Cracticus, but Grallina<br />
is so aberrant that this cannot safely be used as a basis for<br />
comparison.<br />
In the largest forms <strong>of</strong> Strepera (Cracticidae) the plumage is,<br />
almost or entirely black, and the hook on the bill may be absent<br />
in adults. The result is a bird very much like a crow, Corvus,<br />
but this is parallelism, although the families to which these genera<br />
belong may not be very distantly related. For a further discussion<br />
and revision <strong>of</strong> the Cracticidae, see Amadon (in press).<br />
PTILONORHYNCHIDAE; PARADISAEIDAE<br />
Stonor (1937) found some consistent differences between the<br />
skulls <strong>of</strong> bower birds and birds <strong>of</strong> paradise so that the two groups<br />
may be placed in separate families. Similarities in behavior and<br />
in general morphology, however, indicate that they are not distantly<br />
related.<br />
SYSTEMATIC LIST<br />
All living or recent orders and families are given in the following<br />
list, but subfamilies and tribes have been introduced only as<br />
necessary to indicate new groupings. Suborders have been listed<br />
only in the Passeres. For the song birds, vernacular groupings<br />
are used also, to aid in associating related families.<br />
The numbers <strong>of</strong> recent known species for each unit follow the<br />
names, those <strong>of</strong> subfamilies being in parentheses.<br />
Order Struthiones<br />
Family Struthionidae, Ostrich, 1<br />
Order Apteryges, Moas (fossil) and Kiwis<br />
Family Apterygidae, Kiwis, 3
1951 A CLASSIFICATION OF RECENT BIRDS 33<br />
Order Casuarii<br />
Family Casuariidae, Cassowaries, 3<br />
Family Dromaeidae, Emus, 2<br />
Order Rheae<br />
Family Rheidae, Rheas, 2<br />
Order Crypturi (Tinamiformes)<br />
Family Tinamidae, Tinamous, 33<br />
Order Sphenisci<br />
Family Spheniscidae, Penguins, 16<br />
Order Tubinares (Procellariiformes)<br />
Family Diomedeidae, Albatrosses, 13<br />
Family Procellariidae, 73<br />
Subfamily Hydrobatinae, Storm Petrels, (20)<br />
Subfamily Procellariinae, Shearwaters, etc. (53)<br />
Family Pelecanoididae, Diving Petrels, 4<br />
Order Podicipedes (Colymbiformes)<br />
Family Podicipitidae, Grebes, 20<br />
Order Gaviae<br />
Family Gaviidae, Loons, 4<br />
Order Steganopodes (Pelecaniformes)<br />
Family Phaethontidae, Tropic Birds, 3<br />
Family Fregatidae, Frigate Birds, 5<br />
Family Phalacrocoracidae, Cormorants, Anhingas, 31<br />
Subfamily Phalacrocoracinae, Cormorants, (30)<br />
Subfamily Anhinginae, Anhingas, (1)<br />
Family Sulidae, Boobies, Gannets, 9<br />
Family Pelecanidae, Pelicans, 6<br />
Order Falcones or Accipitres<br />
Family Accipitridae, Hawks, Eagles, etc., 205<br />
Family Falconidae, Falcons, etc., 58<br />
Family Pandionidae, Osprey, 1<br />
Family Cathartidae, New World Vultures, 6<br />
Family Sagittariidae, Secretary Bird, 1<br />
Order Gressores (Ciconiiformes)<br />
Family Ardeidae, Herons, etc., 59<br />
Family Threskiornithidae, Ibises, etc., 28<br />
Family Ciconiidae, Storks, 17<br />
Family Scopidae, Hammerhead, 1<br />
Order Phoenicopteri<br />
Family Phoenicopteridae, Flamingos, 6<br />
Order Anseres<br />
Family Anatidae, Swans, Geese, Ducks, 145<br />
Family Anhimidae, Screamers, 3<br />
Order Galli<br />
Family Megapodiidae, Brush Turkeys, 10<br />
Family Cracidae, Guans, etc., 38<br />
Family Phasianidae, Pheasants, etc., 190<br />
Subfamily Phasianinae, Pheasants, Quail, etc., (16-5)<br />
