phylogeny of recent canidae (mammalia, carnivora): relative reliability and utility of morphological...

© The Norwegian Academy of Science and Letters • Zoologica Scripta,

33

, 4, July 2004, pp311–333

311

Zrzav

y

, J. &

R

i

c

ánková, V. (2004). Phylogeny of Recent Canidae (Mammalia, Carnivora):relative reliability and utility of morphological and molecular datasets. —

Zoologica Scripta, 33

,311–333.Phylogenetic relationships within the Canidae are examined, based on three genes (cytb, COI,COII) and 188 morphological, developmental, behavioural and cytogenetic characters. Bothseparate and combined phylogenetic analyses were performed. To inspect the phylogenetic‘behaviour’ of individual taxa, basic phylogenetic analysis was followed by experimentalcladistic analyses based on different data-partition combinations and taxon-removal analyses.The following phylogeny of the Recent Canidae is preferred: (1)

Urocyon

is the most basalcanid; (2)

Vulpes

is a monophyletic genus (including

Fennecus

and

Alopex

); (3) the doglike canids(DC) form a clade (=

Dusicyon + Pseudalopex + Lycalopex + Cerdocyon + Atelocynus + Chrysocyon +Speothos + Lycaon + Cuon + Canis

), split into two subclades, South American and Afro-Holarctic,with uncertain position of the

Chrysocyon + Speothos

subclade; (4)

Canis

is paraphyletic due tothe position of

Lycaon

and

Cuon

.

Otocyon

and

Nyctereutes

are the most problematic canid genera,causing an unresolved branching pattern of

Otocyon

,

Vulpes

,

Nyctereutes

and DC clades. Reclas-sification of the two basal species of ‘

Canis

’ into separate genera is proposed (

Schaeffia

for‘

C

.’

adustus

,

Lupulella

for ‘

C

.’

mesomelas

). Although the morphological dataset ranked poorly inboth separate and simultaneous analyses (measured by number of minimum-length topolo-gies, relative number of resolved nodes in the strict consensus of all minimum-length topologies,consistency and retention indices, nodal dataset influence, and number of extra steps requiredby the data partition to reach the topology of the combined tree), the morphological synapo-morphies represent nearly one quarter of all synapomorphies in the combined tree. Amongthe hidden morphological support of the combined tree the developmental and behaviouralcharacters are conspicuously abundant.

Jan Zrzav

¥

& V

e

ra

‰

i

3

ánková, Department of Zoology, Faculty of Biological Sciences, Universityof South Bohemia, Brani

6

ovská 31, 370 05

1

eské Bud

e

jovice, Czech Republic. E-mail:[email protected], [email protected]

Blackwell Publishing Ltd.

Phylogeny of Recent Canidae (Mammalia, Carnivora): relative reliability and utility of morphological and molecular datasets

J

AN

Z

RZAV

Y

& V

E

RA

R

I

C

ÁNKOVÁ

Accepted: 6 August 2003

Introduction

The Canidae (Mammalia, Carnivora, Caniformia) is a diversegroup of carnivores comprising about 36 extant species and arich fossil record (Van Gelder 1978; Nowak 1999; IUCN/SSC Canid Specialist Group 2001; see Table 1). They areunique among mammals, exhibiting many unusual reproduc-tive and behavioural traits, e.g. monogamy with paternal care,long-term incorporation of young adults into the social group,alloparental care, reproductive suppression in subordinateindividuals, obligate pseudopregnancy, monoestrum, andcopulatory tie (see Moehlman 1989; Geffen

et al

. 1996;Moehlman & Hofer 1997; Asa 1997; Asa & Valdespino 1998;and references therein). Not surprisingly, the evolution ofsuch distinctive physiology, behaviour and social organizationhas attracted a lot of attention. There has been much discussion

of the role of the body mass in determining, via feeding ecology,canid social behaviour and reproduction (see Moehlman1989; Geffen

et al

. 1996; Moehlman & Hofer 1997). Unfor-tunately, the phylogenetic relationships of all big dogs arevery unclear (Bininda-Emonds

et al

. 1999), and it is still vir-tually impossible to formulate and test explicit hypothesesabout the evolutionary transformation of individual relevanttraits. Asa & Valdespino (1998) argued that the adaptive valueof monogamy, monoestrum and obligate pseudopregnancy ismost evident in social cooperative hunters who incorporateadult offspring in the group. Consequently, they hypothesizethat the degree of sociality in the ancestral canids was higherthan that of Recent foxes. Naturally, if the primitive canidswere not pack-living cooperative hunters, the adaptive valueof the unique canid reproduction remains equivocal. As

Phylogeny of Recent Canidae

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ánková

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speculating about the evolution of any trait without referenceto its historical context is pointless, providing a robust phy-logeny of the Canidae is clearly required to address questionsconcerning canid sociobiology and ecology.

Previously published phylogenetic analyses of canids havebeen based on morphology (Clutton-Brock

et al

. 1976; Berta1987; Tedford

et al

. 1995), cytogenetics (Wayne

et al

. 1987a,b),allozyme genetic distances (Wayne & O’Brien 1987), restric-tion sites and restriction fragments (Geffen

et al

. 1992), andmitochondrial DNA sequences (Geffen

et al

. 1992; Wayne

et al

. 1997). However, as shown by Bininda-Emonds

et al

.(1999; and references therein), the 35 canid phylogenies pub-lished between 1967 and 1995 have provided highly unstableresults (Fig. 1). Most clades in the canid supertree (Bininda-

Emonds

et al

. 1999), reconstructed with the ‘matrix represen-tation by parsimony’ (MRP) method, have low Bremer supportvalues, suggesting lack of and/or conflicting evidence.

At present, there are several datasets available for mostcanid species. They include morphology, behaviour, socialorganization and ecology, reproduction and development,cytogenetics, and nucleotide and amino acid sequences of threemitochondrial protein-coding genes, cytochrome

b

(henceforthcytb), cytochrome

c

oxidase subunit I (COI), and cytochrome

c

oxidase subunit II (COII). Phylogeneticists commonlydiscuss the relative utility of different systematic datasets (seeGatesy

et al

. 1999; and references therein); however, theutility of individual datasets for the reconstruction of canidphylogeny has never been tested explicitly. Some workers have

Table 1 List of Recent canid species, with the GenBank accession numbers.

Species Vernacular name cytb COI COII M

Alopex lagopus (Linnaeus, 1758) Arctic fox Geffen et al. (1992) — — +Atelocynus microtis (Sclater, 1883) small-eared zorro AF028135 + AF028159 AF028183 AF028207 +Canis adustus (Sundevall, 1847) side-striped jackal AF028136 + AF028160 AF028184 AF028208 +Canis aureus (Linnaeus, 1758) golden jackal AF028138 + AF028162 AF028186 AF028210 +Canis latrans (Say, 1823) coyote AF028164 + AF028140 AF028188 AF028212 +Canis lupus (Linnaeus, 1758) grey wolf AF028165 + AF028141 AF028189 AF028213 +Canis lycaon (Schreber, 1775)* east Canadian wolf — — — NACanis mesomelas (Schreber, 1775) black-backed jackal AF028166 + AF028142 AF028190 AF028214 +Canis rufus (Audubon & Bachman, 1851)* red wolf NA — — NACanis simensis (Rüppell, 1835) Ethiopian wolf AF028168 + AF028144 AF028192 AF028216 +Cerdocyon thous (Linnaeus, 1766) crab-eating zorro AF028169 + AF028145 AF028217 AF028193 +Chrysocyon brachyurus (Illiger, 1815) maned wolf AF028163 + AF028139 AF028187 AF028211 +Cuon alpinus (Pallas, 1811) dhole AF028161 + AF028137 AF028185 AF028209 +Dusicyon australis (Kerr, 1792) Falkland Island wolf — — — +Fennecus zerda (Zimmermann, 1780) fennec AF028170 + AF028146 AF028194 AF028218 +Lycalopex vetulus (Lund, 1842) hoary zorro AF028148 + AF028172 AF028196 AF028220 +Lycaon pictus (Temminck, 1820) African wild dog AF028147 + AF028171 AF028195 AF028219 +Nyctereutes procyonoides (Gray, 1834) raccoon dog AF028173 + AF028149 AF028197 AF028221 +Otocyon megalotis (Desmarest, 1822) bat-eared fox AF028174 + AF028150 AF028198 AF028222 +Pseudalopex culpaeus (Molina, 1782) culpeo AF028151 + AF028175 AF028199 AF028223 +Pseudalopex fulvipes (Martin, 1837) Darwin’s zorro — — — +Pseudalopex griseus (Gray, 1837) grey zorro AF028152 + AF028176 AF028200 AF028224 +Pseudalopex gymnocercus (Fischer, 1814) Azara’s zorro AF028153 + AF028177 AF028201 AF028225 +Pseudalopex sechurae (Thomas, 1900) Sechuran zorro AF028178 + AF028154 AF028202 AF028226 +Speothos venaticus (Lund, 1842) bush dog AF028179 + AF028155 AF028203 AF028227 +Urocyon cinereoargenteus (Schreber, 1775) grey fox AF028180 + AF028156 AF028204 AF028228 +Urocyon littoralis (Baird, 1857) island grey fox — — — NAVulpes bengalensis (Shaw, 1800) Bengal fox — — — +Vulpes cana (Blanford, 1877) Blanford’s fox Geffen et al. (1992) — — +Vulpes chama (Smith, 1833) Cape fox Geffen et al. (1992) — — +Vulpes corsac (Linnaeus, 1768) corsac fox Geffen et al. (1992) — — +Vulpes ferrilata (Hodgson, 1842) Tibetan sand fox — — — +Vulpes macrotis (Merriam, 1888)† kit fox AF028157 + AF028181 AF028205 AF028229 +Vulpes pallida (Cretzschmar, 1827) pale fox — — — +Vulpes rueppellii (Schinz, 1825) Rüppell’s fox Geffen et al. (1992) — — +Vulpes velox (Linnaeus, 1758)† swift fox NA — — +Vulpes vulpes (Linnaeus, 1758) red fox AF028182 + AF028158 AF028206 AF028230 +

cytb, cytochrome b; COI, cytochrome c oxidase subunit I; COII, cytochrome c oxidase subunit II; M, morphology; NA, not analysed. *For taxonomy of the C. lycaon/C. rufus complex see Wilson et al. (2000), Mech & Federoff (2002), and Nowak (2002). †The V. velox/V. macrotis complex is treated as a single terminal in the present analysis.

J. Zrzav

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Phylogeny of Recent Canidae

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examined a single dataset (often with a quite limited taxonomicsample), others a combination of several; in either case, littleattention has been paid to impact on the framework of simul-taneous (‘combined’, ‘total-evidence’) analysis. For example,Wayne

et al

. (1997) merged the three molecular sequences,concluded that the three genes evolved in a similar fashion, andcompared the resulting ‘molecular tree’ with previously pub-lished morphological, cytogenetic and allozyme-distancephylogenies, emphasizing the several conflicting points (e.g.placements of the raccoon dog, maned wolf and bush dog).Their molecular tree does indeed differ from the morpholog-ical ones (see Berta 1987; Tedford

et al

. 1995). However, thepossibility remains that the combined ‘molecular’ dataset isitself intrinsically heterogeneous. It is still possible that amorphological tree is congruent with one derived frommolecular datasets, and that any ‘morphological-molecular’conflict is, in fact, the product of internal conflict among the

molecular data. Morphological and molecular characters werecombined and analysed simultaneously by Wayne

et al

. (1997)and Muñoz-Durán (2002). Wayne

et al

. (1997) compared thecombined tree topology with those of the morphological andmolecular trees but did not study the relative influence ofindividual data partitions upon the final solution. Muñoz-Durán’s (2002) analysis has not yet been published

in extenso

.In the present paper we have set out to achieve the following:

1

Provide a well-corroborated hypothesis of canid species-levelrelationships based on the combination of all available nonse-quence (morphological, behavioural, developmental and cyto-genetic — abbreviated to ‘morphology’ or M in the figure andtable legends) and molecular sequence (cytb, COI, COII) datasets.

2

Identify ‘problematic species’ whose phylogenetic positionis unstable (due to lack of relevant information, highly conflict-ing signals in individual data partitions, and/or accumulationof numerous aberrant character states) and try to resolve theirphylogenetic placement.

3

Analyse behaviour of the four data partitions (morphology,cytb, COI, COII) in both separate and simultaneous analyses,and their relative influence on the combined-tree topology.

Materials and methods

Species and characters

All species of the Recent Canidae (= the single subfamilyCaninae) were included in the present analysis as ‘operationaltaxonomic units’, or terminals (see Table 1), except for

Urocyonlittoralis

(Wayne

et al

. 1991) and the problematic

Canis lycaon

/

C. rufus

complex (see Reich

et al

. 1999; Wilson

et al

. 2000;Mech & Federoff 2002; Nowak 2002). The

Vulpes velox

/

V

.

macrotis

complex (see Dragoo

et al

. 1990; Mercure

et al

. 1993;Maldonado

et al

. 1997) was treated as a single terminal. Extinctspecies were not included, with the exception of

Dusicyonaustralis

, which was exterminated as recently as 1876. Extinctcanid subfamilies Hesperocyoninae and Borophaginae, aswell as †

Leptocyon

(Caninae), were selected as morphologicaloutgroups (Tedford et al. 1995).

