A PHYLOGENETIC

ANALYSIS

OF RECENT

ANSERIFORM MORPHOLOGICAL

GENERA

USING

CHARACTERS

BRADLEY C. LIVEZEY

Museum NaturalHistoryandDepartment Systematics Ecology, of of and University Kansas, of

Lawrence, Kansas 66045 USA

ABSTRACT.--A phylogeneticanalysisof all Recentgeneraof the Anseriformes using 120 morphologicalcharacters supportsmuch of the current consensus regarding intraordinal relationships. found that (1) Anseranas I shouldbe placedin a monotypicfamily; (2) Dendrocygna,Thalassornis, geeseand swans,and Stictonetta paraphyleticto the rest of the Anatiare dae;(3) Cereopsis the sistergroup to Anserand Branta, is and Coscorobathe sistergroup to is Cygnus and Olor;(4) Plectropterus the sistergroup to the Tadorninae(shelducks) is and the Anatinae (typical ducks);(5) the shelducks monophyleticand include Sarkidiornis are (provisionally), Malacorhynchus, Hymenolaimus, Merganetta, and Tachyeres; the tribe "Cairinini" (6) ("perchingducks")is an unnatural,polyphyleticassemblage is rejected; the dabbling and (7) ducks(includingthe smaller"perchingducks")comprise unresolved, an probablyparaphyletic group;(8) tribal monophylyof the pochards (including Marmaronetta Rhodonessa), and seaducks(including the eiders),and stiff-tailedducks(including Heteronetta) confirmed; is and (9) the retention of Mergellus and resurrectionof Noraonyx recommendedbasedon are clarifications intratribal relationships. of Problematic groups,effects homoplasy, of phenetic comparisons, life-historycorrelates, biogeographic patterns,and fossilspecies discussed, are and a phylogeneticclassification Recentgenera is proposed. of Received November 18 1985,
accepted April 1986. 2
THE order Anseriformes is considered to

1962; Davies and Frith 1964; Raikow 1971; Kear

comprisethe families Anhimidae (2 genera, 3 species) and Anatidae (approximately43 genera and 150 species). The family Anatidae is undoubtedlyone of the best-studied groupsof birds, owing largely to the historical importanceof waterfowl for hunting (Weller 1964a),
domestication (Delacour 1964a), and aviculture (Delacour 1964b).

and Murton 1973). Most authors assumed the

validity of the tribesand usedthem asworking units in phylogenetic analysesof the family (e.g. Johnsgard 1961a, Bottjer 1983). A few workers named additional tribes (Moynihan
1958, Delacour 1959, Woolfenden 1961, Weller

1968b) or attempted to test the naturalnessof those originally proposed(Cotter 1957, Wool-

The classification the Anatidae proposed fenden 1961, Brush 1976). of Behavioral characters have been accorded by Delacour and Mayr (1945) has been followed, with only minor revisions,in recent de- considerable weight in classifications waterof cades (e.g. Delacour 1954, 1956, 1959, 1964c; fowl. Delacour and Mayr (1945) based their Johnsgard 1961a, 1962, 1965a, 1978, 1979; revision on characters they consideredto be Woolfenden 1961; Frith 1967; Bellrose 1976; "non-adaptive," including behavioraldisplays, Palmer 1976; A.O.U. 1983; Bottjer 1983; Scott nesting and feeding habits, and selected mor1985). Perhaps the most innovative aspect of phological characters(e.g. posture, body prothis system (inspiredby the works of Salvadori portions,headshape,syringealbulla). Reliance 1895;Phillips 1922, 1923, 1925;and Peters 1931) on comparative ethology in anatid systematics was the erection of "tribes," groups of genera was furthered by the studiesof Lorenz (1951that were considered be closelyrelatedwith- 1953), McKinney (1953), and Myres (1959) and to in the subfamilies of the Anatidae. These tribes was increased significantly by Johnsgard became primary focusof subsequent the works (1960a-c, 1961a-d, 1962, 1964, 1965a, b, 1966a, on anatid classification, many of which ad- b, 1967, 1978), whose work was largely ethodressedthe tribal assignments problematic logical and influenced profoundly by that of of genera (e.g. Humphrey and Butsch 1958; Delacour (1954, 1956, 1959, 1964c). This emJohnsgard 1960a,1961b;Humphrey and Ripley phasis, work on interspecific hybridization
737 The Auk 103: 737-754. October 1986

738

BRADLEY LIVEZEY C.

[Auk, Vol. 103

(= Metopiana(= Netta), Oidemia(= (Sibley 1957;Gray 1958;Johnsgard 1960d,1963), Casarca Tadorna), (= For and studyof plumagepatterns downy young Melanitta), and Charitonetta Bucephala). Anas, of
speciesfrom several subgenerawere examined: Mallard (Anasplatyrhynchos), Northern Pintail (A. acuta), American Wigeon (A. americana), Green-winged Teal (A. crecca), Northern Shoveler(A. clypeata). and Othcollections. er species Anaswere studied for certain characters. of Other data used in the classification of waSalvadori'sDuck [Anas(Salvadorina) waigiuensis], proterfowl includesyringealanatomy(Humphrey visionally assignedto Anas but consideredproblem1955,1958;Johnsgard 1961e),cytogenetics (Ya- atic by some(Mayr 1931,Kear 1975),was not includmashina 1952), serology (Cotter 1957, Bottjer ed because no skeletal specimens were available (monotypic, 1983), osteology(DeMay 1940, Verheyen 1955, (Wood et al. 1982).Exceptfor Rhodonessa Humphrey and Butsch1958,Woolfenden 1961, probably extinct; one completeskeleton)and Camp(monotypic, extinct; castsof two partial Humphrey and Ripley 1962, Raikow 1971), torhynchus skeletons),all genera analyzed were representedby feather lice (Timmermann 1963), eggshell at least two complete skeletons. For all polytypic structure (Tyler 1964), egg-white proteins (Sibgenera at least two specieswere studied, and a numley 1960, Sibley and Ahlquist 1972), feather ber of the common, diverse, or problematic genera proteins(Brush1976),royology(Zusi and Bentz were representedby large series. 1978), lipids from the uropygial gland (Jacob For Camptorhynchus, character states for unavailand Glaser 1975, Jacob 1982), and mitochon- able elementseither were assumed provisionally(for

(Delacour 1954, 1956, 1959; Frith 1955, 1964b;

Kear 1967) were prompted in part by the opportunity to observe waterfowl in avicultural

drial DNA (Kessler and Avise 1984).

characters invariant within the anatines) or coded as

Howard (1929). Some characterswere rejected becausevariation prevented even modal state assignmentsfor somegeneraor because discretestatescould Woolfenden 1961). I performed a phylogenetic (cladistic) anal- not be distinguished. Sourcesfor data on the postcranialskeleton were ysis of Recent genera of Anseriformes using Wetmore (1951), Rand (1954), Verheyen (1955), 120 morphologicalcharacters. presenta hy- Woolfenden(1961),Humphrey and Clark (1964),and I pothetical evolutionary tree for the order, con- Raikow (1971). Additional sources were (by anatomsider the taxonomic implications, and discuss ical region): integument and molt (DeMay 1940, selectedlife-history and biogeographiccorre- Siegfried 1970, Palmer 1976), trachea and syrinx
lates and the classification of selected fossil

These studies,with the possibleexceptions of thoseby Lorenz (1953) and Kesslerand Avise (1984),estimatedthe evolutionaryrelationships of groups by assessments overall simof ilarities; no attempts were made to determine primitive conditions or to distinguish shared primitive charactersfrom shared derived characters ("special" similarity). Moreover, the "evolutionary trees"presentedin mostof these works lack references the specificcharacters to used to support the branching patterns (e.g. Delacourand Mayr 1945;Johnsgard 1961a,1978;

"missing." Assumption of anatine characters for Camptorhynchus conservative (cf. Humphrey and is Butsch 1958, Zusi and Bentz 1978) and did not alter its positionin the resultanttree (comparedwith analyses without this assumption),but permitted more efficient computation of trees and a shorter final solution.

Analysis characters.--For phylogenetic analof the ysis presented,120 characters were used (Appendix 1); a majority of the osteological characters were described in Woolfenden (1961) and illustrated in

(Wetmore 1926; Niethammer 1952; Wolff and Wolff

species. Many of the characters were described 1952; Humphrey 1955, 1958, unpubl. data; Humfirst in the pioneering work of Woolfenden phrey and Butsch 1958; Johnsgard1961b, e; Humphrey and Ripley 1962; Humphrey and Clark 1964; (1961), to whom I dedicatethis paper.
METHODS

Weller 1968b), and skull (Abbott 1938, Harrison 1958, Raikow 1970a, Olson and Feduccia 1980a). I included

Taxaandspecimens.--Both generaof Anhimidae and all Recent genera of Anatidae were studied. I analyzed separately severalsubgenera (sometimes considered genera), including Olor, Lophonetta, Pteronetta, Amazonetta, Callonetta, Mergellus,Lophodytes, and Nomonyx. Several other "subgenera"were found to
be identical to the taxa with which they generally are merged and are not labeled separatelyin the trees:

only qualitative charactersbecausethe polarities and statesof mensural charactersare especially difficult
to determine.
which fined.

Each character is an anatomical

character

trait for

two or more discrete

states were de-

Derivationof trees.--Polarities of each character (primitive states) were determined by comparison with outgroups--Burhinus and Larus (Charadriiformes), Ortalis and Meleagris(Galliformes), Ciconia

October 1986]

Phylogeny Anseriformes ofthe

739

(Ciconiiformes),and Phoenicopterus (Ciconiiformesor Charadriiformes)--each of which has been proposed as closelyrelated to the Anseriformes(Delacourand Mayr 1945;Delacour 1954;Mainardi 1962;Simonetta 1963; Sibley et al. 1969; Sibley and Ahlquist 1972;
Brush 1976; Feduccia 1977, 1978; Olson and Feduccia

1980a,b; Olson 1985).Generally, the galliformswere most useful for establishing polarities. Outgroups were usedto constructa hypothetical ancestor(a vector of primitive character states) for the Anseriformes,which was usedto root the evolutionarytree; the primitive condition of nine characterscould not be determined and were codedasmissing(Appendix
1). Transformation series were treated as linear un-

tropterus (Peters1931,Delacourand Mayr 1945) and the proposition that Anseranas an aberis rant "true goose"(Daviesand Frith 1964,Frith 1967). Both the anhimids and Anseranas have undergone substantialautapomorphicchange sincedivergence,much of which is unique in
the order.