Subfamily Numidinae, Guineafowl, (7)
34 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
Subfamily Tetraoninae, Grouse, (18)<br />
Family Meleagrididae, Turkeys, 2<br />
Family Opisthocomidae, Hoatzin, 1<br />
Order Cuculi<br />
Family Musophagidae, Turacos, 19<br />
Family Cuculidae, Cuckoos, 128<br />
Order Grues<br />
Family Cariamidae, Seriamas, 2<br />
Family Psophiidae, Trumpeters, 3<br />
Family Gruidae, Cranes, 14<br />
Family Aramidae, Limpkin, 1<br />
Family Eurypygidae, Sun Bittern, 1<br />
Family Heliornithidae, Sun Grebes, 3<br />
Family Rhynochetidae, Kagu, 1<br />
Family Otididae, Bustards, 23<br />
Family Rallidae, Rails, 132<br />
Family Mesoenatidae, Roatelos, 3<br />
Family Turnicidae, Button Quails, 16<br />
Subfamily Turnicinae, Typical Button Quails, (15)<br />
Subfamily Pedionominae, Plains Wanderer or Collared Hemipode, (1)<br />
Order Laro-Limicolae (Charadriiformes)<br />
Family Jacanidae, Jacanas, 7<br />
Family Thinocoridae, Seed Snipe, 4<br />
Family Chionididae, Sheath Bills, 2<br />
Family Dromadidae, Crab Plover, 1<br />
Family Burhinidae, Thick-knees, 9<br />
Family Haematopodidae, Oystereatchers, 6<br />
Family Charadriidae, Plovers, Sandpipers, etc., 152<br />
Subfamily Charadriinae, Plovers, (63)<br />
Subfamily Scolopacinae, Sandpipers, etc., (77)<br />
Subfamily Phalaropinae, Phalaropes, (3)<br />
Subfamily Recurvirostrinae, Avocets, Stilts, (7)<br />
Subfamily Rostratulinae, Painted Snipe, (2)<br />
Family Glareolidae, Pratincoles, Coursers, 16<br />
Family Laridae, Gulls, etc., 89<br />
Subfamily Stereorariinae, Jaegers, Skuas, (4)<br />
Subfamily Larinae, Gulls, (43)<br />
Subfamily Sterninae, Terns, (39)<br />
Subfamily Rynchopinae, Skimmers, (3)<br />
Family Alcidae, Auks, etc., 22<br />
Order Columbae<br />
Family Pteroclidae, Sand Grouse, 16<br />
Family Columbidae, Pigeons, 289<br />
Family Raphidae, Dodos, 3<br />
Order Psittaci<br />
Family Psittacidae, Parrots, 316<br />
Order Striges<br />
Family Strigidae, Owls, 134<br />
Subfamily Striginae, Typical Owls, (123)
1951 A CLASSIFICATION OF RECENT BIRDS 3.5<br />
Subfamily Tytoninae, Barn Owls, (11)<br />
Order Caprimulgi<br />
Family Aegothelidae, Owlet Frogmouths, 8<br />
Family Podargidae, Frogmouths, 12<br />
Family Caprimulgidae, Goatsuckers, etc., 67<br />
Family Nyctibiidae, Potoos, 5<br />
Family Steatornithidae, Oil Bird, 1<br />
Order Trogones<br />
Family Trogonidae, Trogons, 35<br />
Order Coraciae<br />
Family Coraciidae, Rollers, 17<br />
Subfamily Leptosomatinae, Cuckoo Rollers, (1)<br />
Subfamily Brachypteraciinae, Ground Rollers, (5)<br />
Subfamily Coraciinae, Typical Rollers, (11)<br />
Family Alcedinidae, Kingfishers, 87<br />
Family Meropidae, Bee Eaters, 25<br />
Family Momotidae, Motmots, 8<br />
Family Todidae, Todies, 5<br />
Family Upupidae, Hoopoes, 7<br />
Subfamily Upupinae, Typical Hoopoe, (1)<br />
Subfamily Phoeniculinae, Tree Hoopoes, (6)<br />
Family Bucerotidae, Hornbills, 45<br />
Order Colii<br />
Family Coliidae, Mousebirds, 6<br />
Order Macrochires (Apodiformes; Micropodiformes)<br />
Family Apodidae, Swifts, 79<br />
Subfamily Apodinae, Typical Swifts, (76)<br />
Subfamily Hemiprocninae, Crested Swifts, (3)<br />
Family Trochilidae, Hummingbirds; 319<br />
Order Pici<br />
Family Bucconidae, Puffbirds, 32<br />
Family Galbulidae, Jacamars, 14<br />
Family Capitonidae, Barbets, 76<br />
Family Picidae, Woodpeckers, etc., 210<br />
Subfamily Jynginae, Wrynecks, (2)<br />
Subfamily Picumninae, Piculets, (29)<br />