In total, 188 morphological, cytogenetic, and behaviouralcharacters were obtained from the literature (see Appendix 1).Some behavioural patterns were checked by original obser-vation of captive specimens of grey wolves, African wild dogs,bat-eared foxes, dholes, black-backed jackals, maned wolves,and bush dogs (zoos in Prague, Olomouc, Dvur Králové nadLabem, Munich, west and east Berlin). Where possible, datafrom wild populations were preferred over those from captiveones; original observations of the latter were only used whenneeded to fill gaps in the matrix resulting from insufficient orunclear information in the literature (‘orig. observ.’ in Appendix 1).All skeletal, soft tissue and colour pattern characters were lumpedwith cytogenetic, developmental, ecological and behaviouraldata and treated as a single dataset (‘morphology’). Naturally,there is no evident intrinsic reason why that heterogeneous

Fig. 1 Supertree of the Canidae, based on 35 phylogenetic hypotheses(1967–1995) combined by Bininda-Emonds et al. (1999) using the‘matrix representation by parsimony’ method. Bremer supports aregiven for each node. The original authors’ taxonomy is acceptedwithout changes.

Phylogeny of Recent Canidae • J. Zrzavy & V. ‰i3ánková

314 Zoologica Scripta, 33, 4, July 2004, pp311–333 • © The Norwegian Academy of Science and Letters

set of characters should be internally evolutionarily congruent.At present it is impossible to analyse the potential conflictamong ‘morphological’ subsets thoroughly because many speciesare not sufficiently well-represented to be readily comparable(behavioural and developmental characters of most foxes andSouth American canids are unknown).

Nucleotide sequences of cytb, COI, and COII were obtainedfrom GenBank (for accession numbers see Table 1) and fromEli Geffen (see Geffen et al. 1992). The available caniformcarnivorans were selected as outgroups: Ursus americanus (Ursi-dae), Phoca vitulina (Phocidae), Mustela putorius (Mustelidae),and Mephitis mephitis (Mephitidae) for cytb (GenBank accessionnumbers U34266, NC_001325, X94925, X94927, respectively;Phoca vitulina (Phocidae) for COI and COII (NC_001325).As they all are protein-coding genes of highly constrained lengths,the alignments were trivial (no gaps; see Wayne et al. 1997).

Data combination and character congruenceThe different data partitions to be combined in the simulta-neous analyses covered different species spectrums (33 canidspecies for morphology, 28 for cytb, 23 for COI and COII).We performed the following:1 Separate analyses for individual data partitions.2 Combined analyses of all morphological characters andsequences of all 33 taxa, introducing missing values for theabsent sequences (= ‘all-species strategy’).3 Combined analyses of all character partitions and 23 species,from which the taxa whose cytb, COI, and COII sequencesare unknown were removed (= ‘complete-species strategy’).In the ‘experimental’ phylogenetic analyses (see below) the23-species datasets were preferred.

As different partitions were rooted by different outgroups(Hesperocyoninae + Borophaginae + Leptocyon for osteology,noncanid caniforms for cytogenetics, four noncanid caniformsequences for cytb, seal sequences for COI and COII, noavailable outgroup for development, ecology, and behaviour),a single combined ‘outgroup’ was constructed as a strict con-sensus of all outgroup characters.

Phylogenetic analysisThe maximum parsimony analysis was applied to morpho-logical, sequence and combined data matrices (NONA version2.0; Goloboff 1999: heuristics, option ‘hold100000 mult*100hold/1000’, unconstrained ‘mult*max*’ search strategy). Unlessotherwise stated, all characters were equally weighted(morphology:molecules = 1:1, transition:transversion = 1:1).Bremer (decay) indices of branch support (BS) were calculated(NONA: option ‘bsupport100000’) as the difference inlength between the shortest topology that lacks the node ofinterest and the shortest topology that contains it. For the 4-partition combined tree, bootstrap support (1000 replications)was calculated as well. In discussing our results, we refer to BS

values giving weak (1−2), moderate (3−5), good (6−10) andstrong (> 10) support, even though delimiting such qualitativeclasses is inherently subjective. However, all BS values > 20correspond to bootstrap values of > 95% (BS ≥ 15 correspondsto bootstrap > 90%) in the current dataset. For characteroptimization, the ‘unambiguous’ option in NONA was applied.

To investigate the topological changes caused by changesof analytical focus a few experimental cladistic analyses wereperformed. The parameter values of the experiments per-formed here included:1 Fifteen different data-partition combinations (four 1-partition, six 2-partition, four 3-partition, one 4-partition).2 Two different taxon-representing strategies (‘all-species’vs. ‘complete-species’; see above).3 Presence vs. absence of the noncanine outgroups (rootedvs. unrooted analyses).

Relative quality of character partitions in separate and simultaneous analysesThe four character partitions were ranked via four criteriadescribing the separate-partition trees (all tests were madefor the datasets including 23 canid taxa plus a single outgroup).They included:1 Number of minimum-length topologies (Ntree).2 Consistency index (CI, with uninformative charactersexcluded).3 Retention index (RI, with uninformative characters excluded).4 Number of resolved nodes in the strict consensus of allminimum-length topologies divided by the number of nodesin the completely resolved tree (%node).

The same criteria were studied in the 2-, 3- and 4-partitioncombined trees, to test whether combination improved someof the data quality measures. In order to analyse the relativecontribution of individual data partitions to the outcome ofthe simultaneous analysis, two measures were analysed in thecomplete, 4-partition combined tree:1 Nodal dataset influence (NDI), the BS score for a node inthe 4-partition combined tree, minus the score for the node inquestion in the 3-partition combined tree without the partitionunder investigation.2 Congruence of separate and simultaneous trees was mea-sured by the number of extra steps required by the datapartition to reach the topology of the combined tree.

Another measure of the relative performance of individualdata partitions within the simultaneous analysis was the numberof the individual partition’s synapomorphies in the 4-partitioncombined tree.

ResultsDifferent data-partition combinations and canid phylogenyIn the morphological tree (Fig. 2), the foxlike canids (truefoxes, grey fox and bat-eared fox) form a basal stem lineage

J. Zrzavy & V. ‰i3ánková • Phylogeny of Recent Canidae

© The Norwegian Academy of Science and Letters • Zoologica Scripta, 33, 4, July 2004, pp311–333 315

(possibly paraphyletic), and the doglike canids (DC) repre-sent a clade including also the raccoon dog. Within the latter,South American canids (‘zorros’, bush dog and maned wolf)are paraphyletic. The dataset limited exclusively to skeletalcharacters (characters #0−68; similar to the datasets pub-lished by Berta 1987; Tedford et al. 1995) provides 14 trees(length 234, CI 0.41, RI 0.63) in which Speothos groups withCuon and Lycaon (= ‘simocyonine clade’) among the Afro-Holarctic DCs. In the cytb tree (Fig. 3), Urocyon is a sistergroup of all other canids and the DC form a well-supportedclade. Within the latter, only Atelocynus + Cerdocyon + Pseuda-lopex + Lycalopex and Speothos + Lycaon subclades are well sup-ported. The trees derived from COI and COII sequences(Figs 4, 5) share the strongly supported Chrysocyon + Speothosclade, closely related either to the South American (COI), or tothe Afro-Holarctic (COII) canids. In both trees the monophyly

of the DC is relatively well-supported. The strict consensustree derived from the separate analyses is entirely unresolved;only two uncontroversial clades (Vulpes + Fennecus and V. vulpes+ V. velox) are present in their semistrict consensus tree.

The combined morphological-molecular tree (Fig. 6) sug-gests that Urocyon, Otocyon and Vulpes are successive basal out-groups of all other canids. The DC split into South Americanand Afro-Holarctic subclades. Chrysocyon and Speothos arestrongly supported as closely related to each other and bothrepresent a sister group of the South American clade; Lycaonand Cuon form a strongly supported clade within the para-phyletic Canis in the Afro-Holarctic group (Canis adustus andC. mesomelas are the basalmost Afro-Holarctic dogs). Nyc-tereutes is a weakly supported sister group of the DC clade.

The 15 trees based on various combinations of individualdata partitions (four 1-partition, six 2-partition, four 3-partition,one 4-partition) provide a wide spectrum of possible canid

Fig. 2 Strict consensus of 16 trees based on morphological, develop-mental, ecological, behavioural, and cytogenetic data (184 cladisticallyinformative characters, length 659, CI 41, RI 53). The tree is derivedfrom the 23-species dataset. Bremer supports are given for eachnode.

Fig. 3 Single most parsimonious tree found for the cytochrome bdataset (235 cladistically informative characters, length 976, CI 0.36,RI 0.47). The tree is derived from the 23-species dataset. Bremersupports are given for each node.



clades (Fig. 7). The ‘stability tree’, a majority-rule consensus treeincluding all clades that were present in more than 50% (= eightor more) of the source trees, is relatively well resolved (Fig. 8):1 Urocyon is a sister group of all Recent Canidae (67%).2 The DC form a clade (80%) split into South American(Chrysocyon, Speothos, Atelocynus, Cerdocyon, Pseudalopex, Lycalopex;53%) and Afro-Holarctic (Cuon, Lycaon, Canis; 60%) subclades.3 Within the South American canids, two subgroups arepresent: Chrysocyon + Speothos (80%) and unresolved Atelocynus+ Cerdocyon + Pseudalopex + Lycalopex (67%).4 The Afro-Holarctic subclade is unresolved; the ‘higherCanis’ (= Canis less C. adustus and C. mesomelas) is marginallysupported as monophyletic (53%), while the traditional Caniss.l. is not present in any tree.5 Positions of Nyctereutes, Vulpes and Otocyon among thenon-Urocyon canids are unresolved.

The stability tree only differs from the ‘total evidence’combined tree in its lower resolution; all components are

present in the 4-partition combined tree as well. Of the eightclades present in the stability tree, none is present in the mor-phological tree, three are present in the cytb tree, six in theCOI tree, and five in the COII tree. Both cytochrome c oxi-dase subunit genes are then most influential sources of thecombined tree topology, whereas morphology is the leastsuccessful partition in this respect.

‘All-species’ vs. ‘complete-species’ strategies in the simultaneous analysesIn the present analysis, five species (Dusicyon australis, Pseudalopexfulvipes, Vulpes bengalensis, V. ferrilata and V. pallida) are repre-sented exclusively by morphological characters. Five more(Alopex lagopus, Vulpes cana, V. chama, V. corsac and V. rueppellii)are represented by morphological data and fragments of cytbsequences (COI and COII not available). In the all-species4-partition tree (Fig. 9), D. australis and P. fulvipes are placedwithin the unresolved Pseudalopex + Lycalopex + Dusicyon clade,

Fig. 4 Single most parsimonious tree found for the cytochrome csubunit I dataset (180 cladistically informative characters, length653, CI 0.36, RI 0.54). Bremer supports are given for each node.

Fig. 5 Single most parsimonious tree found for the cytochrome csubunit II dataset (194 cladistically informative characters, length689, CI 0.37, RI 0.49). Bremer supports are given for each node.



V. cana as a sister species of Fennecus zerda, V. pallida as a sistergroup of the Vulpes + Alopex + Fennecus clade as a whole, andall other fox species group within the unresolved Vulpes +Alopex + Fennecus clade.

To test the possibility that simultaneous inclusion of all‘incompletely known’ taxa within the combined dataset mightdistort relationships due to too many ‘missing characters’, 10trees including 23 species represented by all data partitionsplus one incompletely known species were constructed. Thelatter group at positions identical to (or compatible with)those situated in the all-species tree, with a single exception.The pale fox (V. pallida) is a sister group of all other true foxesin the all-species tree, and a member of the Afro-Asiatic foxsubclade (V. chama + V. cana + F. zerda + V. ferrilata + V. pallida)in the 24-species tree.

Relative quality of character partitions and their performance in the combined analysisAll molecular partitions share good tree resolution (a singletopology each); morphology, in contrast, ranked highly forlow within-partition homoplasy (high CI and RI) and poorlyfor tree resolution and node support (see Table 2). Combina-tion of data partitions did not improve some of the qualitymeasures because of interpartition character conflict. The 4-partition combined tree has low resolution (although betterthan the morphological tree), low CI and RI. With one par-tition removed, the 3-partition combined trees have betterresolution, and have better or, at worst, equal CI and RI.