Geese, swans, and proto-ducks.--! found that the "Anserinae," as currently defined (e.g. Johnsgard 1978), is paraphyleticto the rest of the family (Fig. 2), in contrast the monophyto ly depictedby Delacourand Mayr (1945),Boetticher (1952), and Woolfenden (1961). Johnsgard (1961a, e) depicted the group as

less they appearedto be nonlinear or problematic; the latter were treated as unordered (Appendix 1). The syringealbulla was given a weight of 2 because it is a locomotion-independentcharactercomplex involving enlargement, symmetry, and fenestration; all other characters were assignedunit weight. The logic and terminology of phylogenetic analysis are discussed Wiley (1981). The tree was dein rived using the PAUP program (Swofford 1984), a program that seekstreesof maximumparsimony(i.e. requires the least number of character-state changes; seeKluge 1984) and that permits the examinationof seriesof "equally short" trees.The large size of the data set prohibited an exhaustivesearchguaranteed to find the shortest tree(s), but two thorough methods-alternate and global branch swapping--were employed and producedidentical topologies. A data matrix for all outgroups and the Anseriformesand a list of specimensexamined are available from the author on request.
RESULTS

paraphyleticto the "Anatinae,"but later (1978) diagrammed asmonophyletic, did Bellrose it as (1976). These variations, however, may reflect
different approaches to tree construction as

muchaschangingperceptions relationships. of The branchingsequence (Fig. 2) differsfrom conventional schemes(Johnsgard1967, 1978;

Kear 1967; Raikow 1971; Brush 1976) in that

Dendrocygna Thalassornis not sistergenand are era but insteadcomprisea grade. !n an equally parsimonioustopology, Thalassornis diverged immediatelyafter the goose-swan branch.Most of the 13 autapomorphies Thalassornis in representadaptationsfor diving and include several convergences with diving ducksin other clades. Until the works of Johnsgard (1967) and
Raikow (1971), Thalassornis was considered to
be an aberrant stiff-tailed duck and allied with

Oxyura (e.g. Peters 1931; Delacour and Mayr

1945; Delacour 1959, 1964c), a treatment re-

General findings.--Of50 equally short trees peatedrecently (Howard and Moore 1984,Scott
examined that resulted from minor changesin characterdistributions, only three distinct to1985).

Ten characterchangesconfirmed the mono-

pologieswere found. The tree illustrated (Fig. 1) has the topology of 46 of thesetrees (consistency index = 0.59). Two treesreversedthe order of branchingof Thalassornis the geese with
and swans, and two others altered relation-

phyly of the geeseand swans(Fig. 2), as hypothesizedby most authorities in recent decades (Delacourand Mayr 1945;Delacour1954; Johnsgard1961a,e, 1965a, 1978; Woolfenden

ships in the goldeneye-merganser clade (Bucephala, Mergellus, Lophodytes, Mergus). Anhimids Anseranas.--Monophylyof the and Anhimidae and the waterfowl and the early branchingof Anseranas were confirmed(Fig. 2). The "primitive" status of Anseranas has been recognized widely (Miller 1919; Boetticher 1943; Delacour 1954;Johnsgard 1961c,e, 1962, 1978,

1961;Bottjer 1983).Also, there is a sister-group relationship between Coscoroba the "typiand cal" swans(Cygnus, Olor; 6 characters) and between Cereopsis the "typical" geese(Anser, and Branta; characters). analysisdemonstrated 2 My monophylyof Olor,but no apomorphies distinguishedCygnus from the commonancestorof Cygnus and Olor (Fig. 2); hence, the topology of Cygnusremains unresolved. The generic 1979; Woolfenden 1961; Olson and Feduccia monophyly of Anserand Brantaalso was not and Cer1980a).Notableexceptions were the placement established.Traditionally, Coscoroba havebeenconsidered be either "links" to of Anseranas with the superficiallysimilar Plec- eopsis

740
ANCESTOR Chauna

BRADLEY C.LIVEZE
PART 1

[Auk, 103 Vol.

Anhima
Anseranas

Dendrocygna
Thalassornis

Cygnus
Coscoroba

Anser
Branta

Cereopsis

SarkidiOFnis
TadoFna

Malacorhynchus Chloephaga Alopochen

Cyanochen Tachyeres
Merganetta
aimus Pteronetta PART

Lophonetta Nettapus
Anas

IGallonefta
Chenonetta Amazonetta

MaFmaronetta Rhodonessa
Netta

Somateria

PART 4

Camptorhynchus
Melanitta

. Clangula Mergellus

Lophodytes Mergus
Nomonyx
Biziura

Fig. 1. Phylogenetic of Recent tree anseriform genera selected and subgenera based 120morphological on characters listedin Appendix1. Lengths horizontal of linescorrespond the numberof character to changes (apomorphies) the lineages. in Sections the tree are detailedin Figs.2-5. of

between tribes, early branchesfrom the com- rived from an earlier "anserine" branch of the mon ancestorto the "true" geese and swans, Anatidae (Verheyen 1955; Johnsgard 1960c,
or, for Cereopsis, aberrant shelduck or the an
1961a,b, 1962, 1965a, b, 1978; Woolfenden 1961;

sole member of a separatetribe (Peters 1931; Frith 1964a, b, 1967; Brush 1976; Olson and Delacourand Mayr 1945;Delacour 1954, 1964c; Feduccia 1980a). Johnsgard 1961a, e, 1978; Woolfenden 1961; Plectropterus theshelducks.--I and found that Frith 1967;Kearand Murton 1973;Bottjer1983). Plectropterus not related closely to the is My analysisshowed Stictonetta be the last "perchingducks"(e.g. Sarkidiornis, to Cairina, and branch in the grade of waterfowl with reticu- Nettapus) generally recognizedsince Delaas late tarsi (Figs. I and 2). Stictonetta long was cour and Mayr (1945). Instead, Plectropterus is
believed to be an aberrant member of the
the earliest branch of the waterfowl with scu-

shelducks (near Tadorna; Peters 1931, Boettich-

er 1952) or the dabbling ducks(e.g. Anas;Delacour and Mayr 1945; Delacour 1956, 1964c). Basedon anatomicalcomparisons, however, otherworkers suggested the genuswasdethat

tellate tarsi, and lacks five synapomorphies uniting more-derivedmembersof the family
(Fig. 3). Woolfenden (1961) concluded that

Plectropterus mostsimilarosteologically was to the shelducks, Tyler (I 964)found that eggand

October 1986]

Phylogeny Anseriformes ofthe

741

106 a-b
I05 b-c

85

a-b

33
79 525564-

a-b
c -d

23

a-b

65
78-

43

a-b

38 a-b +
74 a-b

b-c

45 63

- a-b - a-b

68
72 77 78 79
89

- a-b
- a-b - a-b b-a

53 58 m J= 70

a-b a -b a -b

92 93 96 99

I09

105

a-c

17

a-b

94

a-b

I0
31

a -b
a -b

86 I a-b
4
9

a-b

120

o -b
a -b

40
42 50 91

a-b
a-b a -b a -b

12
18 88 ] O0 ]16

a-b
a-b a-b a -b a-b

Fig. 2. Detailed diagramof Part 1 of the phylogenetictree of the Anseriformesshown in Fig. 1. Characters are listed in Appendix I.

shells of Plectropterus were intermediate between those of anserines and shelducks in

volving Tadorna, Malacorhynchus, two clades and containing three genera each (Fig. 3). I found structure.Bottjer(1983) found that Plectropterus Malacorhynchus be a highly derived shelto differed greatly from other "perching ducks" duck; this genus has been assignedmost freserologically attributed the result to exper- quently to the dabbling ducks (e.g. Delacour but imental error. 1956,Woolfenden 1961,Frith 1967,Johnsgard My analysissupportsthe monophyly of the 1978),although Frith (1955) noted that the patshelducks, although only by a single, possibly tern of its downy young differed greatly from convergentcharacter; further study may show those of Anas and Brush (1976) found that Mathe shelducks be a grade of relatively prim- lacorhynchus distinctlydifferent from Anas to was itive ducks.The clade includesa polytomy in- in its feather proteins.A single synapomorphy

742

BRDI,Y LZVEZEY C.

[Auk,Vol. 103

....

Plectropterus
I
v

Sarkidiornis
Tadorna

: I ', ', : ; Malacorhynchus

FoNeochen .
- L Alopochen

Chloephaa

IICyanochen
' [llllllllll
Tachyeres
..... Merganetta

Hymenolafmus

TOFIGURE4

Fig.3. Detailed diagram Part2 of the phylogenetic of the Anseriformes of tree shown Fig. 1. Characters in are listedin Appendix1. Graphical proximityof branches within the polytomyin the shelducks doesnot
reflect relatedness.

of the skull supportsthe monophyly of the diving ducksin other clades(see Discussion). three genera of "sheldgeese"--Alopochen, Hymenolaimus Merganetta and have been treatChloephaga, Neochen and (Fig. 3). ed as allied either with the shelducks,"perchThe Comb Duck (Sarkidiornis melanotos), tra- ing ducks,"or as exceptional, possiblyprimiditionally placed in the "Cairinini" (Delacour tive dabbling ducks (Delacour and Mayr 1945, and Mayr 1945,Johnsgard 1978),appears be 1946; Delacour 1956; Ripley 1957; Johnsgard to an early branch of the shelducks. However, 1965a,1966a).SomeworkersplacedMerganetta three humeral characters that were important in its own tribe, Merganettini (Woolfenden in distinguishing the "anserines" and shel- 1961; Kear and Steel 1971; Kear 1972, 1975; ducksfrom the "anatines"[capitalshaft ridge Brush1976;Johnsgard 1978).My analysis does (character 22), deltoid crest (25), and external not supportthe suggestion (Olson and Feductuberosity (32)] were of equivocal or "inter- cia 1980a:22) that "... the typical membersof mediate"conditionin this species. addition, the 'subfamily'Anserinaeand the typical memIn the enlarged,uniquely distally directed meta- bers of the 'subfamily' Anatinae are more carpal I of Sarkidiornis, character complex closelyrelatedto one anotherthan to Stictoneta herein considered incorporatetwo aspects ta, Malacorhynchus, Merganetta." to or The third and most derived member of this [orientation(41) and length (42)], proved difficult to characterize. Perhaps Sarkidiornis branchedimmediatelybefore Cairinaor, alternatively, after Plectropterus before the dibut vergence of the shelducks from other "anatines."

clade is Tachyeres (Fig. 3), a neotropical genus

generallyplacedin the shelducks in a sepor

arate tribe allied with the shelducks (Delacour

1954;Moynihan 1958;Johnsgard1965a, 1978; Weller 1976;but see Ripley 1957, Woolfenden Three problematic genera--Hymenolaimus, 1961).Like all shelducks, Merganetta Tachyand Merganetta, and Tachyeres--comprise highly eres show an enlargement of metacarpal I a derived cladeof shelducks (Figs.1 and 3). Sev- (adorned with keratinspursin Merganetta) that

eral of the characters uniting thesegeneraare evidently related to diving and are sharedby

is sexuallydimorphicand age related (Weller 1968a,Livezeyunpubl. data).Cyanochen hyis

October 1986]

Phylogeny Anseriformes ofthe

743

pothesizedto be the sistergenusto the Hymenolaimus-Merganetta-Tachyeres clade, although this relationshipis supportedby only two synapomorphies. Bottjer (1983) suggested that Cyanochen may have branchedbefore the other
shelducks.