Subfamily Picinae, Woodpeckers, (179)<br />
Family Ramphastidae, Toucans, 37<br />
Family Indicatoridae, Honey Guides, 12<br />
Order Passeres<br />
Suborder Eurylaimi<br />
Family Eurylaimidae, Broadbills, 14<br />
Suborder Tyranni<br />
Superfamily Furnarioidea (Tracheophonae)<br />
Family Rhinocryptidae, Tapaculos, 26<br />
Family Conopophagidae, Ant-pipits, 10<br />
Family Formicariidae, Antbirds and Allies, 221<br />
Family Furnariidae, Ovenbirds and Woodhewers, 259<br />
Subfamily Furnariinae, Ovenbirds, (212)
36 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
Subfamily Dendrocolaptinae, Woodhewers, (47)<br />
Superfamily Tyrannoidea (Haploophonae)<br />
Family Pittidae, Pittas, 23<br />
Family Philepittidae, Philepittas, 4<br />
Family Xenicidae, New Zealand Wrens, 4<br />
Family Tyrannidae, Tyrants, Sharp-bills, 366<br />
Subfamily Tyranninae, Tyrant Flycatchers, (365)<br />
Subfamily Oxyruncinae, Sharp-bills, (1)<br />
Family Pipridae, Manakins, 59<br />
Family Cotingidae, Cotingas, 90<br />
Family Phytotomidae, Plant-cutters, 3<br />
Suborder Menurae<br />
Family Menuridae, Lyrebirds, 2<br />
Family Atrichornithidae, Scrub-birds, 2<br />
Suborder Oscines, Song Birds<br />
Family Alaudidae, Larks, 75<br />
Family Hirundinidae, Swallows, 75<br />
Subfamily Hirundininae, Typical Swallows, (74)<br />
Subfamily Pseudochelidoninae, African River-martin, (1)<br />
BULBULS AND ALLIES<br />
Family Pycnonotidae, Bulbuls, 109<br />
Family Irenidae, Fairy Bluebirds and Leafbirds, 14<br />
Family Campephagidae, Caterpillar Birds or Cuckoo-shrikes, 72<br />
PRIMITIVE INSECT EATERS<br />
Family Muscicapidae, 1460<br />
Subfamily Muscicapinae, Old World Flycatchers, (378)<br />
Tribe Muscicapini, Typical Flycatchers<br />
Tribe Monarchini, Monarchs<br />
Tribe Rhipidurini, Fantails<br />
Tribe Pachycephalini, Whistlers<br />
Subfamily Timaliinae, Babblers, (282)<br />
Tribe Pellorneini, Jungle Babblers<br />
Tribe Pomatorhinini, Scimitar and Wren Babblers<br />
Tribe Timaliini, Tit-babblers<br />
Tribe Chamaeini, Wren-tits and Parrotbills<br />
Tribe Turdoidini, Laughing Thrushes, etc.<br />
Tribe Picathartini, Picathartes<br />
Tribe Cinclosomatini, Ground Babblers, etc.<br />
Subfamily Sylviinae, Warblers (313)<br />
Tribe Sylviini, Typical Warblers (includes Regulus)<br />
Tribe Polioptilini, Gnatcatchers, etc.<br />
Subfamily Malurinae, Australian Warblers, (85)<br />
Subfamily Turdinae, Thrushes, (304)<br />
Tribe Turdini, Typical Thrushes<br />
Tribe Zeledoniini, Wren-thrush<br />
Tribe Myiophoneini, Blue Thrushes<br />
Tribe Cochoini, Cochoas
1951 A CLASSIFICATION OF RECENT BIRDS 37<br />
Tribe Enicurini, Fork Tails<br />
Subfamily Miminae, Mockers and Thrashers, (30)<br />
Subfamily Troglodytinae, Wrens, (63)<br />
Subfamily Cinclinae, Dippers, (5)<br />
Family Prunellidae, Hedge Sparrows or Accentors, 12<br />
Family Motacillidae, Wagtails and Pipits, 48<br />
SHRIKES AND ALLIES<br />
Family Laniidae, Shrikes, 67<br />
Subfamily Laniinae, Typical Shrikes, (25)<br />
Subfamily Malaconotinae, Bush Shrikes, (42)<br />
Family Prionopidae, Helmet Shrikes, 14<br />
Subfamily Prionopinae, Helmet Shrikes, (13)<br />
Subfamily Pityriasidinae, Bornean Bristle-head, (1)<br />
Family Vangidae, Vangas, 11<br />
WAXWINGS AND WOOD SWALLOWS<br />
Family Artamidae, Wood Swallows, 10<br />
Family Bombycillidae, Waxwings, etc., 9<br />
Subfamily Hypocoliinae, Hypocolius, (1)<br />
Subfamily Dulinae, Palm Chat, (1)<br />
Subfamily Ptilogonatinae, Silky Flycatchers, (4)<br />