Total Bremer support of the combined tree (i.e. the sum ofall BS scores for all nodes supported by the dataset, TBS =197) contains highly unequal contributions from the individualpartitions. Only two of the 17 nodes present in the 4-partitioncombined tree are present in the morphological tree as well,and the sum of their scores represents 23.1% of TBS of themorphological tree (‘proBS’ in Table 2). In the moleculartrees, the numbers of clades present in the combined tree areconspicuously higher: seven in cytb (proBS = 52.9%), 10 inCOI (78.3%), and eight in COII (73.5%).

Similarly, morphology requires 54 extra steps (= 8.2%) toreach the topology of the combined tree while 32 (= 3.3%),20 (= 3.1%), and 27 (= 3.9%) extra steps are sufficient forcytb, COI and COII, respectively (Table 2).

As measured by the dataset influence (DI), morphology isclearly the least influential partition (DI = 35), while themolecular data partitions are more important sources of thecombined tree topology (DI = 55 for cytb, 85 for COI, and 77for COII; Table 3).

Character support in the combined treeWhile formal inference of the quality of individual data par-titions shows that morphology performs less well than any ofthe molecular partitions both in separate and simultaneousanalyses, its real performance in the context of the 4-partitioncombined tree is not so poor. Of 542 synapomorphies ofthe supraspecific canid taxa in the strict consensus of theminimum-length 23-species combined trees (Table 3, Appendix2A,B), 23.2% originate from morphology (12.4% osteology,4.8% soft anatomy, 4.6% reproduction, development andbehaviour, and 1.5% cytogenetics), 27.1% from cytb, 27.9%from COI, and 21.8% from COII. Morphology is even moresuccessful if only the 115 nonhomoplastic synapomorphiesare counted (morphology 33.9%, cytb 28.7%, COI 17.4%,COII 20.0%).

DiscussionComparison of trees from 15 different data combinations(Figs 2−8) reveals the following:1 The basalmost position of Urocyon.

Fig. 6 Strict consensus of four trees from the combined morpho-logical, cytb, COI and COII characters (793 cladistically informativecharacters, length 3106, CI 0.35, RI 0.47). The tree is derivedfrom the 23-species dataset. Bremer supports are given for eachnode.



2 Monophyly of the DC (Cerdocyon, Atelocyon, Pseudalopex,Lycalopex, Dusicyon, Chrysocyon, Speothos, Canis, Lycaon, Cuon).3 Monophyly of the ‘zorro’ subclade (Cerdocyon + Atelocyon +Pseudalopex + Lycalopex + Dusicyon).4 Monophyly of the Afro-Holarctic subclade (Canis + Cuon+ Lycaon).5 Relationships between basal canid genera (Otocyon, Vulpes,Nyctereutes) and the DC clade, and the phylogenetic positionof Chrysocyon and Speothos are not well established.

If the ‘outgroup’ is removed from the combined dataset,the resulting unrooted tree topology is comparable to that of

the rooted combined tree. The problem of outgroups beingtoo distant (Wheeler 1990) does not apply to the phylogenyof Canidae. The traditional genera of Canidae are mono-phyletic with three exceptions:1 Vulpes s.s. is paraphyletic because Alopex lagopus is a sisterspecies of the V. velox/V. macrotis complex and Fennecus zerdais a sister species of V. cana (see Geffen et al. 1992).2 Lycalopex vetulus does not represent a separate canid lineageand is closely related to the species of Pseudalopex, namely toP. sechurae. The position of D. australis is less certain but itmay be a sister species of P. culpaeus (see below).

Fig. 7 A–J. Simplified phylogenies of theCanidae based on different 2- and 3-partitioncombinations. All trees derived from the 23-species analyses. Double lines indicate thegroup is paraphyletic and/or unresolved.Abbreviations: ATE, Atelocynus; CAD, Canisadustus; CAN, ‘higher Canis’; CER, Cerdocyon;CHR, Chrysocyon; CME, Canis mesomelas; CUO,Cuon; LYC, Lycaon; NYC, Nyctereutes; OTO,Otocyon; PSE, Pseudalopex + Lycalopex clade;SPE, Speothos; URO, Urocyon; VUL, Vulpes(incl. Fennecus and Alopex). —A. M + cytb.—B. M + COI. —C. M + COII. —D. cytb+ COI. —E. cytb + COII. —F. COI + COII.—G. M + cytb + COI. —H. M + cytb+ COII. —I. M + COI + COII. —J. cytb+ COI + COII.



3 Cuon and Lycaon appear to be nested within paraphyleticCanis s.l., while Canis adustus and probably also C. mesomelasare more basal members of the Afro-Holarctic dogs.

Grey foxesThe basalmost position of Urocyon is supported by 32 molec-ular synapomorphies (Table 3). This topology is present incytb and COII trees, in three 2-partition trees (M-COII,cytb-COI, cytb-COII) and in all 3- and 4-partition combinedtrees. On the other hand, no single morphological synapo-morphy supports the monophyly of the non-Urocyon canids(Appendix 2A). The alternative topology, congruent with themorphological dataset (Otocyon + Urocyon, placed within theparaphyletic Vulpes at the base of the canid tree), is supportedby 14 synapomorphies (including 5 nonhomoplastic, viz. 3odontological and 2 cytogenetic); in the combined tree, thesecharacters had to be reinterpreted as plesiomorphic for theCanidae as a whole (or ambiguous). However, when Urocyon

is removed from the combined analysis, the topology of thetree and the character-state optimization are not influenced,so that there is no indication that its basalmost position isartifactual. Although the branch support of the non-Urocyoncanid clade is low (BS = 3), this clade is sufficiently robust tocladistic experiments.

Bat-eared foxThe phylogenetic position of the bat-eared fox, Otocyonmegalotis, is highly uncertain. Urocyon, Nyctereutes, Vulpes, DC,and the Urocyon + Nyctereutes + DC and Vulpes + Nyctereutes +DC superclades were identified in different analyses as potentialsister groups. In the combined analysis, Otocyon groups as a basalcanid just above Urocyon; the clade including all canids lessUrocyon and Otocyon is very weakly supported (BS = 1), with 17synapomorphies (8 morphological, 4 cytb, 4 COI and 1 COII).However, if the transversions are weighted over transitions

Fig. 8 Simplified ‘stability tree’ of the Canidae, a majority-rule consensustree including all clades that were present in more than 50% of the15 1-, 2-, 3- and 4-partition source trees (see Figs 2−5, 6, 8). Frequencylevels are given for each clade including more than one species.

Fig. 9 Strict consensus of 12 trees found for the combinedmorphological, cytb, COI and COII data (806 cladistically informativecharacters, length 3343, CI 0.34, RI 0.48). The tree is derived fromthe 33-species dataset. Bremer supports are given for each node.



(2:1, 3:1; morphology and molecules 1:1), the Nyctereutes +Otocyon clade emerges as a sister group of DC (not shown),further emphasizing the highly unstable position of Otocyon.

True foxesThe monophyly of true foxes (Vulpes s.l. including Fennecusand Alopex) is supported by all analyses (unresolved only inthe morphological tree); character support is good (BS = 10,26 synapomorphies including four morphological). Theinternal hierarchical structure of the Vulpes clade is difficult toassess because only three species (F. zerda, V. velox/macrotis,V. vulpes) are represented by all three molecular partitions,and three (V. pallida, V. bengalensis, V. ferrilata) are representedexclusively by the morphological data.

The combined tree including all 33 species is quite unre-solved: only V. pallida is the sister species of Vulpes as a whole,and small North African-Middle East foxes, F. zerda + V. cana,form a clade. The 28-species combined tree (including thespecies for which at least fragment of one molecule is avail-able) is well-resolved, with two subclades of Vulpes: one includ-ing V. chama and F. zerda + V. cana, the other V. corsac + V.rueppellii + V. vulpes and V. velox/macrotis + A. lagopus. Includ-ing the individual ‘nonmolecular’ species to the 28-speciestree establishes no specific relationships for V. ferrilata andV. bengalensis, but joins V. pallida with V. chama. Our preliminaryconclusion is that two major groups of true foxes may exist:the ‘Afro-Asiatic’ clade (V. chama, V. pallida, F. zerda, V. cana,possibly also V. bengalensis); and the ‘Holarctic’ clade (V. rueppellii,

V. corsac, V. vulpes, V. velox/macrotis, A. lagopus, probablyV. ferrilata). It seems, however, highly improbable that thecontroversial taxonomy of the North American desert foxes,V. macrotis and V. velox, would affect the interrelationshipsamong true foxes. Both taxa are regarded as conspecific bysome authors because of reported interbreeding (see Rohwer& Kilgore 1973; Dragoo et al. 1990), but they are largely para-patric and both are probably monophyletic (Maldonado et al.1997; see Stromberg & Boyce 1986; Mercure et al. 1993).

In their maximum parsimony (MP) and maximum likeli-hood (ML) analyses of the Vulpes cytb sequences, Geffen et al.(1992) show F. zerda + V. cana, V. velox + A. lagopus, and V. velox+ A. lagopus + V. vulpes + V. corsac + V. rueppellii representingthree robust clades. The position of V. rueppellii within thelast clade is unstable, as well as that of V. chama within thegenus as a whole. A similar branching pattern is also corrob-orated by the MPA of the presence/absence matrices ofshared restriction fragments and of shared restriction sites(Geffen et al. 1992). Both support monophyly of the clades ofV. corsac + (V. rueppellii + V. vulpes), V. velox + A. lagopus, sister-grouprelationships between both, and a basal position for F. zerda +V. cana. Again, V. chama is in an uncertain position, as either thebasalmost species (restriction fragments), or as sister speciesto all foxes excluding F. zerda and V. cana (restriction sites).

Monophyly of Vulpes suggests that the basal cytogeneticdivergence of canids into ‘low numbered metacentric species’(Nyctereutes, Vulpes s.s.) and ‘high numbered acrocentric spe-cies’ (Otocyon, Urocyon, DC, Fennecus) (Wayne et al. 1987a,b)

Table 2 Quality of data partitions in separate and simultaneous (3- and 4-partition) trees TI (see Figs 2−7).

comb M cytb COI COII non-M non-cytb non-COI non-COII

Npart 4 1 1 1 1 3 3 3 3Ntax 23 23 23 23 23 23 23 23 23Nchar* 793 184 235 180 194 609 558 613 599Ntree 4 16 1 1 1 1 1 2 4length 3106 659 976 653 689 2378 2090 2434 2404CI 0.35 0.41 0.36 0.36 0.37 0.35 0.36 0.36 0.36RI 0.47 0.53 0.47 0.54 0.49 0.48 0.49 0.46 0.47%node 81.0 71.4 100.0 100.0 100.0 100.0 100.0 95.2 90.5TBS 197 39 85 83 102 241 177 168 140proBS 197 9 45 65 75 201 154 143 120%proBS 100.0 23.1 52.9 78.3 73.5 83.4 87.0 85.1 85.7extrasteps — 54 32 20 27 — — — —%extrasteps — 8.2 3.3 3.1 3.9 — — — —

M, morphology; Cytb, cytochrome b; COI, cytochrome c oxidase subunit I; COII, cytochrome c oxidase subunit II; comb, M + cytb + COI + COII; non-M, cytb + COI + COII; non-cytb, M + COI + COII; non-COI, M + cytb + COII; non-COII, M + cytb + COI; Npart, number of data partitions included; Ntax, number of taxa; Nchar, number of characters; Ntree, number of minimum-length topologies; length, length of the minimum-length topologies; CI, consistency index; RI, retention index; %node, number of resolved nodes in the strict consensus of all minimum-length topologies divided by the number of nodes in the completely resolved tree; TBS, total Bremer support (the sum of all BS scores for all nodes supported by the dataset); proBS, the sum of all BS scores for all nodes supported by the dataset that are present in the 4-partition tree (comb); %proBS, the sum of all BS scores for all nodes supported by the dataset that are present in the 4-partition tree (comb) divided by TBS (i.e. the portion of TBS of the data partition tree that is congruent with the combined tree); extrasteps, number of extra steps required by the data partition to reach the topology of the 4-partition combined tree; %extrasteps, the relative number of extra steps necessary to reach the topology of 4-partition tree (extrasteps divided by length).*only 184 morphological characters are cladistically informative in the 23-species dataset.



is not supported by the combination of several datasets (seealso Graphodatsky et al. 2000, 2001; Nash et al. 2001; Pienkowskaet al. 2002). The similarity of karyotypes of the fennec fox,grey fox, and bat-eared fox are most probably convergent ifnot plesiomorphic (both scenarios seem to be supported byoptimization of individual cytogenetic characters on thecombined tree). However, the karyotypes of most Vulpesspecies are unknown, which precludes making a definitiveconclusion at this time.