28
55 56

a -b

"Perching"ducksand "dabbling" ducks.--The sister group to the shelduckscomprisesfour groups (Fig. 1): a poorly resolved grade of "perching" and "dabbling" genera (Pteronetta, Cairina, Aix, Lophonetta, Nettapus, Anas,Callonetta, Chenonetta,and Amazonetta), which in turn

33

b-c

gave rise to the pochardsand independently to

the sea ducks and stiff-tailed ducks. The first

97 o-b

group (Fig. 4), henceforth termed "dabbling ducks," is a paraphyletic group of genera previously allocatedto either the "Anatini" or the
"Cairinini" (Delacour and Mayr 1945;Delacour
1978). Fig. 4. Detailed diagramof Part 3 of the phyloCharacters listed in Appendix 1. Graphical proxare

1956;Johnsgard 1960c,d, 1961a,e, 1962,1965a, genetic tree of the Anseriformesshown in Fig. 1.

within the polytomyin the dabThe polyphyleticcharacter the "Cairinini" imity of branches of was inferred by Woolfenden (1961). The tribe bling ducksdoesnotreflectrelatedness. hasbeen recognizedby subsequent workers in spiteof the equivocal allocationof severalgenera (e.g. Callonetta and Amazonetta; Johnsgard

1960a, 1965a, 1978), the widely recognized heterogeneity of its members in behavior, morphology, and biochemistry(Johnsgard1960c, 1961a,1962, 1965a,1978;Woolfenden 1961;Tyler 1964; Brush 1976; Bottler 1983), the lower incidenceof interspecifichybridization within
the tribe than between its members and those

Anas, Callonetta, Chenonetta,Amazonetta, and the

of other tribes (Johnsgard1960d), and the conspicuouslack of a single character(or combination of characters) that uniquely distinguishes its members from other anatines. Johnsgard (1965a, 1978) admitted that retention of the tribe was partly a taxonomic convenienceto avoid creationof "a comparativelylarge tribe" (1978: xxi) and omitted it as a suprageneric taxon in his latest list (Johnsgard1979). The genera of dabbling ducks (on the basis of three variable multistate characters) form a grade from relatively primitive (e.g. Cairina, Lo-

sea ducks and stiff-tailed ducks (Fig. 4), suggestingthat the pochardsaroseindependently of other diving ducks.I found that Rhodonessa is the sistergroup to Netta and Aythya, which agrees with most studies (Verheyen 1955; Johnsgard 1961a,e, 1962, 1978, 1979;Woolfenden 1961; Humphrey and Ripley 1962; Brush 1976) sinceDelacour and Mayr (1945, 1946) and Delacour(1956) provisionally placedRhodonessain the dabbling ducks.Marmaronetta, genus a believed to "link" the Anatini with the pochards but retained within the Anatini (Johns-

gard 1961a, b, e, 1978; Delacour 1964c;Brush 1976), is supported in my study as the sister genusto the pochardsby two osteological synapomorphies (Fig. 4). This relationship is corroborated the secondary by lossof metallic coloration in the speculum (Delacour and Mayr

phonetta) more-derivedforms(e.g. Anas, to Cal- 1946). Seaducks.--The sea ducks comprise a monolonetta). A single osteologicalsynapomorphy supportsa close relationship between Cairina phyletic group relatedto the stiff-tailedducks, and Aix (Fig. 4), a relationship suggestedpre- although this relationshipis supportedby only viously by karyotypic and serological compar- a few, possibly convergent charactersrelated
isons (Yamashina 1952, Cotter 1957, Bottjer
1983).

to diving (Fig. 5). My result contradictsthe previouslyproposed closerelationshipbetweensea Pochards.--Thepochardsare a monophyletic ducks and "perching ducks" (Delacour and groupin an unresolved polytomythat involves Mayr 1945, Delacour 1959, Bottler 1983). With

744

BRADLEYLIVEZEY C.
main tentative because

[Auk,Vol. 103
of the limited material

available. Humphrey and Butsch(1958) placed Camptorhynchus Melanitta but before Clanafter gula,and Zusi and Bentz (1978) allied the genus with eiders, evidently on the basisof shared primitive characters.The very close relationship of goldeneyesand mergansershas had unanimous support in recent decades(e.g. Delacour and Mayr 1945; Boetticher 1952; Humphrey 1955; Delacour 1959; Myres 1959; Johns== ss o-b ,,,.-gard 1960b, d, 1961a, 1978; Brush 1976). This analysisshowsthe Sinew (Mergellus albellus) be either the sistergenusto Bucephala to (Fig. 5) or the sister group to the LophodytesMergusclade. The former topologyis corroborated by the relatively high frequency of Mergellusx Bucephala hybridsin the wild (Phillips 1925,Ball 1934,Gray 1958,Nilsson 1974,Johnsgard 1978). Previous workers either listed the Sinew between the goldeneyes(Bucephala) and the mergansers (Mergus, Lophodytes) a monoas typic genus (Peters 1931, Woolfenden 1961, Fig. 5. Detailed diagramof Part 4 of the phylo- A.O.U. 1983), or merged it (with Lophodytes) genetictree of the Anseriformes shownin Fig. 1. into Mergus(Delacourand Mayr 1945;BoettichCharacters listed in Appendix 1. Placementof er 1952;Humphrey 1955;Delacour 1959, 1964c; are Camptorhynchustentative. is Johnsgard1960c, 1961a, d, 1965a, 1978, 1979). Stiff-tailedducks.--My study supports the monophyly of the stiff-tailed ducks, wherein the possible exception of a few workers who Heteronetta the sister genus to the more typis advocated(largely on the retention of primi- ical members(Fig. 5). The position of the clade tive Anas-like characters)the tribal separation as closely related to the sea ducks and highly of the eiders(Somateria Polysticta) and from the derived (especiallyBiziura)agreeswith recent other sea ducks (Humphrey 1955, 1958; Dela- orderingsof generaby taxonomists (e.g.Johnscour 1959; Brush 1976; Todd 1979), the mono- gard 1979) but disagreeswith suggestionsof a phyly of the seaduckshas not been questioned pre-dabbling duck (Raikow 1970b, Johnsgard recently (e.g. Johnsgard1960b, 1961a, e, 1964, 1978)or pre-shelduck(Johnsgard 1965b,Bottjer 1983) origin for the group. A few workers have 1978;Woolfenden 1961;Bottjer 1983). The proposed sequences generawithin the expressed doubts about the relationships of of grouphasvaried (e.g. Delacourand Mayr 1945; Heteronetta(Johnsgard 1960c, Brush 1976), and Delacour 1959;Johnsgard1960b, 1961a, 1965a, others accepted the relationship between Het1978, 1979).My analysis(Fig. 5) indicatesthat eronettaand other stiff-tailed ducks but sugPolysticta, Somateria, Histrionicus, and Campto- gestedthat the genus be accordedtribal rank rhynchus comprise lessspecialized, a basalgrade (Weller 1967, 1968b; Rees and Hillgarth 1984). of genera retaining primitive, unfenestrated My analysis shows that the Masked Duck syringealbullae. This seriesof generagivesrise [Nomonyx (Oxyura)dominica] the sistergroup is to a well-supported clade of, in order of in- to the highly derived Oxyura-Biziura clade (Fig. creasing relatedness, Melanitta,Clangula, the 5), i.e. Oxyurais relatedmore closelyto Biziura and goldeneye-merganser clade. The eiders (Poly- than to the very similar Nomonyx.This topolsticta,Somateria) appear to be paraphyleticto ogy is supportedas well by the derived lossof the other seaducks;this paraphyly is support- a speculum in Oxyura and Biziura,which is reed only weakly, and downy patterns suggest tained in Nomonyx(Delacour 1959). This findthat the eidersmay be monophyletic(seeDis- ing supports the resurrection of Nomonyx cussion). Placementof Camptorhynchus re- as advocatedby Woolfenden (1961), a recommust

October 1986]

Phylogeny theAnseriformes of

745

mendationrejected Delacour(1964c),Johnsby gard (1967), and most subsequent workers. I did not examineskeletons all species Oxyof of ura, so monophyly of the genus was not establishedwith certainty.
DISCUSSION

osteological evidence,appear monophyletic in their dusky undersides,obsoletedorsal spots, and dark cheeks;(2) other Mergini lack the supraorbitalstripe and dorsalspottingof the dabbling ducks(loral spot and vestigial back spots
retained in Histrionicus); (3) Melanitta and its

sister genera are synapomorphicin their dark breast bands, a character secondarily lost in Diving habit and homoplasy.--Considerable Mergus; the BlackScoter(Melanittanigra)and (4) homoplasy (convergence) characters shown Surf Scoter (M. perspicillata) united by the of is are in the tree (Figs.1-5) and by the consistencies derived darkening of the lower breastand belof characters (Appendix 1). The majorityof the ly; (5) Bucephala, Mergeflus, Lophodytes, Merand convergencesare associatedwith adaptations gussharea reversalin (presence dorsalspotof) for diving, and most involve the leg elements ting; (6) obscured (Lophodytes) dark (Mergus) to (characters 55, 56, 64, 65, 69, 75), pelvis (119), cheeksunite the mergansers; 52, and (7) Mergusis and skeletal pneumaticity(28, 78). These fea- derived further in the anteriorly incomplete tures tend to co-occur,especiallywithin ele- breastband and pale suborbitalstripe. ments. Convergence between Thalassornis and Similarityvs. relatedness.--Recognition the of the stiff-tailed ducks is particularly pervasive different types of characterchange is important (Figs. 2 and 5). It appears,however, that the in light of the conspicuously unequal rates of moderately large number of characters includ- morphologicalevolution in different lineages, ed in this analysisreduced the impact of such e.g. autapomorphies Brantavs. Cereopsis of (Fig. homoplasy on the resultant tree, although 2) and Oxyura vs. Biziura(Fig. 5). The inadedeletionsor heavy weighting of selectedchar- quacy of simple distancetechniqueswas demacterscanproducetopological changes. ex- onstratedusing thesedata through a compariFor ample, becauseof a number of diving-related son of phylogenetic relationshipswith "path homoplasies, postulation of the Hymenolaimus- lengths" or patristic distances. Selectedresults Merganetta-Tachyeres as the sister group were: (1) Anseranas roughly equidistantfrom clade is