Subfamily Bombycillinae, Waxwings, (3)<br />
CREEPERS, NUTHATCHES, AND TITMICE<br />
Family Certhiidae, Typical Creepers, 6<br />
Family Sittidae, Nuthatches, etc., 29<br />
Subfamily Salporninae, Spotted Creepers, etc., (12)<br />
Subfamily Sittinae, Nuthatches, etc., (16)<br />
Subfamily Hyposittinae, Coral-billed Nuthatch, (1)<br />
Family Paridae, Titmice, 64<br />
Subfamily Parinae, Typical Titmice and Chickadees (46)<br />
Subfamily Remizinae, Penduline Titmice and Verdins, (8)<br />
Subfamily Aegithalinae, Long-tailed Titmice and Bush Tits, (7)<br />
OLD WORLD NECTAR EATERS<br />
Family Dicaeidae, Flowerpeckers, 54<br />
Family Nectariniidae, Sunbirds, 104<br />
Family Meliphagidae, Honeyeaters, 160<br />
Subfamily Meliphaginae, Australian Honeyeaters, (159)<br />
Subfamily Promeropinae, Sugar-bird, (1)<br />
Family Zosteropidae, White-eyes, 80<br />
VIREOS, FINCHES, AND ALLIES<br />
Family Vireonidae, Vireos, 41<br />
Family Drepaniidae, Hawaiian Honeycreepers, 22<br />
Family Thraupidae, Tanagers, etc., 474<br />
Subfamily Parulinae, Wood Warblers, (109)
38 AMERICAN MUSEUM <strong>NOVITATES</strong> NO. 1496<br />
Subfamily Coerebinae, Honeycreepers, (36)<br />
Subfamily Catamblyrhynchinae, Plush-capped Finch, (1)<br />
Subfamily Thraupinae, Tanagers, (196)<br />
Subfamily Pyrrhuloxiinae, Cardinal Grosbeaks, (132)<br />
Family Tersinidae, Swallow-tanager, 1<br />
Family Fringillidae, Typical Finches, 293<br />
Subfamily Emberizinae, Buntings and <strong>American</strong> Sparrows, (171)<br />
Subfamily Fringillinae, Chaffinches, Linnets, and Allies, (122)<br />
Family Icteridae, Troupials, <strong>American</strong> Blackbirds, and Allies, 88<br />
WEAVERBIRDS, STARLINGS, AND ALLIES<br />
Family Ploceidae, Weaverbirds, 263<br />
Subfamily Bubalornithinae, Buffalo-weavers, (3)<br />
Subfamily Passerinae, Sparrow-weavers, (35)<br />
Subfamily Ploceinae, Typical Weavers, (109)<br />
Subfamily Estrildinae, Waxbills and Allies, (107)<br />
Subfamily Viduinae, Widow-birds, (9)<br />
Family Sturnidae, Starlings, 103<br />
Subfamily Sturninae, Starlings, (101)<br />
Subfamily Buphaginae, Tick-birds, (2)<br />
Family Oriolidae, Old World Orioles, 34<br />
Family Dicruridae, Drongos, 20<br />
CROWS AND AUSTRALIAN CROW-LIKE BIRDS<br />
Family Corvidae, Crows, Jays, Magpies, 100<br />
Family Cracticidae, Bell Magpies, etc., 11<br />
Family Grallinidae, Magpie-larks, etc., 4<br />
Subfamily Grallinae, Magpie-larks, (2)<br />
Subfamily Corcoracinae, White-winged Chough; Gray Jumper, (2)<br />
Family Callaeidae, Wattlebirds, 3<br />
Family Ptilonorhynchidae, Bowerbirds, 17<br />
Family Paradisaeidae, Birds <strong>of</strong> Paradise, 43<br />
LITERATURE CITED<br />
AMADON, DEAN<br />
1943. The genera <strong>of</strong> starlings and their relationships. Amer. Mus. Novitates,<br />
no. 1247, 16 pp.<br />
1944. The genera <strong>of</strong> Corvidae and their relationships. Ibid., no. 1251, 21 pp.<br />
1950a. Australian mud nest builders. Emu, vol. 50, pp. 123-127.<br />
1950b. The Hawaiian honeycreepers (Aves, Drepaniidae). Bull. Amer. Mus<br />
Nat. Hist., vol. 95, pp. 151-262, 23 figs., pls. 9-15, 15 tables.<br />
[In press.] Le pseudo-souimanga de Madagascar (Neodrepanis). L'Oiseau.<br />
[In press.] Taxonomic notes on the Australian butcher-birds. Amer. Mus.<br />
Novitates.<br />
ARVEY, M. DALE<br />
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1951 A CLASSIFICATION O RECENT BIRDS 39<br />
BEECHER, WILLIAM J.<br />
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