Raccoon dogsThe raccoon dog is, phylogenetically, one of the most prob-lematic canids. In the 4-partition combined tree, Nyctereutesis a sister group of the DC, while in the morphological tree,it is deeply nested within the DC clade (close to Atelocynus,Speothos and Cerdocyon; Berta 1987 and Tedford et al. 1995).The cytb tree supports its basal position, the COI tree its

sister-group relationship to the DC and the COII tree its prox-imity to Otocyon. Similar instability appears in the 2- and 3-partition trees as well. Branch support of the Nyctereutes + DCsuperclade shown in the combined tree is very low (BS = 1).On the other hand, it is supported by 14 morphological, 6cytb, 14 COI and 2 COII characters, although most of theseare homoplastic except for 11 morphological synapomor-phies (including posterior expansion of paroccipital process,morphology of frontal process and orbital border of thezygoma, enlarged mastoid process, shortened ears, monogyny,and orientation of agonistic behaviour towards the neck; seeAppendix 2A) and one COI synapomorphy. It is evidentthat placement of the raccoon dog is heavily influenced bymorphological characters; however, the molecular partitionsprovide no decisive information about the position of Nyctereutes.

Moreover, the raccoon dog is most probably a compositeof at least two allopatric species that profoundly differ in

Table 3 Relative performance of data partitions in the combined tree (see Fig. 6 and Appendix 2).

clade

M cytb COI COII comb

sBS NDI Ns Nsnh sBS NDI Ns Nsnh sBS NDI Ns Nsnh sBS NDI Ns Nsnh BS Ns Nsnh %bs

UcinSG −4 −9 0 0 6 +5 13 12 −5 −5 9 6 2 +3 10 6 3 32 24 52OmegSG 0 +9 8 5 −3 +3 4 2 −5 +9 4 3 −2 0 1 1 1 17 11 < 50Vulpes 0 0 4 1 1 +5 5 2 1 +4 7 3 5 +5 10 3 10 26 9 87Vvul + Vvel 0 −4 5 0 5 +17 14 3 11 +14 23 0 13 +21 18 3 37 60 6 99Npro + DC 4 +8 14 11 −3 −4 6 0 4 +16 14 1 −4 −11 2 0 1 36 12 51DC −7 −6 4 3 10 +14 17 7 8 +3 10 0 7 +13 12 3 22 43 13 99South Am. −10 +3 6 2 −23 −4 4 0 4 −2 9 0 10 −2 2 0 2 21 2 72Sven + Cbra −14 −5 9 3 −30 +2 18 2 22 +19 21 2 20 +24 27 5 44 75 12 100zorros −11 −14 4 0 7 −3 7 1 6 +10 11 3 10 −4 4 0 4 26 4 74Ps. + Lyc. −3 +1 8 2 13 +12 21 1 1 +5 13 0 10 +26 10 1 27 52 4 100Afro-Hol. 0 +14 12 6 −19 −1 7 1 4 +4 8 0 −10 0 2 0 15 29 7 91CaduSG −1 +2 4 0 −19 −1 8 1 1 −6 9 1 −6 −3 3 0 1 24 2 63CmesSG −2 +12 7 2 −17 −3 1 0 −6 −5 3 0 −6 −1 3 0 1 14 2 < 50Lpic + Calp 5 +23 28 3 −15 −9 4 0 −6 +4 3 0 −6 0 6 1 15 41 4 92hCanis −10 −9 4 1 −2 +10 4 0 3 +9 5 1 8 +5 7 0 7 20 2 80CaurSG −10 +6 4 0 3 +7 9 1 −2 +6 1 0 −3 +2 0 0 4 14 1 53Clat + Csim −5 +4 5 0 −2 +5 5 0 −1 0 1 0 −1 −1 1 0 3 12 0 < 50Di 35 55 85 77

M, morphology; cytb, cytochrome b; COI, cytochrome c oxidase subunit I; COII, cytochrome c oxidase subunit II; comb, M + cytb + COI + COII; sBS, Bremer support of the combined-tree node in the separate analysis of the partition; NDI, nodal data influence (the BS score for a node in the combined tree, minus the BS score for that node in the combined tree without the partition); Ns, number of partition synapomorphies in the combined tree (for morphological synapomorphies see Appendix 2); Nsnh, number of partition nonhomoplastic synapomorphies in the combined tree (for morphological synapomorphies see Appendix 2); BS, Bremer support of the node in the combined tree; %bs, bootstrap support value (in percentage, 1000 replications); DI, dataset influence (the sum of Bremer support scores for all nodes in the combined tree, minus the sum of BS scores for all nodes in the combined tree without the partition).Clades present in the 23-species 4-partition combined tree (comb): UcinSG, Urocyon sister group (= monophyly of Otocyon + Vulpes + Nyctereutes + DC); OmegSG, Otocyon sister group (= monophyly of Vulpes + Nyctereutes + DC); Vulpes, monophyly of Vulpes clade (incl. Fennecus); Vvul + Vvel, monophyly of Vulpes clade less Fennecus; Npro + DC, monophyly of Nyctereutes + DC, DC, monophyly of the ‘dog-like’ clade (Speothos + Chrysocyon + Atelocynus + Cerdocyon + Pseudalopex + Lycalopex + Dusicyon + Canis + Lycaon + Cuon), South Am., monophyly of the South American clade (Speothos + Chrysocyon + Atelocynus + Cerdocyon + Dusicyon + Pseudalopex + Lycalopex), Sven + Cbra, monophyly of the Speothos + Chrysocyon clade; zorros, monophyly of the ‘zorro’ clade (= Atelocynus + Cerdocyon + Dusicyon + Pseudalopex + Lycalopex); Ps. + Lyc., monophyly of the Dusicyon + Pseudalopex + Lycalopex complex; Afro-Hol., monophyly of the Afro-Holarctic clade (Canis + Lycaon + Cuon); CaduSG, Canis adustus sister group (= monophyly of C. mesomelas + Lycaon + Cuon + ‘higher Canis’); CmesSG, Canis mesomelas sister group (= monophyly of Lycaon + Cuon + ‘higher Canis’); Lpic + Calp, monophyly of the Lycaon + Cuon clade; hCanis, monophyly of ‘higher Canis’ (C. aureus + C. lupus + C. simensis + C. latrans); CaurSG, C. aureus sister group (= monophyly of C. lupus + C. latrans + C. simensis); Clat + Csim, monophyly of C. latrans + C. simensis.



cytogenetics (Ward et al. 1987; Wurster-Hill et al. 1988;Wada et al. 1998): viz., mainland N. procyonoides (Gray 1834)with 2n = 54 and Japanese N. viverrinus (Temminck, 1839),or tanuki, with 2n = 38. Both taxa differ also in skull andteeth morphology, ecology and behaviour (Kauhala et al.1998).

Doglike canids (DC)Monophyly of the DC clade, including all Recent canidsexcept Urocyon, Otocyon, Vulpes and Nyctereutes, is corrobo-rated by all trees excepting the purely morphological tree andthe combined M-COI tree (where the raccoon dog is placedwithin the clade). It is supported by 43 synapomorphies andhas strong branch support (BS = 22). However, there are onlyfour morphological synapomorphies: high neocortex ratio,high neonate mass, straight and horizontal tail posture indominant animals, and absence of metacentric chromosomes(which seem to reappear in Cerdocyon) (Appendix 2A). Thebulk of the character support of this clade is based on mole-cular characters (predominantly cytb and COII). It is highlyrobust and is present in the stability tree (80%) as well.

South American canidsThe South American subclade of the DC (Chrysocyon, Speot-hos, Cerdocyon, Atelocynus, Dusicyon, Pseudalopex, Lycalopex) isone of the more weakly supported components of the com-bined tree (BS = 2). It is present also in the COI tree as wellas in the M-COII, cytb-COI, COI-COII, M-cytb-COI, M-cytb-COII, and M-COI-COII combined trees. It is corrob-orated by 21 synapomorphies (6 morphological, 4 cytb, 9COI, 2 COII). The few morphological synapomorphies arerather dubious, as they apply predominantly to developmen-tal characters (in general, they suggest ‘retardation’ of someontogenetic processes; see Appendix 2A), which have not yetbeen studied in most of the species of interest.

The problematic nature of the South American subclade iscaused by the uncertain and conflicting position of Chrysocyonand Speothos (see below). The remaining five genera (Cerdo-cyon, Atelocynus, Dusicyon, Pseudalopex and Lycalopex) usuallyform a ‘zorro clade’. Monophyly of this group is primarilyderived from cytb, COI and COII sequences but remains inmost combined analyses. However, in some combined trees(M-COII, M-cytb-COII, M-COI-COII), the Speothos +Chrysocyon clade is more deeply nested among the SouthAmerican zorros as a sister group of Atelocynus (together withNyctereutes in M-COI). Phylogenetic relationships within the‘zorro clade’ are unclear: Atelocynus is either a sister specieseither of the Dusicyon + Pseudalopex + Lycalopex complex,leaving Cerdocyon as a basal genus (COII, M-cytb, cytb-COII,COI-COII, M-cytb-COI, cytb-COI-COII), or the basalmostzorro (cytb, COI, cytb-COI). If the transversions are weightedover transitions (2:1, 3:1), Atelocynus and Cerdocyon form a clade.

Phylogenetic relationships among the species of the Dusicyon+ Pseudalopex + Lycalopex complex are uncertain owing to absenceof molecular information in D. australis and P. fulvipes. In the all-species combined tree, there is no hierarchical structure withinthe genus except for the P. sechurae + L. vetulus clade (see Berta1987), which is retained also in the 23-species and ‘experimental’trees. Most of partition combinations trees include the P. griseus+ P. gymnocercus + P. culpaeus clade. When the ‘nonmolecular’species were appended to the 23-species combined tree, P. fulvipesgroups with P. griseus and D. australis with P. culpaeus. Two groupsof the Dusicyon + Pseudalopex + Lycalopex complex therefore appearto be corroborated: (1) P. sechurae + L. vetulus; (2) D. australis(?)+ P. culpaeus + P. gymnocercus + P. griseus + P. fulvipes.

Maned wolf and bush dogIn the combined tree, Speothos and Chrysocyon form a cladewith very strong branch support (BS = 44), corroborated by75 synapomorphies (9 morphological, 18 cytb, 21 COI, 27COII). The few morphological synapomorphies are quitedubious, including shortened tail, straight caecum, dark-coloured legs, agouti-coloured dorsal guard hairs, course ofpostembryonic development, and loss of chromosome 22(Appendix 2A). Not surprisingly, this clade is not present inthe purely morphological tree (morphologically, Speothosbelongs to South American canids and Chrysocyon to theAfro-Holarctic clade as its single Neotropical representative;Fig. 2). It is also absent in the cytb tree, but is supported byCOI, COII and most combined trees (excluding M-cytb).Overall, the Speothos + Chrysocyon clade is almost exclusivelybased on very strong COI and COII evidence.

The phylogenetic position of the clade is uncertain. Itbelongs to the South American canids in the COI, M-COI(with Nyctereutes), M-COII, cytb-COI, COI-COII, M-cytb-COI, M-cytb-COII, and M-COI-COII trees, and to theAfro-Holarctic canids in the COII, cytb-COII, and cytb-COI-COII trees. Its position within the latter appears to besupported by five morphological characters (including mor-phology of temporal ridges, female body mass ≥ 6 kg, laterattainment of adult body mass, and a derived form of playwith prey). In the absence of Chrysocyon, Speothos groups withLycaon within the Afro-Holarctic clade in the 4-partitioncombined tree; by contrast, in the absence of Speothos, theposition of Chrysocyon is uncertain within the clade. It appearsthat Speothos is strongly attracted to Chrysocyon (predomin-antly by the COI and COII sequences) as well as to socialhypercarnivorous Afro-Holarctic dogs (by morphology).

The taxon-removal approach therefore appears to supportclose relationships of the bush dogs with two specialized car-nivorous and highly social species, Lycaon and Cuon (all threeforming the traditional ‘Simocyoninae’; see Nowak 1999). Onthe contrary, after removal of Lycaon, Cuon and Chrysocyon fromthe combined tree, the bush dog groups with the ‘zorro clade’.



Moreover, all Speothos fossil remnants are known from SouthAmerican deposits while Lycaon and Cuon fossils are knownexclusively from the Old World (McKenna & Bell 1997; andreferences therein). At present, we consider Chrysocyon andSpeothos extremely divergent sister genera, representingaberrant members of the South American clade, althoughtheir position deserves further attention.

Afro-Holarctic canidsMonophyly of the clade including several species of Canis s.l.,together with the African wild dog (L. pictus) and the dhole(C. alpinus), is supported by 29 synapomorphies (12 morpho-logical, 7 cytb, 8 COI, 2 COII), with relatively strong branchsupport (BS = 15). This clade is present in the COI tree, aswell as in the M-COI, M-COII, cytb-COI, and all 3- and4-partition combined trees. The Afro-Holarctic canids arecharacterized by a few osteological characters and by elon-gated limbs (Appendix 2A); however, there are no evidentdevelomental and behavioural synapomorphies, whichindicates a conspicuous diversity of life strategies within thegroup.