to the seaducksand stiff-tailedducksis only the anhimids and other anatids [corroborated slightly lessparsimonious than the topology immunologically Bottjer(1983)],but is the by presented (Fig. 1). Similarly, heavierweighting sistergroup to the latter (Fig. 2); (2) Thalassornis
of appendicularcharacters placesHeteronetta as is most similar to Dendrocygna but is more the sistergroup to both the seaducksand other closelyrelated to other anatids,excludingAnstiff-tailed ducks. seranas (Fig. 2); (3) Heteronetta appears"nearer" Patterns downy of young.--Althoughan anal- to Anasthan to Oxyura,a member of the sister ysis of downy patterns for the entire order is group of Heteronetta (Figs.4 and 5); and (4) benot possibleat present,a cladisticreevaluation causeof autapomorphiesin Oxyura, Biziurais of the downy young illustrated in Delacour phenetically "closer" to Nomonyxthan it is to (1954, 1956, 1959) permits an independent test its sistergenus Oxyura(Fig. 5). of two parts of my phylogenetic hypothesis Life-history correlates.--Diving,at least as an behavior, occursthroughout the order (using Anas the outgroup). as Patterns downy escape in stiff-tailed ducksagree well, wherein (1) Het- exceptin the anhimids and possiblyAnseranas eronetta retains virtually all dabbling-duck (Johnsgard 1962, Todd 1979). Groupsthat roucharacters;(2) Nomonyx, Oxyura, and Biziura tinely dive for food are fewer, but occur in six sharea synapomorphic, dark cheek stripe; (3) lineagesthroughoutthe family (Weller 1964b): Oxyura and Biziuraare united by the derived Dendrocygna, Thalassornis, Hymenolaimus-Merlossof the pale supraorbitalstripe;and (4) Bi- ganetta-Tachyeres, pochards, ducks,and stiffsea ziurasharesa lossof dorsaland wing spotting tailed ducks. with the Peruvian Ruddy Duck (O. ferruginea) Perchinghabit, the probablyprimitive charand dark cheeks with the Australian Blue-billed

Duck (O. australis). Patternsof downy Mergini

also are informative: (1) eiders retain the su-

acterusedtraditionallyto define the polyphyletic "perchingducks,"occurs many genera, in includingAnseranas, Dendrocygna, Plectropterus,
Sarkidiornis, Tadorna, Malacorhynchus, Cairina, Aix,

praorbital stripe of dabblersbut, in contrastto

746

BRADLEY LIVEZEY C.

[Auk, Vol. 103

Mayr 1945:51), most previous biogeographers 1960,Johnsgard 1978,Todd 1979).A relatedtrait contendedthat the Anseriformesoriginated in (alsousedto justify the "Cairinini"), nestingin the Northern Hemisphere,probably the Paletree cavities, occurs in Dendrocygna, some arctic (Howard 1950, Weller 1964d). Although shelducks(Sarkidiornis, Neochen, Alopochen, Ta- the fossilrecordof waterfowlis morecomplete dorna,Malacorhynchus),number of dabbling for the Northern Hemisphere (Howard 1964), a ducks(e.g.Aix, someAnas),and someseaducks early forms are known from both hemispheres, (Histrionicus, Bucephala, Mergellus,Lophodytes, and the apparent disparity in representation Mergus). probably reflects intensity of paleontological Use of terrestrial cavitiesfor nesting also oc- research.The only essentiallynorthern groups Cyanochen, Rhodonessa, cursin someshelducks (Tadorna, Hymenolaimus,are Olor, Anser-Branta, Merganetta, Tachyeres), someAnas,and in the and the sea ducks;genera that have roughly in sea ducks cited above (Hobbs 1957, Warham equal distributions in both hemispheresare 1959, Johnsgard 1962, Johnson 1963, Weller Cygnus, Tadorna, Alopochen, Anas,Netta,Aythya, 1964c,Kear 1970,Moffett 1970,Humphrey and and Oxyura.The remaining 20 genera are limin Livezey 1985).Other species nest on the ground ited to or mostspeciose the SouthernHemior over water (Weller 1964c,Kear 1970). Only sphere,and, with the anhimids, include most the Black-headed Duck (Heteronetta atricapilla) of the early branches in the order. Furthermore, the earliestbranches(Fig. 2) in the swan is an obligate nest parasite, although infrequent nest parasitismoccursin a number of and goose clades are genera limited to the other genera including Dendrocygna, Branta, SouthernHemisphere.The numerousholarctic Anas, Aythya,and Mergus (Weller 1959,1968b). speciesof Anas, Aythya, and the sea ducks Although ground nesting appearsto be prim- (Weller 1964d) may represent radiations hasitive for the order (Johnsgard 1965a,Kear 1970), tened by widespread glaciations(cf. Ploeger nestinghabit is probablyunreliable for intraor- 1968). Consequently, I agree with Cracraft dinal phylogenetic inferences. Clutch size, (1980) that the Anseriformes probably origiproportion of yolk in eggs,incubation period, nated in the Southern Hemisphere. parental carrying of young, and sexualdimorTaxonomic implications.--The tribes of Anatiphism also appear to be quite plastic (Johns- dae originally proposedby Delacourand Mayr gard 1961f,1966b;Lack1967,1968,1974;Johns- (1945) were defined primarily in terms of gengard and Kear 1968; Kear 1970; Livezey and era of the Northern Hemisphere. Although most tribes were assignedmembersfrom both Humphrey 1984). Selected reproductive characteristics, how- hemispheres, only the monotypic"Merganetever, show distinct primitive-to-derived se- tini" was limited to the Southern Hemisphere. quences(Kear 1970). Most changenear the di- A number of "aberrant" southern genera were vergence of the goose-swan clade but may be sorted tentatively among these tribes: Anserand Plectropterus the Cairinini; Cereopsis, to confoundedby an evolutionary trend toward anas and to reducedbody size: (1) nest bowl unlined vs. Tachyeres, Lophonetta the Tadornini; Sticlined with down (secondarilylost in stiff-tailed tonetta, Malacorhynchus, Hymenolaimus the and to to ducks); (2) biparental nest construction,incu- Anatini; and Thalassornis the Oxyurini. Since bation, and attendance of young vs. female then four of these genera have been placed in alone responsible; (3) broodingperiodand their own subfamilies or tribes, one has been and pair bond long ("anserines," roughly 6 months) moved to another subfamily, and the others vs. moderately long (shelducks, roughly 4 have remained problematic(Wolfenden 1961, months)vs. comparatively short (dabblersand Johnsgard 1978).Severalfindings in my study divers, less than 2 months). Participation of involve thesegenera,and suggest revision of a malesin brood rearing is variable within Anas, the classification waterfowl (Appendix 2). of The dabbling ducks are paraphyletic and however, wherein severalneotropicalspecies are characterizedby protracted, perhaps per- should be considered a phylogenetically unmanent pair bonds(Johnsgard 1978). resolvedgroup. I therefore place these genera Biogeographic patterns.--Despitethe early ac- in a provisionaltaxon, the "Anatini" (Appenknowledgmentof the diversityof "aberrantand dix 2). If paraphyly of this group is corroboprimitive" genera in Australia (Delacour and rated by further work, the erectionof addition-

Chenonetta, Rhodonessa,and Amazonetta (All

October 1986]

Phylogenythe of Anseriformes

747

caral tribes corresponding the branchesin the ALI, $. 1960. The Pink-headed Duck Rhodonessa to yophylacea (Latham).Wildfowl 11: 55-60. grade would be warranted. AMERICAN ORNITHOLOGISTS' UNION. 1983. Check-list Classification fossilgroups.--Severalearly of of North Americabirds, 6th ed. Washington,D.C., fossilscan be classified tentatively on the basis Amer. Ornithol. Union. of published descriptions (Howard 1964). RoBALL, C. 1934. Hybrid ducks,including descrip$. mainvillia(upper Eoceneor lower Oligocene), tions of two crosses Bucephala Lophodytes. of and Cygnopterus (upper Oligocene),and Paranyroca PeabodyMus. Nat. Hist. Bull. 3: 1-26. (lower Miocene) possess primitive procor- B^UMEI,, J. (Ed.). 1979. Nomina anatomicaavium. the J. acoidal foramen (character92) retained among New York, Academic Press. Recentgeneraonly by anhimids and Anseranas. BELLROSE, C. 1976. Ducks, geese and swans of F.

North America. Harrisburg, Pennsylvania, StackpoleBooks. H. anyroca derived with respectto Anseranas, BOETTICHER, VON. 1943. Die phylogenetisch-sysare tematische stellungyon Anseranas. Zool. Anz. 142: and the hypotarsus(72) of Paranyroca showsit

Conformation

of the tarsometatarsal

trochlea

(68) indicates that at least Romainvilliaand Par-

to be more primitive than Dendrocygna. Ac--. 1952. Ganse-und Entenv6gelausaller Welt. cordingly, these fossils should be listed after Leipzig, Germany, Geese&Portig K.-G. Anseranas and before Dendrocygna and se- BOTTJER, P.D. 1983. Systematic relationshipsamong quenced (provisionally by epoch of occurthe Anatidae: an immunological study: with a rence) as Romainvillia, Cygnopterus, Paranyand history of anatid classification, and a systemof

55-58.

roca; fossils the may be given familial namesor classification.Unpublished Ph.D. dissertation, New Haven, Connecticut, Yale Univ. be designated"piesions" (Wiley 1981) at familial rank. Phylogeneticreappraisals sev- BRVSH,A. H. 1976. Waterfowl feather proteins: of analysis of use in taxonomic studies. J. Zool. eral other fossilAnseriformes [e.g.Anas(?) blanchardi, the tadornines Anabernicula and
London 179: 467-498.

Brantadorna, and Chendytes; Howard 1964], and the probableanseriformPresbyornis (Olson and

COTTER, B., JR. 1957. A serologicalanalysisof W.

some anatid classifications. 300. Wilson Bull. 69: 291-

Feduccia 1980a), shouldprovideminimum ages of branchpointsin the phylogeny.

ACKNOWLEDGMENTS

CRACRAFT, 1980. Avian phylogeny and interconJ. tinental biogeographicpatterns. Pp. 1302-1308 in Acta 17th Congr. Intern. Ornithol., vol. II (R. Nohring, Ed.). Berlin, DeutschenOrnithologenGesellschaft.

This work was supported in part by National Science Foundation grant BSR-83-19900and the University of KansasMuseum of Natural History. I am most grateful for the enthusiasmand insights of P.

DAVIES,$. J. J. F., & H. J. FRITH. 1964. Some com-

ments on the taxonomicposition of the Magpie GooseAnseranas semipalmata (Latham). Emu 63:
265-272.