The internal cladistic structure of the clade is highly unsta-ble, and four subclades are present in most trees. They areC. adustus, C. mesomelas, the other Canis species (‘higher Canis’,henceforth HC), and the Lycaon + Cuon clade (see below). Thegroup of the hypercarnivorous species (Lycaon + Cuon) — ifmonophyletic — is either basal (close to C. adustus andC. mesomelas), or highly derived (close to C. lupus) in various datacombinations. The resolution present in the 4-partition com-bined tree — C. adustus as the basalmost species, C. mesomelasas second basal branch, Lycaon + Cuon as a sister group of HC— is weakly supported in terms of BS scores and numbers ofsynapomorphies. Despite the basal position of Lycaon in somemolecular trees, the side-striped jackal (C. adustus) is mostprobably the basal member of the clade, retaining manyplesiomorphic character states shared with other canids.

The taxonomy of the HC species (C. aureus, C. lupus, C.simensis, C. latrans) needs reevaluation. Genetic data suggestthat red wolves (Canis rufus) resulted from a hybridizationbetween coyotes and grey wolves that probably occurredduring the past 2500 years (Roy et al. 1994, 1996; Reich et al.1999); the eastern Canadian timber wolf (C. lupus lycaon) hasalso been suspected of hybridization with coyotes. In contrast,morphological analysis of the fossil and Recent North Ameri-can Canis spp. suggests that the red wolf did not originate asa hybrid of the (western) grey wolves and coyotes, and thatC. lupus lycaon may have resulted from natural hybridizationof C. rufus and western C. lupus (Nowak 2002). Based on DNAprofiles at eight microsatellite loci and the control region ofthe mitochondrial DNA, Wilson et al. (2000) suggested thatboth the red wolf and the eastern Canadian timber wolfevolved in North America, sharing a common ancestor with

the coyote (about 150 000−300 000 years ago), and proposedclassifying them as a species, Canis lycaon. On the other hand,α1-antitrypsin polymorphism shows that the eastern Cana-dian wolf is either a grey wolf of Eurasian origin, or possiblya hybrid between the latter and the C. lycaon/rufus complex(Mech & Federoff 2002). The phylogenetic structure of theHC clade is highly ambiguous and probably influenced byhybridization (or introgression) events.

African wild dog and dholeBoth these social and hypercarnivorous species form a stronglysupported clade (BS = 15) among the Afro-Holarctic dogs inthe combined tree, supported by 41 synapomorphies (28morphological, 4 cytb, 3 COI, 6 COII). This clade is presentin the 4-partition and morphological trees, as well as in theM-COI, M-COII, M-cytb-COI, M-cytb-COII, and M-COI-COII combined trees, but not in the molecular trees.The tree topology may be compromised by the occurrence oftwo or more ‘problematic species’ that include numerousaberrant character states. Indeed, in the absence of Cuon,Lycaon is a sister group of all Afro-Holarctic dogs (includingC. adustus) even in the 4-partition tree, while the absence ofLycaon does not influence the position of Cuon within para-phyletic Canis. It is possible that the African wild dog is abasal Afro-Holarctic canid resembling the dhole due to itsshared aberrant morphology and behaviour, while the latteris closely related to the HC species. The monophyly of theLycaon + Cuon clade is hence corroborated almost exclusivelyby the morphological evidence, which gives rise to concernthat their similarity may be convergent. The morphologicalsynapomorphies of the Lycaon + Cuon clade include morpho-logy of rostrum/palate part of the skull, zigzag Hunter-Schreger bands of enamel and similar hypercarnivorous teethadaptations, high ratio of neocortex volume, a few develop-mental characteristics, and specialized hunting of large prey.Many of these characters are shared with Speothos as well,some others also with the grey wolf (see Appendix 2A,B).

Phylogeny of the Recent CanidaePreviously published phylogenetic analyses were summa-rized by Bininda-Emonds et al. (1999) using the MRP super-tree method. There is considerable discussion concerningthe utility of this method (see Bininda-Emonds et al. 2002,who support it and e.g. Gatesy et al. 2002, who strongly rejectit). Even if the supertrees do not present an accurate phylo-genetic reconstruction, they can nonetheless present a usefulsummary of the previous trees. Only five supertree clades havea Bremer support of 3 (DC, foxlike canids) or more (Vulpesincl. Alopex and Fennecus 6, Urocyon 7, Canis s.l. 10). Amongthe three nodes with the highest support in the canid super-tree, two are uncontroversial (monophyly of Urocyon, Vulpes)while the third highly supported clade (monophyly of Canis



s.l.) is contradicted by the present analysis. Our preferred treehence differs from the supertree due to (1) paraphyly of thefoxlike canids, (2) better resolution of the DC, and (3) para-phyly of both Canis s.l. and jackals.

The only previously published morphological-molecularanalysis (Wayne et al. 1997) combined the cytb, COI, andCOII sequences of 23 canid species (only three foxes, Vulpesvulpes, V. macrotis, and F. zerda, were included) with 57osteological characters from Tedford et al. (1995). Becausethe genus-level data matrix by Tedford et al. (1995) doesnot agree with the probable paraphyly of Canis s.l. indicatedby the molecular data, only grey wolf sequences werecombined with the ‘Canis’ morphology. The other Canis spe-cies were excluded from the analysis. Similarly, V. vulpes wasthe only Vulpes species included in the combined analysis.The morphological characters published by Tedford et al.(1995), when analysed separately, provided a tree with twobasal clades, one foxlike (Urocyon, Otocyon, Vulpes), the otherdoglike. The DC included the Afro-Holarctic and SouthAmerican groups (the latter with sister clades of Cerdocyon +Nyctereutes and Atelocynus + Speothos, more basal Chrysocyonand basalmost paraphyletic Pseudalopex + Dusicyon + Lycalopexassemblage). Although no sensitivity analysis was performedby Wayne et al. (1997), it is evident that the combination ofmolecular characters and osteological dataset producedthe following: (1) withdrawal of Nyctereutes from the SouthAmerican DC clade, (2) origin of the Speothos + Chrysocyon cladeat the base of the South American canids, and (3) breakup ofthe Lycaon + Cuon branch. The resulting combined tree wasunrooted (neither molecular nor morphological outgroupswere included) and, in accord with the phylogeny presentedhere. There was a clade of South American canids ((Speothos+ Chrysocyon) (Pseudalopex (Atelocynus + Cerdocyon))) as a sistergroup of the Afro-Holarctic clade (Lycaon (Cuon + Canis lupus));Nyctereutes was a sister group of the DC, while Urocyon, Vulpesand Otocyon were more basal outgroup(s).

Muñoz-Durán (2002) reported a canid phylogeny based ona combined analysis of morphological and mtDNA data, buthas yet to provide details enabling direct comparison withother results. The tree presents Urocyon, Nyctereutes andOtocyon + Vulpes as three successive outgroups of the DC.The DC are split into two groups, the ‘zorros’ and the cladeincluding Chrysocyon, Speothos and all the Afro-Holarcticspecies. Within the latter, Cuon belongs to Canis ( just aboveC. adustus, while C. mesomelas is a sister species of HC),Speothos and Lycaon form a clade, and Chrysocyon is a sister speciesof the whole group. The position of Nyctereutes and Otocyonis disputable in our analysis and the placement preferred byMuñoz-Durán (2002) cannot be excluded; the same appliesto the Afro-Holarctic affinities of Chrysocyon and Speothos, andto possible polyphyly of the Lycaon + Cuon cluster. On theother hand, the Speothos + Lycaon clade is most probably based

on convergent osteological and sociobiological charactersjoined with the hypercarnivory of both species.

ClassificationWe propose that the current supraspecific classification ofRecent canids be amended as follows (Fig. 10):1 Alopex Kaup, 1829 and Fennecus Desmarest, 1804 should beincluded in Vulpes Frisch, 1775, as proposed, e.g. by Clutton-Brock et al. (1976: Fennecus), Geffen et al. (1992: both), andBininda-Emonds et al. (1999: Fennecus). Both V. lagopus (Linnaeus,1758) and V. zerda (Zimmermann, 1780) represent derivedspecies of true foxes and their separation to a full-genus levelwould make Vulpes s.s. obviously paraphyletic.2 In the South American subclade, Pseudalopex spp. andLycalopex vetulus should probably be classified under a singlegeneric name (as suggested by Berta 1987; Zunino et al. 1995;Bininda-Emonds et al. 1999), because Pseudalopex s.s. is certainly

Fig. 10 Preferred phylogeny of the Canidae, with a new, revisedgeneric classification. Elevation of ‘Canis’ adustus and ‘C.’ mesomelasto full-level genera, and placement within Schaeffia and Lupullella,respectively, are proposed in the text; all Lycalopex species mayeventually be classified under Dusicyon (see text).



paraphyletic. As the generic name Lycalopex Burmeister, 1854is older than Pseudalopex Burmeister, 1856, all the speciesshould be classified as Lycalopex spp. (see Zunino et al. 1995).The situation is made more complex because the extinct wolf-sized Dusicyon australis appears to represent another memberof the ‘zorro clade’, and is possibly closely related to its largestspecies, L. culpaeus. If this hypothesis is confirmed by furtherresearch, all species of Lycalopex should belong to DusicyonHamilton-Smith, 1839 (see Clutton-Brock et al. 1976). However,the relationships of D. australis await clarification.3 Canis Linnaeus, 1758 is paraphyletic in the traditionalbroad sense. Either Lycaon and Cuon should return to Caniss.l., or ‘Canis’ adustus should be classified under a new genericname (Schaeffia Hilzheimer, 1906). Because of the highconservation priority of both African wild dogs and dholes weprefer to preserve their separate generic names and to elevatethe side-striped jackal to a full genus. The phylogenetic positionof ‘Canis’ mesomelas is more equivocal, and its placementwithin a separate genus (Lupulella Hilzheimer, 1906) is apossible solution.

Combination of different data partitionsThe analysis clearly shows that many canid clades are neitherwell-supported nor stable. Evaluation of the relative qualityof data partitions and their performance in the combinedanalysis produced a rather surprising result: while the mor-phological dataset ranked very poorly in both separate andsimultaneous analyses, morphological synapomorphies rep-resented nearly one quarter of those in the combined tree andone-third of the nonhomoplastic synapomorphies in that tree.This indicates that there is a strong secondary signal hidden in themorphological data matrix, which is influential in constructingthe combined tree. From 35 nonhomoplastic synapomorphiesof the morphological tree (Appendix 2C), 18 (51%) representskeleton, 9 (26%) soft anatomy, body proportion, pelage andcolour patterns, 1 (3%) reproduction and development, 2 (6%)ecology and behaviour, and 5 (14%) cytogenetics. Themorphological synapomorphies that have survived combina-tion of the morphological dataset with three molecular sequences(Appendix 2A,C) exhibit the same pattern (8, 4, 0, 1, 2, respect-ively). On the other hand, the 25 hidden synapomorphies(i.e. the nonhomoplastic morphological synapomorphies ofthe combined tree that are not present in the morphologicaltree; see Gatesy et al. 1999) exhibit a different pattern:purely morphological characters are less common (in total48%: 8 osteological, 4 soft-anatomical) while the ‘biological’characters are relatively more abundant (52%: 3 reproduc-tion and development, 6 ecology and behaviour, 4 cytogenet-ics). Consequently, the latter, which are often overlooked,appear to contribute substantially to the outcome of simul-taneous analysis of the morphological and molecular datapartitions.

While the morphological and molecular approaches arenot congruent in separate analyses, combining them suggeststhat conflicts do not exist throughout the tree. Differencesin the placement of a few taxa (e.g. Nyctereutes, Chrysocyon+ Speothos) are sufficient to create significant incongruence,although excluding morphology would have prevented thisdataset from having an influence over all the remainingrelationships which do not conflict significantly with thesequence data. The hidden support among datasets cannotemerge in separate analyses of individual data partitions. Bycombining all of the datasets, however, we have identifiednodes that may potentially change with the addition ofnew data, i.e. those where different data partitions conflictstrongly with each other (basalmost canid phylogeny, posi-tion of Nyctereutes, position of Speothos and Chrysocyon),and those that are unlikely to change with future newdata, i.e. where all data partitions are in agreement(monophyly of Vulpes, DC, ‘zorros’, Afro-Holarctic canids,and HC clades). The combined analysis suggests that thereis a lot of extra information in morphology that is notcaptured in the restricted set of genes currently beingsequenced.

Note added in proofLyras & van der Geer (2003) describe the external cerebrumanatomy of most Recent Canidae. Two distinctive features ofpossible phylogenetic value are recognized, viz., the sulcalpattern medial to the coronal sulci and the shape and relativesize of the proreal gyrus. The authors hypothesize thatNyctereutes is close to Cerdocyon, Speothos to Atelocynus, andChrysocyon to Dusicyon and Pseudalopex. The combinedmorphological-molecular trees published in the presentstudy are not changed if these two characters are includedto the data matrix.

Lyras, G. A. & van der Geer, A. A. E. (2003). External brainanatomy in relation to the phylogeny of Caninae (Carnivora:Canidae). Zoological Journal of the Linnean Society, 138, 505–522.

AcknowledgementsEli Geffen (Tel Aviv) kindly made available unpublished cyto-chrome b sequences of a few fox species. Oldrich Nedv´dand Magda Vítková (both Ceské Bud´jovice) provided tech-nical assistance. Eli Geffen, Olaf Bininda-Emonds (Leiden),Patricia Moehlman (Arusha), Hynek Burda (Essen), andthree anonymous referees discussed and/or corrected earlierversions of the manuscript. Late Jumper the degu and Rikithe cocker spaniel were a source of support and inspiration.The study was funded by the Czech Ministry of Education(123100003) and Academy of Sciences of the Czech Republic(K6005114).



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Appendix 1: List of morphological charactersMorphological, cytogenetic, developmental and behaviouralcharacters were obtained from the following sources:

Osteological characters (#0–68) predominantly fromClutton-Brock et al. (1976), Berta (1987), Gittleman (1991)and Tedford et al. (1995).

The other morphological characters (body proportion,anatomy, pelage, colour patterns; #69–115) predominantlyfrom Clutton-Brock et al. (1976) and Ortolani & Caro (1996).

Reproduction and developmental characters (#116–125)predominantly from Hayssen et al. (1993), Moehlman &Hofer (1997) and Asa & Valdespino (1998).



Ecological and behavioural characters (#126–173) pre-dominantly from Fox (1969a,b, 1970: Vulpes, Alopex, Urocyon,Canis), Clutton-Brock et al. (1976), Macdonald (1980: Vulpesvulpes), Biben (1982a,b, 1983: Cerdocyon, Chrysocyon, Speothos),Nel & Bester (1983: Otocyon), Estes (1991: African species),Geffen & Macdonald (1992: Vulpes cana), Macdonald (1996:Speothos), Moehlman & Hofer (1997), Hudáková (1998: Speothos),and Sillero-Zubiri & Macdonald (1998: Canis simensis).

Cytogenetic characters (#174–187) largely from Wayneet al. (1987a,b), Park (1996: Nyctereutes), Graphodatsky et al.(2000), and Nash et al. (2001).

Unless otherwise stated, all multistate characters are treatedas ordered (‘additive’). Polymorphic coding is applied; inap-plicable characters are treated as missing information (‘?’). Quan-titative data were coded according to the gap-coding method,modified to minimize character-state overlaps (for discussionsee Swiderski et al. 1998) and to minimize the number ofautapomorphic (unique) character states. Numbers ofchromosomes are referred to the karyotype of Canis lupus(Wayne et al. 1987a,b). The original homology of the cytogeneticcharacters has recently been discussed by Graphodatsky et al.(2000, 2001) and Nash et al. (2001). However, their papersinclude a few model species only and do not allow the overallreorganization of the canid karyological data (note thatincluding/removing the cytogenetic characters does notconsiderably change the combined-tree topology).

The ancestral character states for each character (derivedfrom parsimony optimization of the character in question onthe combined morphological-molecular tree) are indicated inbrackets [anc.]. If the ancestral character state differs fromthe character state in the outgroups, the conflict is indicatedas well [anc. × outg.].

Characters whose rescaled consistency indices (rc = ci*ri )are equal to 1.00 (i.e. those best congruent with the combined-tree topology) appear in boldface (viz., characters #17–19,24, 26, 28, 44–46, 52, 60, 62, 63, 128, 135, 140, 141, 147,157, 160–162, 165, 166, 170, 172–175, 177–179, 181, 186,187).

List of characters0 Temporal ridges size from (0) absent to (3) highlydeveloped (Clutton-Brock et al. 1976) [anc. = 1?].1 Temporal ridges proximity from (0) wide apart to (3) fused(Clutton-Brock et al. 1976) [anc. = 0].2 Interparietal crest from (0) absent to (2) well developed(Clutton-Brock et al. 1976) [anc. = 2?].3 Parietal bones rugosity from (0) smooth to (2) distinctlyrugose (Clutton-Brock et al. 1976) [anc. = 2].4 Postorbital processes convexity (0) concave (1) flat (2)strongly convex (Clutton-Brock et al. 1976) [anc. = 0].5 Mandible, size of subangular lobe from (0) small to (2)large (Clutton-Brock et al. 1976) [anc. = 1].

6 Condylobasal length (0) 90–139 (1) 144–188 (2) 213–226(Clutton-Brock et al. 1976) [anc. = 0].7 Palate, greatest width as percentage of length of palate (0)49–57 (1) 58–67 (2) 70–76 (Clutton-Brock et al. 1976)[anc. = 0?].8 Rostrum, width as percentage of length of palate (0) 24–31 (1) 32–36 (2) 40–47 (Clutton-Brock et al. 1976)[anc. = 0?].9 Rostrum, width as percentage of width of palate (0) 45–49(1) 50–53 (2) 54–57 (3) 59–62 (Clutton-Brock et al. 1976)[anc. = 1?].10 Premaxillae, anterior palatine length as percentage ofwidth of rostrum (0) 61–82 (1) 83–98 (2) 115–124 (Clutton-Brock et al. 1976) [anc. = 1].11 Zygomatic width as percentage of condylobasal length(0) 51–56 (1) 57–59 (2) 61–68 (Clutton-Brock et al. 1976)[anc. = 0].12 Bullae, maximum length as percentage of condylobasallength (0) 12–14 (1) 15–17 (2) 18–20 (3) 26 (Clutton-Brocket al. 1976) [anc. = 1].13 Horizontal ramus (0) deep and thick (1) shallow and thin(Tedford et al. 1995) [anc. = 1 × outg. = 0].14 Paroccipital process width (0) narrow mediolaterally (1)broad, closely appressed to bulla, short free tip turnedlaterally, rarely extends below body of process (Tedford et al.1995) [anc. = 1 × outg. = 0].15 Nasal length (0) long, usually extending posteriorlybeyond maxillary-frontal suture (1) short, rarely extend tolevel of most posterior position of maxillary-frontal suture(Tedford et al. 1995) [anc. = 0].16 Paroccipital process posterior expansion (0) no or littleexpansion (1) expanded posteriorly from bulla, usually withprominent free tip (2) large, greater posterolateral expansion(Tedford et al. 1995) [anc. = 0].17 Frontal sinus (0) absent, presence of a depression ondorsal surface of postorbital process (1) present, lacks a depressionon dorsal surface of postorbital process (2) present, large,penetrates postorbital process and expands posteriorly towardthe frontal-parietal suture [unordered] (Tedford et al. 1995)[anc. = 0].18 Mastoid process (0) small, crestlike (1) enlarged, knob- orridgelike (Tedford et al. 1995) [anc. = 0].19 Zygoma, orbital part (0) presence of a lateral flare andeversion of dorsal border (1) lack of lateral flare and dorsalborder thickened (Tedford et al. 1995) [anc. = 0].20 Scars of medial masseteric muscle (0) narrow and uniformwidth on zygomatic arch and on lateral surface of angularprocess (1) wide on zygomatic arch and enlarged on mandible(Tedford et al. 1995) [anc. = 0].21 Coronoid process (0) short at base relative to dorsoventralheight (1) long at base relative to dorsoventral height (Tedfordet al. 1995) [anc. = 0].



22 Angular process (0) slender, attenuated, with dorsal hook,inferior pterygoid fossa not expanded (1) large, usually bluntwithout dorsal hook, fossa for inferior branch of medialpterygoid muscle expanded (2) deep, short process, fossae forthe pterygoid muscle are expanded (Tedford et al. 1995)[anc. = 0].23 Palate (0) not widened (1) widened (Tedford et al. 1995)[anc. = 0].24 Angular process, superior fossa (0) not expanded (1)expanded with large fossa for superior branch of medialpterygoid muscle (Tedford et al. 1995) [anc. = 0].25 Palatine length (0) extends posteriorly to or just anteriorto end of tooth row (1) extends beyond end of tooth row(Tedford et al. 1995) [anc. = 0].26 Strongly arched zygoma with inverted jugals (0)absent (1) present (Berta 1987) [anc. = 0].27 Mandibular condyle above level of alveolar border ofcheekteeth (0) absent (1) present (Berta 1987) [anc. = 0].28 Supraoccipital shield (0) rectangular or fan-shaped inposterior view, inion may not overhang condyles (1) triangularin shape, inion usually pointed and overhangs condyles (Tedfordet al. 1995) [anc. = 0].29 Average height of olfactory bulb (mm) (0) 1.10–1.46 (1)1.53–1.80 (2) 1.94–2.33 (Gittleman 1991) [anc. = 1].30 Average width of olfactory bulb (mm) (0) 0.83–1.20 (1)1.21–1.56 (2) 1.67–2.12 (Gittleman 1991) [anc. = ?].31 Hunter-Schreger bands in enamel ultrastructure (0)undulating (1) zigzag (Stefen 1999) [anc. = 0].32 P3 posterior cusp from (0) absent to (2) well-developed(Clutton-Brock et al. 1976) [anc. = 1].33 M1 (carnassial) two cusp on heel (0) absent (1) present(Clutton-Brock et al. 1976) [anc. = 1].34 M3 (0) absent (1) present (Clutton-Brock et al. 1976)[anc. = 1].35 DP3 protocone developed as a cusp (0) absent (1) present(Clutton-Brock et al. 1976) [anc. = 1].36 DP4 posterior border concave, so that metacone appearsas a separate lobe (0) absent (1) present (Clutton-Brock et al.1976) [anc. = 0].37 C1 height as percentage of conbylobasal length (0) 8–9(1) 10–11 (2) 12–14 (Clutton-Brock et al. 1976) [anc. = 0?].38 C1 alveolar length as percentage of height (0) 35–40 (1)41–46 (2) 47–53 (Clutton-Brock et al. 1976) [anc. = 2?].39 P4 (carnassial), length as percentage of condylobasallength (0) 5–8 (1) 9 (2) 10–11 (Clutton-Brock et al. 1976)[anc. = 0].40 M2, greatest width as percentage of condylobasal length(0) 4–5 (1) 6 (2) 7–8 (Clutton-Brock et al. 1976) [anc. = 2?].41 M1 hypocone (0) small, barely differentiated from lingualcingulum (1) enlarged (Tedford et al. 1995) [anc. = 1].42 m1 entoconid (0) conical, enlarged, may coalesce withbase of hypoconid to block talonid basin (1) joining with

hypoconid by cristids to form transverse crest (Tedford et al.1995) [anc. = ?].43 m2 metaconid (0) equal to or lower than protoconid (1)enlarged, taller than protoconid (2) greatly reduced or lost[unordered] (Tedford et al. 1995) [anc. = 1 × outg. = 0].44 I1–3 medial cusplets (0) present (1) cusplet in I3 absent(2) cusplet in I1–2 weak or absent (Tedford et al. 1995)[anc. = 1? × outg. = 0].45 Crown height of premolars (0) low-crowned (1) high-crowned (Tedford et al. 1995) [anc. = 0?].46 p2 position (0) not isolated (1) isolated by relativelylarger diastemata than other premolars (Tedford et al. 1995)[anc. = 0?].47 p4 anterior cusplet (0) weak or absent (1) strong (Tedfordet al. 1995) [anc. = 0].48 Canine shape (0) long, slender with recurved crown (1) short,slender, crown not recurved (Tedford et al. 1995) [anc. = ?].49 m2 protostylid (0) absent (1) buccal cingulum bearsprotostylid (Tedford et al. 1995) [anc. = 0].50 m1 protostylid (0) absent (1) present (Tedford et al. 1995)[anc. = 0].51 M1–2 shape (0) transversely wide for their labial length (1)narrow for their labial length (Tedford et al. 1995) [anc. = 0].52 p4 second posterior cusplet (0) absent (1) present(Tedford et al. 1995) [anc. = 0].53 P4 protocone (0) salient, located medial to anteriorborder of paracone (1) reduced (2) further reduced or absent,located posterior to anterior border of paracone (Tedfordet al. 1995) [anc. = 0].54 m1 metaconid and entoconid (0) not reduced (1) greatlyreduced or absent (Tedford et al. 1995) [anc. = 0].55 m2 metaconid and entoconid (0) not reduced (1) greatlyreduced or absent (Tedford et al. 1995) [anc. = 0].56 M1–2 hypocones (0) not reduced (1) greatly reduced orabsent (Tedford et al. 1995) [anc. = 0].57 M1–2 paracones (0) not enlarged (1) enlarged relative tometacone (Tedford et al. 1995) [anc. = 0].58 M1–2 buccal cingulum (0) not reduced (1) reduced or lost(Tedford et al. 1995) [anc. = 0].59 M2 (0) triple-rooted (1) double-rooted, or absent (Tedfordet al. 1995) [anc. = 0].60 I3 (0) small crown extending to or just below level of I1–2,posteromedial cingulum weak or absent (1) large crown extendingmarkedly below level of I1–2, cingulum enlarged, medial crestof I1–2 present merges with cingulum (Tedford et al. 1995)[anc. = 0].61 P3 and p2+3 posterior cusplets (0) present (1) weak orabsent (Tedford et al. 1995) [anc. = 1 × outg. = 0].62 m1+2 with mesoconid (0) absent (1) present (Berta 1987)[anc. = 0].63 m2 with strong paracristid (0) absent (1) present (Berta1987) [anc. = 0].