S. Humphrey; his comments the manuscript, on and those of G. E. Woolfenden, D. Siegel-Causey, M. R. Chandler, and W. Hoffman are greatly appreciated.I alsobenefitedfrom discussions with E. O. Wiley, D. L. Swofford,L. M. Witruer, D. Frost,R. L. Mayden, L. D. Martin, D. E. Seibel,and D. M. Hillis. R. Mengel and M. Jenkinsonpermitted access specimens to and
work space, and loans of specimenswere arranged by S. L. Olson (National Museum of Natural History), R. W. Storer (Museum of Zoology, University of Michigan), G. E. Woolfenden (Department of Biolo-

DELACOUR, 1954. The waterfowl of the world, vol. J.

1. London, Country Life.

1956. The waterfowl of the world, vol. 2.

London, Country Life.

1959. The waterfowl of the world, vol. 3.

London, Country Life.

1964a. Domestic waterfowl. Pp. 154-166 in The waterfowl of the world, vol. 4 (J. Delacour,

gy, Universityof SouthFlorida),and P. Devillers(Institut Royal des Sciences Naturelies de Belgique).J. Cracraftand A. H. Bledsoe provided useful criticisms of the manuscript. thank M. A. Masten for graphical I help and K. McManness typing the manuscript. for
LITERATURE CITED

Ed.). London, Country Life. 1964b. Aviculture. Pp. 145-153 in The waterfowl of the world, vol. 4 (J. Delacour, Ed.). London,Country Life. 1964c. Correctionsand additions. Pp. 327354 in The waterfowl of the world, vol. 4 (J. De-

lacour,Ed.). London, Country Life. -, & E. MAYR. 1945. The family Anatidae. Wilson Bull. 57: 3-55.

AOTT, C.E.

1938. The skulls of ducks. Wilson Bull.

,& . 1946. Supplementary noteson the family Anatidae. Wilson Bull. 58: 104-110.

50: 249-253.

DEMAY,I. $. 1940. A study of the pterylosisand

740

BIDLE LIVEZE C.
pneumaticity of the screamer.Condor 42: 112118

[Auk,Vol. 103

FEDUCCIA, 1977. Hypothetical stagesin the evoA. lution of modernducksand flamingos. Theor. J.
Biol. 67: 715-721.

-, & A. GLASER. 1975. Chemotaxonomy Anof seriformes.Blochem. Syst. Ecol. 2: 215-220. JOHNSGARD, P.A. 1960a. The systematic position of

the Ringed Teal. Bull. Brit. Ornithol. Club 80:

165-167.

--.

1978. Presbyornis the evolution of ducks and

1960b. Classification and evolutionary re-

and flamingos. Amer. Sci.66: 298-304. FRITH, H.J. 1955. The downy ducklingsof the Pinkeared and White-eyed ducks. Emu 55: 310-312. 1964a. Taxonomicrelationship of Stictonetta naevosa (Gould). Nature 202: 1352-1353. 1964b. The downy young of the Freckled
Duck Stictonetta naevosa. Emu 64: 42-47.

lationships the seaducks.Condor62:426-433. of 1960c. Comparativebehaviour of the Anatidae and its evolutionaryimplications.Wildfowl
11: 31-45.

1967.

Waterfowl

in Australia.

Honolulu,

Hawaii East-West Center Press.

1960d. Hybridization in the Anatidae and its taxonomic implications. Condor 62: 25-33. 1961a. The taxonomy of the Anatidae--a behavioural analysis.Ibis 103a:71-85. 1961b. The systematic position of the Marbled Teal. Bull. Brit. Ornithol. Goose. Wildfowl 12: 92-103. Club 81: 37-43.

GRAY, A.P. 1958. Bird hybrids:a checklistwith bibliography. Bucks, England, CommonwealthAgr.
Bur.

1961c. Breeding biology of the Magpie

1961d. The behavior and systematicposition of the Hooded Merganser.Wilson Bull. 73:
227-236.

HARRISONG. 1958. Skull pneumaticityin wildJ.

fowl in relation to their mode of life. Wildfowl
9: 193-196.

HOBBS,J. N.

1957. Notes on the Pink-eared Duck.

Emu 57: 263-268.

1961e. Tracheal anatomy of the Anatidae

HOWARD,H.
shellmound.
394.

1929. The avifauna of Emeryville

Univ. California Fossil evidence Publ. Zool. 32: 301of avian evolution.

and its taxonomic significance. Wildfowl 12:5869.

1961f. Evolutionary relationships amongthe

North American Mallards. Auk 78: 3-43.

1950.

Ibis 92: 1-21.

1962. Evolutionary trends in the behaviour

1964. Fossil Anseriformes. Pp. 233-326 in

The waterfowl of the world, vol. 4 (J. Delacour,

and morphologyof the Anatidae.Wildfowl 13:

130-148.

Ed.). London, Country Life. HOWARD & A. MOORE R., 1984. A completechecklist of the birds of the world. London Macmillan.

1963. Behavioralisolating mechanisms in the family Anatidae. Pp. 531-543 in Proc. 13th Intern. Ornithol. Congr.,vol. I (C. G. Sibley,Ed.).
Amer. Ornithol. Union.

HUMIHRE, P.S. 1955. The relationships of the seaducks(Tribe Mergini). Unpublished Ph.D. dissertation Ann ArborUniv. Michigan. 1958. Classification and systematic position
of the eiders. Condor 60: 129-135.

1964. Comparativebehavior and relationshipsof the eiders.Condor 66: 113-129.

1965a. Handbook London, Constable.
tralian

of waterfowl

behavior.

& R. S. BUTSCH.1958. The anatomy of the Labrador Duck Camptorhynchuslabradorius (Gmelin). Smithsonian Misc. Coil. 135: 1-23. , & G. A. CLARK, JR. 1964. The anatomy of waterfowl. Pp. 167-232 in The waterfowl of the worldvol. 4 (J. Delacour, Ed.). LondonCountry
Life.

1965b. Observations on some aberrant AusAnatidae. Wildfowl 16: 73-83.

1966a. The biology and relationships the of

Torrent Duck. Wildfowl 17: 66-74. 1966b. Behavior of Australian Musk Duck and Blue-billed Duck. Auk 83: 98-110.
1967. Observations on the behaviour and

relationships the White-backed of Duck and the

stiff-tailed ducks. Wildfowl 18: 98-107.

-, & B.C. LIVEZE. 1985. Nest, eggs,and downy young of the White-headedFlightlessSteamerDuck.Pp. 949-954in Neotropicalornithology(P. A. Buckley, M. S. Foster, E. S. Morton R. S. Ridgely, and F. G. Buckley, Eds.). Ornithol. Monogr. No. 36.
& S. D. RIPLEY. 1962. The affinities of the

1978. Ducks, geese swans the world. and of

Lincoln, Univ. Nebraska Press.

1979. Order Anseriformes. Pp. 425-506 in

Checklist of birds of the world, 2nd ed., vol. 1

Pink-headed Duck (Rhodonessa caryophyllacea). --,

(E. Mayr and C. W. Cottrell, Eds.). Cambridge, Massachusetts Mus. Comp. Zool. & J. KEAR. 1968 A review of parental car-

Postilia 61: 1-21. rying of youngby waterfowl.Living Bird 7: 80102. JACOB, 1982. Integumentlipide--ihre chemische J. A. Struktur und ihre Bedeutungals systematisches JOHNSON, W. 1963. Notes on the distribution, reproduction and displayof the AndeanTorrent Merkmal in der Zoologie. Funkt. Biol. Med. 1: 83-90. Duck Merganetta armata.Ibis 105: 114-116.

October 1986]

Phylogeny Anseriformes ofthe

749

KEAR, 1967. Notes of the eggsand downy young J.

of Thalassornis leuconotus. Ostrich 38: 227-229.

tematischen Stellung der Sturzback-Ente. OrJ.

nithol. 93: 357-360.

1970. The adaptiveradiationof parentalcare in waterfowl. Pp. 357-392 in Socialbehaviour in birds and mammals(J. H. Crook, Ed.). New York,
Academic Press.

NILSSON, 1974. The behaviourof wintering Smew L.

in southern Sweden. Wildfowl 25: 84-88.

OLSON,S. L. 1985. The fossil record of birds. Pp. 79-238 in Avian biologyvol. 8 (D. S. Farner, J.

1972. 1975.
Wildfowl

The Blue Duck of New Salvadori's

26: 104-111.

Zealand.

Liv-

R. King, and K. C. Parkes,Eds.).New York, Academic Press.

ing Bird I1: 175-192.

Duck of New Guinea.

--,

& R. K. MURTON. 1973. The systematic status of the CapeBarrenGoose judgedby its photo- --, as responses. Wildfowl 24: 141-143. Smithsonian Contrib. Zool. 316: 1-73. T. H. STEEL.1971. Aspectsof social behaviour in the Blue Duck. Notornis 18: 187-198. PALMER,R. S. (Ed.). 1976. Handbook of North American birds vols. 2 and 3. New Haven, ConKESSLER, G., & J. C. AVISE. 1984. Systematic L. relanecticut, Yale Univ. Press. tionships among waterfowl (Anatidae) inferred PETERS, L. 1931. Check-list of birds of the world, J. from restriction endonuclease analysisof mitovol. 1. Cambridge, Massachusetts, Harvard Univ. chondrial DNA. Syst.Zool. 33: 370-380. Press. KLVGE, G. 1984. The relevance of parsimony to A. J. phylogenetic inference. Pp. 24-38 in Cladistics: PHILLIPS, C. 1922. A natural history of the ducks, vol. I. Boston,Houghton Mifflin. perspectives the reconstruction evolutionon of 1923. A natural history of the ducks,vol. 2. ary history (T. Duncan and T. F. Stuessy,Eds.). --. Boston,Houghton Mifflin. New York, Columbia Univ. Press. 1925. A natural history of the ducks,vol. 3. LACK,D. 1967. The significance of clutch-size in Boston,Houghton Mifflin. waterfowl. Wildfowl 18: 125-128. P.L. 1968. Geographicaldifferentiation in 1968. The proportion of yolk in the eggsof PLOEGER, arctic Anatidae as a result of isolation during the waterfowl. Wildfowl 19: 67-69.

& A. FEDUCCIA. 1980a. Presbyornis the and origin of the Anseriformes(Aves: Charadriomorphae). Smithsonian Contrib. Zool. 323: 1-24. & --. 1980b. Relationships and evolution of flamingos (Aves: Phoenicopteridae).

1974. Evolution illustrated by waterfowl. Oxford, England,BlackwellSci. Publ.

LIVEZEY, B.C., & P.S. HUMPHREY 1984. Sexual di-

Last Glacial.

Ardea

56: 1-159.

RAIKOW, R.J. 1970a. The function and evolution of

the supraorbitalprocess ducks.Auk 87: 568in

572.

morphism in continental steamer-ducks. Condor

86: 368-377.

1970b. Evolution of diving adaptationsin

the stifftail
94: 1-52.

LORENZ, 1951-1953. Comparativestudieson the K. behaviour of Anatinae. Avicult. Mag. 57: 157182; 58: 8-17, 61-72, 86-94, 172-184; 59: 24-34,
80-91.

ducks.

Univ.

California

Publ.

Zool.

--.

1971. The osteologyand taxonomicposition

of the White-backed
tus. Wilson Wilson

Duck, Thalassornis leucono-

MAINARDI,D. 1962. Immunological data on the phylogeneticrelationshipsand taxonomicposition of flamingoes(Phoenicopteridae). Ibis 104:
426-428.

Bull. 83: 270-277.

RAND,A. L. 1954. On the spurs on birds' wings.

Bull. 66: 127-134.

MAYR,E. 1931. Anatomie und systematischen Stellung der Salvadori-Ente (Salvadorina waigiuensis
Rothsch.& Hartert). Ornithol. Monatsberichte 39:
69-70.

REMS, C., & N HILLGARTH. 1984. The breeding E. biology of captive Black-headedDucks and the behavior of their young. Condor 86: 242-250.

RIPLEY, S.D. 1957. A paddlingof ducks.New York,

Harcourt, Brace, and Co.

MCKINNEY, F. 1953. Studies on the behaviour of the

Anatidae. Unpublished Ph.D. dissertation BristolEngland, Univ. Bristol.

MILLER, W. DEW. 1919. Notes on the structure of

Anseranas semipalmata. 36: 562-564. Auk MOFFETT M., JR. 1970. A studyof nestingTorrent G. Ducksin the Andes. Living Bird 9: 5-28.
MOYNIHAN, M. 1958. Notes on the behavior of the

SALVADORI, 1895 Catalogue of the ChenomorT. phae (Palamedeae, Phoenicopteri, Anseres), Crypturi and Ratitaein the collection the Britof ish Museum. London, Longmans. SCOTT, 1985. Duck. Pp. 157-161 in A dictionary P. of birds (B. Campbell and E. Lack, Eds.).Vermillion, South Dakota, Buteo Books.

SIBLEY, C.G. 1957. The evolutionary and taxonomic significanceof sexual dimorphism and hybridization in birds. Condor 59: 166-191.

Flying SteamerDuck. Auk 75: 183-202. MYRE$, M.T. 1959. Display behavior of Buffiehead, scoters, and goldeneyes at copulation. Wilson
Bull. 71: 159-168.

1960. The electrophoreticpatternsof avian egg-white proteinsas taxonomiccharacters. Ibis

102: 215-284.

NIETHAMMER, G.

1952. Zur Anatomie und sys-

--,

& J.E. AHLQUIST. 1972. A comparative study

750

BRADLEY LIVEZE C.
of the egg white proteins of non-passerinebirds. PeabodyMus. Nat. HistBull39: 1-276. K. W. CORBIN,& J. H. HAAVIE. 1969. The relationships of the flamingos as indicated by the egg-white proteins and hemoglobins Condor 71: 155-179.

[Auk, Vol. 103

gentina,Paraguay, Uruguayand Chile. U.S. Natl.

Mus. Bull. 133.

SIEGFRIED, R. 1970. Double wing-moult in the W.

Maccoa Duck Wildfowl 21: 122.

1951. Observations on the genera of the swans.J. WashingtonAcad. Sci. 41: 338-340. WILEY,E. O. 1981. Phylogenetics:the theory and practiceof phylogeneticsystematics. New York, J. Wiley and Sons.
WOLFF, & E. WOLFF1952. La dterminisme de la E.,

SILEO,L., C. H. TUITE, & J. B. D. HOPCRAFT. 1977

diffrenciation sexuelle de la syrinx du canard

cultive in vitro. Bull. Biol. 86: 325-349.

Plumages and primary moult in LesserFlamingos Wildfowl 28: 139-142. SIMONETTA, A.M. 1963. Cinesi e morfologia del
cranio negli uccelli non passeriformi. Studio su
varie tenenze evolutive. Parte 1. Arch Zool. Ita-

WOOD, D. S., R. L. Zusi, & M. A. JENKINSON. 1982.

World inventory of avian skeletal specimens,

1982. Norman, Oklahoma, Amer. Ornithol. Union
and Oklahoma Biol. Surv.

lia (Torino) 48: 53-135.

WOOLFENDEN, E. 1961. Postcranialosteologyof G.

the waterfowl. Florida State Mus. Bull. 6: 1-129. SWOFFORD, 1984. PAUP: phylogenetic D.L. analysis YAMASHINA, Y. 1952. Classification of the Anatidae using parsimony.Illinois Nat. HistSurv. (Mim-

eo0 based on cytogenetics.Pap. Coord. Comm. Res. Gen. 3: 1-24. TIMMERMANN, 1963. Fragen der AnatidensysteG. matik in parasitologischer Sichr. Pp. 189-197 in ZusI, R. L., & G. D. BENTZ. 1978. The appendicular Proc. 13th Intern. Ornithol. Congr., vol. I (C. G. royologyof the LabradorDuck (Camptorhynchus labradorius).Condor 80: 407-418. Sibley, Ed.). Amer. Ornithol. Union TODD, F.S. 1979. Waterfowl:ducks,geeseand swans

of the world New York, Harcourt Brace Jovanovich

TYLER, 1964. A study of egg shellsof the AnatiC.

dae. Proc. Zool. Soc. London 142: 547-583

APPENDIX

VERHEYEN, 1955. La systmatiquedes AnsriR.

formesbasesur l'ostologiecompare.Bull Inst

RoyalSoc.Natur. Belgique 31(35):1-18, (36): 116, (37): 1-22, (38): 1-16. WARHAM,J. 1959. The nesting of the Pink-eared
Duck Wildfowl 9: 118-127.

WELLER, W. 1959. Parasiticegg laying in the M. Redhead(Aythya americana) and other North American Anatidae. Ecol. Monogr. 29: 333-365. 1964a. Fowling. Pp. 121-127 in The waterfowl of the world, vol. 4 (J. Delacour, Ed.). Lon-

The 120 characters used in this analysisare listed below and are numberedand grouped anatomically. Character states are lettered and correspond to the character changes Figs.2-5. Plesiomorphic in (primitive) conditions generally are designated "a" and derived character statesare ordered alphabetically thereafter (implying a linear transformationseries); characters followed by a "U" were analyzed as unordered. Characters for which the primitive state was
not determined are marked with "U*." Characters

don, Country Life 1964b. General habits. Pp. 15-34 in The waterfowl of the world, vol. 4 (J. Delacour, Ed.). London, Country Life

judged to be unusually variable, generally necessitating determinations of modal conditions, are indicatedwith a "V." Taxa with problematicstate determinations are listed in parentheses after the corresponding character. Consistency indices (CI) follow each character.Anatomical terminology follows Howard (1929) and Woolfenden (1961) unless
annotated otherwise.

1964c. The reproductivecycle. Pp. 35-79 in

The waterfowl of the world vol. 4 (J. Delacour,

Ed.). London Country Life. 1964d. Distribution and speciesrelationships 108-120in The waterfowlof the world, Pp. vol. 4 (J. Delacour,Ed.). London,Country Life 1967. Notes on plumages and weights of the Black-headed Duck, Heteronettaatricapilla.
Condor 69: 133-145.

Integument

1. Molt of remiges:(a) sequential; synchronous, (b) once annually; (c) synchronous, twice annually. (Variable in Phoenicopteridae;
Sileo et al. 1977.) CI = 1.0.

1968a. Plumagesand wing spurs of Torrent

--.

DucksMerganetta armata. Wildfowl 19: 33-40 1968b. The breedingbiologyof the parasitic
Black-headedDuckLiving Bird 7: 169-207.

2. Tarsalsheath:(a) scutellate anteriorly and posteriorly;(b) reticulate anteriorly and posteriorly;(c) scutellateanteriorly (at least distally)and reticulateposteriorly. = 1.0. CI 3. Spongysubcutaneous layer:(a) absent; present. = 1.0. (b) CI 4. Interdigitalwebbingof feet (excluding hallux):(a) lacking(slight webbingin anhimids);(b) incomplete(semipalmate); complete (c) (incised in Cereopsis Brantasandwicensis). = 1.0. and CI 5. Apteria: (a) present;(b) obsolete.CI = 1.0.
Trachea

1976. Ecology and behaviour of steamer

ducks Wildfowl 27: 45-53

WETMORE, 1926. Observations on the birds of ArA.

6. Bulla ossea males (U, weight = 2): (a) not enlarged;(b) symof metrically enlarged;(c) asymmetricallyenlarged,unfenestrated;

October 1986]

Phylogeny Anseriformes ofthe

751

vated,essentially parallels shaft;(b) elevated, angleddistally; (c) (d) asymmetrically enlarged, fenestrated; enlargementre(e) elevated, angled medially. CI = 0.40. ducedand symmetrical, obsolete. or (Thalassornis, Malacorhynchus, 27. Internaltuberosity: proximally (a) rotated, exposing completely Nettapus.)CI = 0.67. the pneumaticforamenin anconalview, lackinga distinct,dis7. Extrasternal, subdermal 1ooping trachea males(U*): (a) presof in
ent; (b) absent. CI = 0.33. 8. Inflatable tracheal air sacs:(a) absent;(b) present.CI = 1.0
Skull

tally directed prominence; produced (b) distally asto largely so

or completely obscure pneumatic foramenin anconalview, typically with prominent,distallydirectedpoint.CI = 0.50. 28. Pneumatic fossa (U): (a) open,usuallycontainingnumerous bony struts; closed bonyshellexcept a smallcentral (b) by for opening;

9. Occipitalfontanelles: absent; present. = 1.0. (a) (b) CI 10. Lacrymals (a) not fusedto skull;(b) fusedto skulldorsally, (U): small,nonpneumatic; fuseddorsally, (c) moderately thick,long,
ventrally directed (lacking flange), nonpneumatic;(d) fused to

(c)completely closed; closed perforated numerous (d) but by small

holes.(Lophodytes.) = 0.43. C1 29. Attachment of M. latissimus site dorsiposterioris: well medial (a) to externaledge of pectoralattachment; in line with outer (b) edgeof pectoral attachment, anconal on surface shaft;(c) in of line with outer edge of pectoral attachment, on raised ridge.
CI = 1.0.

skull dorsally, and (typically)also fusedto postorbital process, slightly pneumatic;(e) fused dorsally, of variable shape and pneumaticity, posterioventrally directed, oftenwith flangedventral terminus. CI = 1.0.

11. Supraorbital process bestdeveloped adultmales): absent (U; in (a) or small,straight, essentially coplanar with dorsalsurface skull; of (b) large, flat, medially appressed dorsalmargin of orbit; (c) to large,thick,rugose, dorsolaterally directed; long,slender, (d) dorsallydirected, often curved.(Aythya, Bucephala albeola.) = 0.38. CI 12. Anterior terminusof premaxillae: stronglyventrally hooked, (a) typically pointed; (b) strongly ventrally hooked, moderately rounded;(c) not ventrally hooked(directedanteriorly), rounded,
spatulate.C1 = 1.0.