64 Canines small relative to cheekteeth (0) absent (1) present(Berta 1987) [anc. = 0].65 Carnassials small relative to cheekteeth (0) absent (1)present (Berta 1987) [anc. = 0].66 Scapula, shape of teres major muscle scar on posteriorangle (0) on posterior border only, with plane at right anglesto lateral face (1) intermediate (2) whole scar on lateral face[unordered] (Clutton-Brock et al. 1976) [anc. = 0/1?].67 Scapula, extent of scar of serratus magnus muscle on medialside (0) small (1) large (Clutton-Brock et al. 1976) [anc. = 0].68 Baculum, anterior end bifurcate (0) absent (1) present(Clutton-Brock et al. 1976) [anc. = 0].69 Ears, length as percentage of length of head and body (0)7–13 (1) 14–18 (2) 20–25 (Clutton-Brock et al. 1976)[anc. = 1].70 Tail, length as percentage of length of head and body (0)22–38 (1) 41–59 (2) 60–76 (Clutton-Brock et al. 1976) [anc. = ?].71 Hind feet, length as percentage of length of head andbody (0) 17–19 (1) 20–22 (2) 23–26 (Clutton-Brock et al.1976) [anc. = 2].72 Fore legs, length as percentage of length of body spine(cervical to lumbal vertebrae) (0) 52–62 (1) 63–71 (2) 92(Clutton-Brock et al. 1976) [anc. = ?].73 Hind legs, length as percentage of length of body spine(cervical to lumbal vertebrae) (0) 58–70 (1) 73–83 (2) 103(Clutton-Brock et al. 1976) [anc. = 1].74 Neck, length of cervical vertebrae as percentage ofcombined length of thoracic and lumbar vertebrae (0) 34–38(1) 39–42 (2) 43–47 (Clutton-Brock et al. 1976) [anc. = 1].75 Pelvis, width as percentage of length (0) 53–63 (1) 67–77(Clutton-Brock et al. 1976) [anc. = ?].76 Femur, length as percentage of length of tibia (0) 80–89 (1) 93–97 (2) 100–108 (Clutton-Brock et al. 1976)[anc. = 1].77 Femur, minimum width of shaft as percentage of length(0) 6 (1) 7 (2) 8 (Clutton-Brock et al. 1976) [anc. = 2?].78 Third metatarsal, length as percentage of length of femur(0) 37–42 (1) 43–46 (2) 48–53 (Clutton-Brock et al. 1976)[anc. = ?].79 Baculum, length as percentage of condylobasal length(0) 34–41 (1) 43–49 (2) 50–58 (Clutton-Brock et al. 1976)[anc. = 1].80 Relative length of fore- to hindlimbs (0) short, radius/tibia ratio < 90% (1) long, radius/tibia ratio > 90% (Tedfordet al. 1995) [anc. = 0].81 Forelimb (humerus-radius) (0) longer than 30% of head-body length (1) less than 30% of head-body length (Tedfordet al. 1995) [anc. = 0].82 External auditory meatus very short and of a smalldiameter (0) absent (1) present (Berta 1987) [anc. = 0].83 Caecum shape (0) straight (1) nearly straight (2)convoluted (Berta 1987) [anc. = 2].

84 Number of mammae (0) 4–7 (1) 8 (2) 10–14 (Clutton-Brock et al. 1976) [anc. = 0].85 Ratio of neocortex volume to the of rest of brain (totalbrain volume minus neocortex volume) (0) 1.22–1.31 (1)1.40–1.62 (2) 1.66–1.83 (Dunbar & Bever 1998) [anc. = 0].86 Overall colour, intensity of black pigment (0) absent (1)grey or banded hairs (2) general appearance dark (3) very dark(Clutton-Brock et al. 1976) [anc. = 1].87 Overall colour, intensity of red pigment (0) absent (1) presentas yellow or red underfur (2) general appearance reddish ortan (3) extensive red colour (Clutton-Brock et al. 1976) [anc. = 1].88 Muzzle (0) light (1) dark (Clutton-Brock et al. 1976)[anc. = 0].89 Facial mask between nose and eye (0) absent (1) present(Clutton-Brock et al. 1976) [anc. = 0?].90 Facial mask behind and below eye (0) absent (1) present(Clutton-Brock et al. 1976) [anc. = 0].91 Mystacial vibrissae, length and thickness from (0) low to(2) high (Clutton-Brock et al. 1976) [anc. = 2].92 Neck and back (0) no crest (1) crest (2) mane [unordered](Clutton-Brock et al. 1976) [anc. = 0].93 Back dark longitudinal band (0) absent (1) narrow stripe(2) wide stripe (3) saddle [unordered] (Clutton-Brock et al.1976) [anc. = 0].94 Guard hairs’ coarseness from (0) soft to (2) coarse(Clutton-Brock et al. 1976) [anc. = 1].95 Dorsal guard hairs, length in relation to body size from(0) short to (2) long (Clutton-Brock et al. 1976) [anc. = 1].96 Dorsal guard hairs banded (agouti) (0) present (1) absent(Clutton-Brock et al. 1976) [anc. = 1].97 Underfur, density from (0) low to (2) high (Clutton-Brock et al. 1976) [anc. = 2?].98 Ear pinnae (0) pointed (1) rounded (Clutton-Brock et al.1976) [anc. = 0].99 Ear pinnae (0) light (1) dark (Clutton-Brock et al. 1976)[anc. = 0].100 Ear pinna dark rim (0) absent (1) present (Clutton-Brock et al. 1976) [anc. = 0].101 Tail bushiness from (0) low to (2) high (Clutton-Brocket al. 1976) [anc. = 1].102 Tail dark patch on dorsal surface (0) absent (1) short (2)long [unordered] (Clutton-Brock et al. 1976) [anc. = 0?].103 Tail tip (0) white (1) same as rest of tail (2) black[unordered] (Clutton-Brock et al. 1976) [anc. = 2].104 Fore legs (0) light (1) entirely dark (Clutton-Brock et al.1976) [anc. = 0].105 Fore legs with black line on front (0) absent (1) present(Clutton-Brock et al. 1976) [anc. = 0].106 Hind legs (0) light (1) dark (Clutton-Brock et al. 1976)[anc. = 0].107 Hind feet, dark plantar surface (0) absent (1) present(Clutton-Brock et al. 1976) [anc. = 0].



108 Skin darkly pigmented from (0) light to (2) dark(Clutton-Brock et al. 1976) [anc. = 0].109 Face contrasting (0) absent (1) present (Ortolani & Caro1996) [anc. = 0].110 Face (0) darker (1) unicolour (2) lighter (Ortolani &Caro 1996) [anc. = 1].111 Eye contour (0) unicolour (1) light ring (2) dark around[unordered] (Ortolani & Caro 1996) [anc. = ?].112 Throat and neck (0) darker (1) unicolour (2) lighter (3)white [unordered] (Ortolani & Caro 1996) [anc. = ?].113 Below eyes (0) unicolour (1) dark patch (Ortolani &Caro 1996) [anc. = 0?].114 Eye pupil (0) elliptical (1) circular (e.g. Nowak 1999)[anc. = 0].115 Female body mass (0) < 5 kg (1) 6–9 kg (2) 10–12 kg (3)13–16 kg (4) > 20 kg (Moehlman & Hofer 1997) [anc. = 0].116 Sexual dimorphism (M/F) (0) < 1 or equal (1) > 1(Moehlman & Hofer 1997) [anc. = 0].117 Gestation time (0) 50–58 days (1) 60–65 days (2) > 67 days(Hayssen et al. 1993; Moehlman & Hofer 1997) [anc. = 1].118 Neonate mass (0) 25–150 g (1) 160–350 g (2) more than350 g (Hayssen et al. 1993; Moehlman & Hofer 1997) [anc. = 0].119 Age when eyes open (0) 5–8 days (1) 9–12 days (2) morethan 13 days (Moehlman & Hofer 1997) [anc. = 1].120 Age when teeth erupt (0) 7–11 days (1) 12–15 days (2)more than 21 days (Moehlman & Hofer 1997) [anc. = 1].121 Age when eating first solids (0) 17–21 days (1) 24–28 days (2) 32–40 days (Moehlman & Hofer 1997)[anc. = 0].122 Age at weaning (0) 49–63 days (1) 90–120 days (Moehlman& Hofer 1997) [anc. = 1?].123 Age when reaching adult body mass (0) 90–190 days (1)more than 200 days (Moehlman & Hofer 1997) [anc. = 0].124 Age at sexual maturity (0) 240–270 days (1) 285–330 days(2) 360–365 days (Hayssen et al. 1993; Moehlman & Hofer1997) [anc. = 1?].125 Life span (0) 40–80 months (1) 100–155 months (2)more than 165 months (Hayssen et al. 1993; Moehlman & Hofer1997) [anc. = ?].126 Communal denning and nursing (0) absent (1) present(Moehlman & Hofer 1997) [anc. = 1].127 Polygyny (0) absent (monogamy) (1) present(Moehlman & Hofer 1997) [anc. = 1].128 Multiple pairs breeding (0) absent (1) present(Moehlman & Hofer 1997) [anc. = 0].129 Polyandry (0) absent (monogamy) (1) present (Moehlman& Hofer 1997) [anc. = 0].130 Helpers and/or nonreproductive adults (0) absent (1)females (2) both males and females [unordered] (Moehlman& Hofer 1997) [anc. = ?].131 Territorial behaviour (0) absent (1) present (Moehlman& Hofer 1997) [anc. = 1?].

132 Multiple litters (0) absent (1) present (Moehlman &Hofer 1997) [anc. = 1].133 Close contact (resting in heaps) (0) absent (1) present(see, e.g. Estes 1991; Macdonald 1996) [anc. = 0?].134 Adult regurgitation of food for young (0) absent (1)present (see, e.g. Estes 1991) [anc. = 1].135 Raising lip to show fangs as a threat behaviour (0)absent (merely opening their mouths slightly) (1) present(Fox 1970; see Clutton-Brock et al. 1976) [anc. = 0].136 Low level of food caching (0) absent (1) present (see, e.g.Macdonald 1996) [anc. = 0?].137 Handstand urination (0) absent (1) present (see Estes1991) [anc. = 0].138 Scratching ground before or after urination (0) absent(1) present (see Estes 1991) [anc. = 0].139 Laying during submissive display (belly crawl) (0)absent (1) present (Estes 1991; Hudáková 1998) [anc. = ?].140 Licking during greeting ceremony (0) absent (1)present (see Fox 1970) [anc. = 0].141 Tucking tail between hindlegs as a subordinationposture (0) absent (1) present (see, e.g. Estes 1991;Hudáková 1998) [anc. = 0].142 Frequency of oestrus phases (0) once a year (1) aseasonal(Clutton-Brock et al. 1976) [anc. = 0].143 Size of prey relative to body size from (0) small to (2)large (Clutton-Brock et al. 1976) [anc. = 0].144 Social grooming (0) rare and only between pairs (1) welldeveloped (Clutton-Brock et al. 1976) [anc. = ?].145 Howling (0) absent or only as long-distance contact call(1) present but no physical contact (2) close contact call, socialhowling in unison [unordered] (Clutton-Brock et al. 1976)[anc. = 0].146 Defecation at specific sites (0) absent (1) present(Clutton-Brock et al. 1976) [anc. = 0?].147 Tail posture in dominant animals (0) no distinctposture (1) straight and horizontal (2) raised in a J-shape(3) inverted U-shape [unordered] (Clutton-Brock et al. 1976)[anc. = 3].148 Frequency of tail-wagging in submissive posture from(0) low to (2) high (Clutton-Brock et al. 1976) [anc. = ?].149 Oestrus duration from (0) short to (2) long (Estes 1991;Asa & Valdespino 1998; Nowak 1999) [anc. = 0].150 Duration of copulatory tie from (0) short to (2) long (seeEstes 1991; Alderton 1998; Nowak 1999) [anc. = 0].151 Average number of adults per group during breedingseason from (0) low to (2) high (Moehlman & Hofer 1997;Muñoz-Durán (2002) [anc. = ?].152 Treatment of prey (0) aggressive defense of all food (1)sharing of some foods (2) sharing of most or all foods (Biben1982b) [anc. = 0].153 Treatment of play object (0) aggressive competition(playmates discouraged) (1) mock competition (playmates