30. Distalportionof anconal surface bicipitalcrest: poorlydeof (a) veloped shelflike; produced or (b) medially with distinct proximal

cuplike depression, visible translucent as window palmar in view.

CI = 0.50.

13. Bill lamellae: absent; present. (a) (b) (Nonhomologous lamellae

in flamingos,vestigial in anhimids;Olson and Feduccia1980a.)
CI = 1.0.

31. Distal terminusof deltoid crest:(a) essentially continuous with lateraledgeof shaft;(b) produced into prominenttuberosity on palmarsurfaceof shaft.CI = 1.0. 32. External tuberosity: prominent, (a) buttressed, typicallywith attachmentsite elevated,parallel to shaft, not slopingaway with ancohal surface deltoidcrest; reduced,lackingbuttress, of (b) with attachment sloping essentially site and flushwith ancohal surface of deltoid crest. (Stictonetta, Sarkidiornis, Cyanochen, Merganetta,
Oxyura.)CI = 0.33.

14. Retroarticular processes mandible:(a) lacking, small, or reof curvedand rounded;(b) recurved,pointed,and bladelike.CI =
1.0.

15. Quadrate, lateralview: (a) not squarish, with variablydeeplycurved

dorsalmarginbetweenorbitaland otic processes; squarish, (b)

with dorsalmargin straight.CI = 0.33.

33. Relativeanconal heightsof ectepicondyle entepicondyle: and (a) ectepicondyle distinctly higherthanentepicondyle; condyles (b) essentially equallyhigh; (c) ectepicondyle lower than entepicondyle. (Hymenolaimus.) = 0.40. CI 34. Pit for attachment M. flexor carpi ulnaris:(a) prominent;(b) of
reduced to obsolete. CI = 1.0.

16. Frontonasal regionof skull (U): (a) essentially continuous with profiledefinedby premaxillae frontals; enlarged and (b) into conspicuous, laterally compressed, dorsal prominence (largerin males); (c)with rounded, pneumatic swelling(especially adultmales). in
CI = 0.50.

35. External condyle brachial and depression (palmar side, distalend): (a) separated smooth by strip of bone;(b) connected rounded by
ridge. CI = 1.0.

17. Frontals: without dorsallydirectedhornlike prominence; (a) (b)

with small, ossified "horn" on midline. CI = 1.0.

36. Attachment forexternal site headof triceps: immediately (a) distal to head,typically excavation in underhead; displaced (b) distally
on lobe of bone and obscures externalterminusof capitalgroove.
CI = 1.0.

18. Pterygoid-palatine articulation: a simpleabutment; a ball(a) (b) and-socket arrangementinvolving two extensions the pteryof Carpometacarpus gold. CI = 1.0. 37. Distal end of internal rim of carpaltrochlea(externalview): (a) 19. Dorsumof upper bill in region of externalnares:(a) essentially with prominentswelling;(b) without prominentswelling;(c) continuous with curvature of skull to somewhat convex; (b) subdeeplyexcavated. (Marmaronetta, Hymenolaimus, Clangula, Mergelstantiallydorsallybowed. CI = 1.0. lus,Heteronetta.) = 0.33. CI 20. Basipterygoid processes: lacking or (in Galliformes)present (a) 38. Externalrim of carpaltrochlea: essentially (a) continuous, unbutwithoutbasal supports; present, (b) lipped,almost pedicellate. notched; with prominentnotchdistally.CI = 1.0. (b)
CI = 1.0.

39. Dorsal surfaceof metacarpal (a) flattened proximally (can apII: pearangular);(b) roundedproximally.CI = 1.0.

Vertebrae

40. Tuberosity metacarpal (a) small; prominent, of II: (b) spurred. = CI 41. Angle of process metacarpal (a) perpendicular or proxiof I: to mally directedrelativeto shaft;(b) angleddistally.(Plectropterus.)
CI = 1.0.

21. Number of cervical vertebrae (U, V): (a) 17; (b) 18-20; (c) 21; (d) 22-25; (e) 16. (Some Tadorninae.) CI = 0.67.
Humerus

22. Capitalshaftridge:(a) prominentand directedtowardhead;(b) prominent and directed towardexternaltuberosity; obsolete (c) proximallyor absent completely. (Sarkidiornis, Cyanochen, Hymenolaimus,Chenonetta.) = 0.50. CI

42. Process metacarpal (U): (a) not enlarged,length lessthan of 1 width of trochlea; an enlarged, (b) pointed spur,longerthanwidth of trochlea; enlarged, (c) blunt, typicallywith rugose-capped spur, longerthanwidth of trochlea. (Plectropterus, Hymenolaimus, Lophonetta.) CI = 0.50.

23. Capitalgroove: short,essentially (a) directed distally; extend(b) ing laterally toward external tuberosity, undercutting head. (Thal
assornis.) 0.50. CI

43. Attachmentsite of M. extensor metacarpiulnaris(Zusi and Bentz 1978;"flexor" of Woolfenden 1961) (U, V): (a) completelyproximal to proximalfornix of metacarpal and III: (b) opposite, II at

24. Proximo-anconal region: (a) variably rounded by shaft; (b) traversedby a deep,uninterrupted trenchlikedepression from under head to internal edge immediatelydistal to bicipital crest.
CI = 0.33.

leastpartly, fornix; (c) completely distal to fornix. (Tachyeres.)

CI = 0.29.

44. Lower proximalsurface metacarpal (a) ungrooved, of III: rounded; (b) distinctly grooved. CI = 0.50. 45. Facetsfor digits II and III (U*): (a) facet for digit III extending

25. Deltoid crest(V): (a) margin rounded,laterally flaring, concave anconally; marginangular squared, (b) or depressed around shaft toward palmar side, convexanconally.(Sarkidiornis, Hymenolaimus.) CI = 0.50. 26. Surfaceof attachment for anterior articular ligament: (a) not ele-

fartherdistallythan facetfor digit II; (b) facets essentially equal

in distal extent. CI = 0.33.

46. Cuneiformfossa: shallowto moderatelydeep;(b) deep,round(a)

ed, ovate, with distinct rim. CI = 1.0.

752
APPENDIX 1. Continued.

BDLE LIVE C.

[Auk,Vol. 103

47. Distal portion of internal rim of carpal trochlea: (a) of uniform thicknesswith proximal portion; (b) distinctly thickened. CI 0.50.

48. Internal rim of carpal trochlea(posteriorview): (a) in line with internal margin of shaft;(b) sharplydeflectedlaterally. (Callonetta.) CI 1.0.

Radial carpal
49. Size and shape:(a) small, short,and blunt; (b) elongatedinto large pointed spur. CI - 1.0.

Appendicular pneumaticity 50. Distal alar and pelvic elements: essentially (a) nonpneumatic; (b) pneumatic with one or more large foramina. CI - 1.0.
Femur

72. Calcaneal ridges of hypotarsus:(a) 2, lateral to midline of shaft, borderedmedially by depression (deep in Anseranas); 3 or 4, (b) situated on midline of shaft, without depressionon internal margin. CI = 1.0. 73. Wing on trochlea for digit lI: (a) not prominent medially; (b) medially prominent, thickened. CI = !.0. 74. Groovein trochleafor digit II: (a) absent; present,but posterior (b) terminus of groove variable in extent. CI - 1.0. 75. Anterior extent of internal and externalridgesof shaft:(a) essentially equal, no twisting of shaft about its long axis;(b) internal ridge lessprominentanteriorlythan external,becoming flushwith shaft immediately distal to proximal foramen, associated with moderate twisting of shaft; (c) internal edge of shaft depressed below level of shaftanteriorly,associated with strongtwisting of shaft. (Hymenolaimus, Merganetta, Amazonetta.) = 0.33. CI 76. External margin of shaft: (a) concave(in anterior profile), curving smoothly to external surface of trochlea for digit IV; (b) essentially straight, trochlea for digit IV internally deflected. CI = 1.0.

51. Head, relative to plane of external surfaceof shaft:(a) oriented posteriorly;(b) perpendicular.CI - !.0. 52. Anterior extent of trochanter: (a) relatively great, such that anterior-posterior depth of trochanter substantially exceeds depth of head; (b) reduced,such that depth of trochanteronly equals
that of head. CI = 0.33.

77. Posterior openingof distalforamen: directed (a) posteriorly, flush with surfaceof shaft; (b) directed distoposterinrly, recessed in depression immediatelyproximal to symphysis trochleafor of
digits III and IV. (Anseranas.) = 1.0. CI
Sternum

53. Distal extentof internal condyle:(a) distinctlylessthan that of externalcondyle; equalto that of externalcondyle. (b) CI 1.0.
54. Rotular depression:(a) shallow to moderately deep, margin rel-

78. Pneumatic foramen(U*): (a) open, ovoid;(b) pitted, largely occluded by medial bar; (c) closed (sometimesmarked by small
depression).CI = 0.25. 79. Ventral manubrial region (U*, V): (a) keel-like, laterally compressed medial flange;(b) thick medialwedge;(c) lackingmedian protuberance(s); long, peglike spine;(e) small lump; (f) a pair (d) of small pointed prominencesseparatedat midline by a deep excavation, typically with an ovoid pit at base;(g) a wide, moderatelylong, dorsoventrally compressed flange;(h) a pair of points partially separatedby a shallow midline excavation;(i) a small, unforked, dorsoventrallycompressed flange.(Cyanochen, Hymen olaimus, Cairina, Lophonetta, Chenonetta, Polysticta.) = 0.50. CI 80. Carina (keel) shape, lateral profile: (a) well developed, ventral margin curved throughout length; (b) reduced,ventral margin essentiallystraightfor posteriorhalf. CI - 0.33. 81. Posterior-lateral processes (U*): (a) extendwell posteriorto postpectoral line of sternal plate; (b) approximatelyequal to postpectoral line in posterior extent. CI = 0.33. 82. Dorsal manubrial region (U, V): (a) rounded notch; (b) rounded notch with small point on midline; (c) rounded notch with moderately large point on midline; (d) even shelf. (Malacorhynchus, Hymenolaimus, Marmaronetta, Callonetta.) = 0.43. CI 83. Abdominal plate (dense,symmetricalextensionof sternal plate

atively indistinct; deep,distinctlyborderedproximally.CI = (b)

1.0.

55. Curvatureof shaft,lateralview (U): (a) straightto slight;(b) moderate;(c) strong,subangular. - 0.22. CI 56. Poplitealfossa: shallow;(b) deep, typically pitted. CI -- 0.25. (a) 57. Lobe at midpoint of posterior surfaceof shaft: (a) not prominent; (b) prominent. CI - 1.0. 58. Posteriorintermuscularline (U): (a) relatively distinct,following internal edge of shaft;(b) relatively distinct, swingslaterally toward trochanter; (c) indistinct. CI = !.0.

59. Internal edge of distal end of shaft: (a) smoothly curving and continuouswith proximal portion; (b) leveled by raisedridge to internal condyle. CI = 1.0. 60. Posteriorintermuscular line: (a) distinguishable only asfine etching; (b) forming overhanging ridge proximally. CI = 1.0.
Tibiotarsus

61. Proximal articulating surface:(a) in line with shaft, squareswith distal condyles; stronglyrotatedcounterclockwise (b) aboutshaft (proximal view). CI = 0.50. 62. Rim of internal condyle:(a) distinctly notched;(b) lackingnotch.
CI = 1.0.

posterior both post-pectoral and posterior-lateral to line processes):(a) absent;(b) present. CI !.0. 84. Sternal notches(posteriormargin of plate medial to posteriorlateralprocesses): typicallyopen posteriorly; typicallyclosed (a) (b) posteriorly, forming fenestrae.CI 1.0. 85. Xiphial area:(a) posteriormargin approximatelystraightor concave;(b) with medial, irregularly shaped,roughly circular extension of thin bone (anterior to posterior-lateralprocesses). = CI
1.0.

63. Inner cnemial crest:(a) not deflectedlaterally; (b) laterally deflected. CI = 1.0.

64. Anterior extent of condyles:(a) internal distinctly greater than external; (b) approximately equal. (Aythya.) CI = 0.25.

65. Inner cnemialcrest: lackingdistinctridge extendingdistally (a) along anterior surfaceof shaft; (b) continued by distinct ridge distally alonganteriorsurface shaftto point well beyondproxof
imal terminus of fibular crest. CI = 0.25.

66. Internal condyle, posterior view: (a) with relatively rounded internal edge;(b) flared, with squaredmedial edge. CI - !.0. 67. External ligamental prominence: (a) essentially continuouswith

curvature shaft;(b) produced of laterally,ridgelike.CI = !.0.

Tarsometatarsus

68. Trochleafor digit 11: approximately (a) equal to trochleafor digit IV in distal extent;(b) proximal to trochleafor digit IV. CI - 1.0. 69. Anterior (of two) ligamental passages between trochleafor digits
III and IV (in distal wall of distal foramen) (V): (a) obscured from view anteriorly by bone; (b) largely or completely exposedante-

86. Costal margin: (a) comprises less than half of basin length; (b) comprisesmore than half of basin length. CI - 1.0. 87. Carina: (a) lacking pneumatic foramen in anterior margin, apparently solid; (b) with small pneumatic foramen in anterior edge, but carinauninflated;(c) hollow, containingloop of trachea, with large pneumatic foramen in anterior edge. CI 1.0. 88. Intermuscularline: (a) anglesmedially to carlhal basewell anterior to posterioredgeof plate;(b) extends posteriorlyto posterior margin of plate. CI 0.33.
89. Foramina of basin (U*): (a) limited to midline and anterior mar-

gin; (b) essentiallyabsent;(c) present on anterior margin, midline, and scatteredacrossplate (often among transversebony
striatinns). CI = 0.50.

riorly because reductionof bony covering.CI - 0.25. of 70. Internal calcaneal ridge of hypotarsus: slightly to moderately (a) exceeds other calcanealridges in posteriorextent;(b) greatly exceedsother (more external) calcanealridges. CI - 0.50. 71. Facetfor metatarsalI: (a) deep; (b) obsolete.CI - 1.0.

90. Midpoint of coracoidalsulcus:(a) solid; (b) having oval pneumatic foramen.

Costae

CI = !.0.

91. Uncinate processes: present; (b) absent. CI - 1.0. (a)

October 1986]
Coracold

Phylogeny Anseriformes ofthe

753

92. Procoracoidalforamen: (a) present (variable in Chauna);(b) absent. CI = 1.0.

93. Pneumatic foramen on dorsal surface anterior to sternal facet: (a)

present;(b) absent.CI = 1.O. 94. Dorsal sternalfacet:(a) with anteriorborderessentially smoothly curving;(b) with prominentcircularinternallip. CI = 1.0. 95. Brachialtuberosity(U): (a) essentially without foraminaunder posterioredge;(b) with small foraminaunder posterioredge;(c) with small foramina,typically containedwithin larger foramina, under posterior edge. CI = 0.50. 96. Depressionon ventral surfaceanterior to sternal facet (U*): (a) present,typically deep;(b) absent.(Anseranas.) = 0.25. CI 97. Furcular facet:(a) with posteriormargin completeor slightly reduced; posterior (b) margin deeplynotched. = 1.0. CI 98. Angle of head:(a) coplanar slightlyventral to planeof blade; to (b) distinctlyventralto plane of blade.CI = 1.0. 99. Sternocoracoidal process: wide, long, and rounded flange,ex(a) tending farther laterally than sternal facet;(b) variably shaped, rounded or angular process, approximatelyequal to sternalfacet in lateral extent;(c) long pointed process, extending farther laterally than sternalfacet.CI = 0.67. 100. Ventral (external) sternal facet(V): (a) anterior margin moderately raised or continuouswith blade; (b) anterior margin with distinct
buttress. CI 0.33.

118. Dorsolateral crests:(a) distinct to caudal margin of pelvis; (b) becomesobsoletecranial to caudal margin. CI = 1.0. 119. Anterior terminus of shield (posterior terminus of fusion of median dorsal ridge): (a) cranial to acetabula;(b) essentially coincident with acetabula; (c) well caudad to acetabula. CI = 0.67. 120. Recessus iliacus (Baumel 1979; a pneumatic pocket at caudal terminus of renal depression): present;(b) absent.CI = 1.0. (a)

APPENDIX 2.

A Linnean

classification

of the Recent

genera of Anseriformes.I follow the conventions of Wiley (1981), with the exceptionof the provisional recognition of the paraphyletic "Anatini"
(annotated incertaesedis).I have retained, where

possible,the names and taxonomicranks of previous classifications. Names of subtribesare given endingsof -eae after the namesfor the "sections"
of Boetticher (1952) and are derived from the old-

Furculum

101. Coracoidaltuberosities(U*): (a) present;(b) obsolete.CI = 0.25. 102. Furcular process: a flattened point; (b) variable,but reduced, (a)

est included genus.Sedis mutabilis follows taxa in which the order of included groupsis unresolved. * = two subfamiliesmay be in reverseorder; ** = subtribes possiblyare sistergroups;*** = probable
sister groups.
Order Anseriformes Anhimae Suborder

essentially continuous with curvatureof clavicles; swollen (c)

truncate lobe. CI = 1.0.

103. Clavicularsymphysis: without foramina;(b) with medial fo(a)

ramina. CI 1.0.

104. Clavicles: roughlycircularin cross-section; distinctlyflat(a) (b) tened antero-posteriorly. = 1.0. CI

105. Lateral surfacesof clavicles: (a) smooth, unperforated; (b) with depressions containing severalsmall foramina;(c) with depression containinglarge pneumaticforamen.CI 0.50.

Family Anhimidae
Genus Anhima

Genus

Chauna

Suborder
Genus

Anseres
Anseranas

Family Anseranatidae Family Anatidae Subfamily Dendrocygninae

Genus Dendrocygna

106. Regionof clavicularsymphysis: a continuous (a) smoothcurve; (b) markedlyextendedposterodorsally, forming a U-shapedaccommodation trachealloop and associated for modificationof carina. CI = 1.0.

SubfamilyThalassorninae*
Scapula
Genus Thalassornis

107. Coracoidal articulation: (a) flush with blade; (b) base protruding

SubfamilyAnserinae*
Tribe Anserini

ventrallyas roundedhump. CI = 1.0. 108. Taper (profile) of blade (U): (a) of uniform width or tapering continuously throughout length;(b) width maximalat midpoint;
(c) width maximal at terminus. (Cygnus, Olor.) CI = 1.0. 109. Coracoidal articulation:(a) equal to acromionin proximalextent;

Genus Cereopsis
Genus Genus Anser Branta

Tribe Cygnini
Genus Coscoroba

(b) distinctlydistalto acromion. = 1.0. CI 110. Internal surface, immediatelyposterior glenold facet:(a) essento tially smooth; having deepdepression. - 1.0. (b) CI 111. Anterior edge (U*): (a) containingpneumaticfossa; without (b)
pneumatic fossa.(Cairina.)CI = 0.25. 112. Dorsal surfaceof neck: (a) marked by single distinct raised at-

Genus Cygnus
Genus Olor

Subfamily Stictonettinae
Genus Stictonetta

tachmentscar;(b) markedby two prominentraisedattachment

scars. CI
Pelvis

1.0.

Subfamily Plectropterinae GenusPlectropterus Subfamily Tadorninae

Tribe Sarkidiornini

Genus Sarkidiornis Tribe Tadornini

Genus

sedis mutabilis

113. Preacetabulariliac fossa: smoothly curved surface;(b) contain(a) ing a deep, irregularly shapeddepression. = 1.0. CI 114. Caudal margin: (a) ischiumextending well caudadto ilium; (b) variable,but ischiumand ilium roughly equal in caudalextent,

Subtribe

Tadorneae
Tadorna

SubtribeMalacorhyncheae
Genus Malacorhynchus

forming an obliquelysloping margin, with elementstypically separated posteriorly a distinctnotch.CI - 1.0. by 115. Body of pubis (V): (a) concavedorsally (rarely almost straight); (b) convexdorsally.CI = 0.50. 116. Orientation of postischiac pubis:(a) directed posteriorly;(b) directedventrally. CI = 1.0. 117. Shapeof postischiac pubis:(a) of uniform width or evenly widening caudally;(b) widened into roughly circularflange,especially extensiveanteroventrally.CI = 1.0.

SubtribeChloephageae sedis mutabilis Genus Alopochen

Genus Neochen

Genus Chloephaga

Subtribe Cyanocheneae**
Genus Cyanochen SubtribeMerganetteae** Genus Hymenolaimus

754
APPENDIX 2. Continued GenusMerganetta GenusTachyeres SubfamilyAnatinae
[Tribe] "Anatini" incertaesedis
Genus Pteronetta
Genus Genus Cairina A/x

BRADLEYLIVEZE C.
APPENDIX 2. Continued
Tribe Mergini GenusPolysticta
Genus Somateria Genus Histrionicus

[Auk,Vol. 103

GenusCamptorhynchus
Genus Melanitta

GenusLophonetta Genus Nettapus

Genus Genus Genus Genus Anas Callonetta Chenonetta Amazonetta

GenusClangula GenusBucephala* ** GenusMergellus*** GenusLophodytes GenusMergus

Tribe Oxyurini
Genus Heteronetta

Tribe Aythyini
Genus Marmaronetta

GenusNomonyx GenusOxyura
Genus Biziura

Genus Rhodonessa
Genus Netta

Genus Aythya