encouraged) (2) sharing without competition (Biben 1982a;orig. observ. Lycaon 2) [anc. = 0].154 Cooperative hunting (0) absent (1) noncooperative, butforaging in proximity (2) cooperation [unordered] (see, e.g.Estes 1991; Sillero-Zubiri & Gottelli 1995; Macdonald &Courtenay 1996; Nowak 1999) [anc. = 0].155 Diet (0) opportunist (1) specialized to small prey (2)specialized to large prey [unordered] (see, e.g. Estes 1991;Malcolm 1986; Kauhala 1996; Nowak 1999; Juarez &Marinho 2002) [anc. = 0].156 Ontogeny of aggressivity over food (0) aggressive (1)youngsters more aggressive than adults (2) nonaggressive(Biben 1982a, 1983) [anc. = 0].157 Age of first killing and eating of the prey (0) less than35 days (1) more than 50 days (Fox 1969a; Biben 1983)[anc. = 0].158 Age of first social interactions (pawing, licking,monting) (0) less than 15 days (1) more than 20 days (Biben1983; Fox 1970) [anc. = ?].159 Age of first roll over as a submissive expression (0) lessthan 23 days (1) 26–35 days (Fox 1969b) [anc. = 1].160 Jaw wrestling as a part of ritualized fighting (0)absent or rare (1) common (see Fox 1969b, 1970)[anc. = 0].161 Age of first jaw wrestling (0) 14 days (1) 22–23 days(2) 35–45 days (see Fox 1969b, 1970; Biben 1983)[anc. = ?].162 Period of fighting begins at age (0) 24 days (1) 30–32 days (2) 40–45 days (Fox 1969b; Biben 1983) [anc. = ?].163 Play bow (a posture with lowered forequarters andelevated hindquaters) (0) absent or rare (1) common (Fox1970; orig. observ. Lycaon 1, Chrysocyon 1) [anc. = 1].164 Adults initiate interactions by assuming a submissiveposture (0) absent (1) present (see, e.g. Estes 1991) [anc. = 0].165 Male urinary behaviour is related to dominance (0)absent (1) present (in social species the top male marks mostoften; see Estes 1991) [anc. = ?].166 Orientation of attack during agonistic interactions(0) towards the scruff of the neck (1) towards cheeks, muzzleor lower jaw (Fox 1969b) [anc. = 1].167 Scruff bite intention or scruff-biting associated withmating (0) absent (1) present (Fox 1969b) [anc. = ?].168 Dead shake movements (0) absent (1) present (Fox1969a) [anc. = ?].169 Play with prey terminated when a conspecific attemptsto take the prey from its owner (0) absent (1) present (Fox1969b, 1970; orig. observ. Lycaon 0) [anc. = 1].170 Playing together at the same time with one object(‘tugs-of-war’) (0) rare (1) common (see, e.g. Estes 1991;Macdonald 1996) [anc. = 0].171 Gape (facial expression with opened mouth) (0) absent(1) less developed (2) well developed (Fox 1970) [anc. = 2].

172 ‘Cut off’ response during higher level of intraspecificconflict (facial expression used to stop aggressive behaviour)(0) absent (1) present (Fox 1970) [anc. = 0].173 Head turning and avoidance of eye contact as asubmissive response (0) absent (1) present (Fox 1970)[anc. = 1].174 Chromosome diploid number (0) less than 74 (1) 74(2) 76 (3) 78 [unordered] (Wayne et al. 1987a,b) [anc. = 0].175 Acrocentric chromosomes (0) present (1) absent(Wayne et al. 1987a,b) [anc. = 0].176 Metacentric chromosomes (0) present (1) rare (2) absent[unordered] (Wayne et al. 1987a, b) [anc. = 1 × outg. = 0].177 Chromosome 40 (0) absent (1) present (Wayne et al.1987a,b) [anc. = 1].178 Chromosome 34 (0) absent (1) present (Wayne et al.1987a,b) [anc. = 0?].179 Chromosome 36 (0) absent (1) present (Wayne et al.1987a,b) [anc. = 0?].180 Chromosome 28 (0) present (1) absent (Wayne et al.1987a,b) [anc. = 0].181 Chromosome 22 (0) present (1) absent (Wayne et al.1987a,b) [anc. = 0].182 Terminal segments added to chromosomes 12, 18, 24,and 30 (0) absent (1) present (Wayne et al. 1987a,b)[anc. = 0].183 Chromosome 31 (0) present (1) absent (Wayne et al.1987a,b) [anc. = 0].184 Chromosome 37 (0) present (1) absent (Wayne et al.1987a,b) [anc. = 1? × outg. = 0].185 Chromosome 38 (0) absent (1) present (Wayne et al.1987a,b) [anc. = 1?].186 Reciprocal arm translocation (0) absent (1) present(Wayne et al. 1987a,b) [anc. = 0].187 Heterochromatic short arms (0) absent (1) added tochromosome 4 (Wayne et al. 1987a,b) [anc. = 0].

Appendix 2: Lists of synapomorphies and apomorphies#character:#state (characters listed in Appendix 1). Non-homoplastic synapomorphies are asterisked. Character statesthat represent nonhomoplastic synapomorphies in the mor-phological tree are italicized. DC = doglike canids; HC =‘higher Canis’

(A) Morphological synapomorphies of canid clades in the 4-partition combined tree (23 species)Otocyon + Vulpes + Nyctereutes + DC (no morphologicalsynapomorphy)Vulpes + Nyctereutes + DC (0:1, 37:1*, 38:1, 39:1, 84:1*,122:0*, 178:1*, 179:1*)Vulpes (15:1, 38:0, 117:0, 175:1*)Vulpes vulpes + V. velox (39:2, 89:1, 101:2, 176:0, 185:0)



Nyctereutes + DC (1:2, 4:1, 14:0, 16:1*, 17:1*, 18:1*, 19:1*,26:1*, 69:0*, 71:1*, 94:2*, 114:1*, 127:0*, 166:0*)DC (85:1, 118:1*, 147:1*, 176:2*)South American DC (97:1, 121:1, 122:1, 156:1, 157:1*, 177:0*)Speothos + Chrysocyon (70:0, 83:0, 94:1, 96:0, 104:1, 106:1,121:2*, 162:2*, 181:1*)‘zorros’ (= Atelocynus + Cerdocyon + Pseudalopex + Lycalopex)(1:1, 66:1, 102:2, 117:0)Pseudalopex + Lycalopex (51:0, 62:1*, 63:1*, 83:2, 91:2, 94:1,102:1, 114:0)Afro-Holarctic DC (0:2, 1:3, 6:1, 17:2*, 24:1*, 28:1*, 52:1*,57:1, 60:1*, 67:1*, 80:1)Canis mesomelas + Lycaon + Cuon + HC (8:1, 11:1, 75:1, 143:1)Lycaon + Cuon + HC (4:2, 29:2, 32:2, 38:2, 87:2, 115:3*, 128:1*)Lycaon + Cuon (7:2*, 8:2, 9:2, 11:2*, 23:1, 31:1, 33:0, 35:0,40:0*, 41:0, 47:1, 48:1, 55:1, 56:1, 58:1, 61:0, 71:2, 74:0, 78:0,84:2, 85:2, 95:0, 96:0, 119:2, 120:2, 143:2, 155:2, 164:1)HC (66:1, 105:1, 135:1*, 138:1)Canis lupus + C. latrans + C. simensis (12:0, 121:1, 150:1, 156:1)Canis latrans + C. simensis (8:0, 9:0, 11:0, 37:2, 38:1)

(B) Morphological autapomorphies of canid species in the 4-partition combined tree (23 species)Urocyon cinereoargenteus (49:1, 50:1, 51:1)Otocyon megalotis (5:2*, 8:1, 9:3, 10:0, 29:0, 32:0, 47:1, 66:1,68:1, 69:2, 79:2, 88:1, 90:1, 95:2, 99:1, 101:2, 104:1, 106:1,109:1, 119:0, 154:1, 155:1, 182:1, 183:1)Fennecus zerda (0:0, 2:0*, 3:0, 12:3*, 29:0, 66:2, 69:2, 76:0*,78:2, 94:0*, 110:2, 124:0, 142:1, 182:1, 183:1)Vulpes vulpes (1:2, 9:2, 37:2, 74:0, 85:1, 95:2, 99:1, 103:0,105:1, 111:0)Vulpes velox (70:0, 151:0, 184:0)Nyctereutes procyonoides (4:2, 8:1, 9:3, 11:1, 27:1, 49:1, 65:1,66:2, 70:0, 79:2, 87:2, 90:1, 95:2, 98:1, 100:1, 103:1, 104:1,106:1, 109:1, 146:1, 148:0, 151:0, 184:0)Speothos venaticus (3:2, 4:2, 8:2, 9:3, 10:0, 11:1, 13:0, 15:1,16:2, 23:1, 31:1, 32:2, 33:0, 34:0, 38:2, 41:0, 43:2, 54:1, 55:1,56:1, 57:1, 58:1, 59:1, 71:0, 74:0, 75:1, 76:2, 77:2, 78:0, 81:1,91:0, 95:0, 97:0, 98:1, 101:0*, 108:1*, 111:0, 117:2, 133:1,142:1, 143:2, 145:2, 151:2, 154:2, 155:2)Chrysocyon brachyurus (0:3, 1:3, 6:2*, 7:0, 12:0, 29:2, 30:2,32:0, 39:0, 66:2, 72:2*, 73:2*, 74:2, 77:0, 78:2, 80:1, 87:3,88:1, 92:2*, 93:2, 95:2, 101:2, 115:4*, 116:0, 130:0, 138:1,146:1, 147:2*, 151:0, 174:2*)Atelocynus microtis (0:2, 6:1, 8:1, 9:2, 11:1, 15:1, 81:1, 83:0,95:0, 104:1, 106:1, 111:0, 112:1*, 116:0, 148:0)

Cerdocyon thous (0:0, 2:1, 27:1, 36:1, 37:0, 48:1, 65:1, 78:2,92:1, 93:1, 107:1*, 118:0, 142:1, 154:1, 176:0)Pseudalopex culpaeus (43:0, 50:1, 53:1, 79:0, 110:2, 115:1,143:1)Pseudalopex griseus (0:0, 2:1, 3:0, 9:0, 69:1, 70:2, 71:2, 77:0,91:1, 124:2)Pseudalopex gymnocercus (116:0, 143:1)Pseudalopex sechurae (4:2, 8:1, 9:2, 11:1, 29:0, 36:1, 87:0)Lycalopex vetulus (4:2, 8:1, 9:2, 11:1, 12:2, 32:0, 39:0, 51:1,155:1)Canis adustus (12:0, 38:0, 84:0, 105:1, 114:0)Canis mesomelas (69:1, 76:2, 77:0, 79:0, 93:3*, 101:2, 110:2,116:0, 120:0, 124:2, 127:1)Lycaon pictus (3:0, 9:3, 16:2, 30:2, 37:2, 53:1*, 68:1, 86:2, 88:1,94:1, 98:1, 100:1, 103:0, 108:2*, 110:0*, 111:2, 112:0*, 116:0,117:2, 125:1, 131:0)Cuon alpinus (0:1, 34:0, 36:0, 43:2, 54:1, 59:1, 66:2, 67:0, 72:0,75:0, 76:2, 77:2, 86:0, 87:3, 124:2, 150:1)Canis aureus (71:0, 79:0, 110:2, 111:0, 127:1, 146:1)Canis lupus (0:3, 31:1, 35:0, 78:0, 84:2, 85:2, 93:2, 100:1,120:2, 143:2, 150:2*, 152:2, 164:1)Canis latrans (40:2, 103:1)Canis simensis (0:1, 1:2, 4:1, 7:0, 32:1, 38:0, 39:1, 86:0, 87:3,94:1, 105:0, 129:1, 132:0, 143:0, 145:1, 146:1)

(C) Nonhomoplastic synapomorphies of canid clades in the morphological tree (23 species)Urocyon + Otocyon (44:2*, 45:1*, 46:1*, 178:0*, 179:0*)Vulpes (present only in the majority-rule consensus tree;

15:1*, 38:0*, 117:0*, 175:1*)Nyctereutes + DC: 4:1*, 16:1*, 17:1*, 18:1*, 19:1*, 26:1*,174:1*, 176:2*)Lycalopex + Pseudalopex sechurae + Nyctereutes + Cerdocyon + Atelocynus+ Speothos + Chrysocyon + Afro-Holarctic canids (present onlyin the majority-rule consensus tree; 36:1*, 69:0*, 71:1*)Lycalopex + Pseudalopex sechurae + Nyctereutes + Cerdocyon+ Atelocynus + Speothos (73:0*)Nyctereutes + Cerdocyon + Atelocynus + Speothos (22:2*, 64:1*, 82:1*)Atelocynus + Speothos (81:1*)Chrysocyon + Afro-Holarctic canids (74:2*, 80:1*, 115:[2,3]*)Canis (excl. C. simensis) + Cuon + Lycaon (67:1*)Canis aureus + C. latrans + C. lupus + Cuon + Lycaon(135:1*)Canis latrans + C. lupus + Cuon + Lycaon (149:1*)Canis lupus + Cuon + Lycaon (84:2*)Cuon + Lycaon (7:2*, 11:2*, 40:0*)

phylogeny of recent canidae (mammalia, carnivora): relative reliability and utility of morphological...

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